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J. Paleont., 79(5), 2005, pp. 907–926 Copyright q 2005, The Paleontological Society 0022-3360/05/0079-907$03.00

UPPERMOST MISSISSIPPIAN BRACHIOPODS FROM THE BASAL ITAITUBA FORMATION OF THE AMAZON BASIN, BRAZIL ZHONG-QIANG CHEN,1 JUN-ICHI TAZAWA,2 G. R. SHI,3

AND

NILO SIGUEHIKO MATSUDA4

School of Earth and Geographical Sciences, The University of Western Australia, 35 Stirling Highway, Crawley 6009, Australia, ,[email protected]., 2Department of Geology, Faculty of Science, Niigata University, Niigata 950-2181, Japan, 3 School of Ecology and Environment, Deakin University, Melbourne Campus, 221 Burwood Highway, Burwood, Victoria 3125, Australia, and 4 PetroBras/Un-Eep/St/Msp, 65/1301 Chile Avenue, Rio de Janeiro, Centro CEP-20031-912, Brazil 1

ABSTRACT—This paper describes 19 brachiopod species (including six indeterminate species) in 15 genera and one indeterminate genus from the basal Itaituba Formation at the Caima Quarry 1 section of Itaituba, Amazon Basin, Brazil. The faunal correlations of the brachiopods and the associated fusulinids and conodonts indicate a late Chesterian (late Serpukhovian) age for the described fauna, therefore confirming for the first time the presence of uppermost Mississippian rocks in the Amazon Basin. A new species, Composita caimaensis, is created, and two species, Inflatia cf. gracilis and Marginovatia cf. catinulus, are described for the first time from the Amazon Basin. The Amazon brachiopods appear to be of strong affinity with coeval faunas of the North American midcontinent.

INTRODUCTION

prospecting in the Amazon Basin, northern Brazil, reveals that Carboniferous sedimentary successions are extensively present in outcrop and also in the subsurface. In general, these rocks have been referred to the Itaituba Formation, which attains more than 1,000 m in thickness in some boreholes (Mendes, 1959; Rocha Campos, 1985). Brachiopods from the Itaituba Formation are extremely diverse and abundant and have been dealt with in several previous studies (i.e., Derby, 1874; Katzer, 1903; Reed, 1933; Mendes, 1952, 1956a, 1956b, 1957, 1959, 1961; Dresser, 1954). However, a zonal scheme has not been developed for the Itaituba brachiopod fauna, therefore preventing a satisfactory comparison with coeval faunas outside the Amazon Basin on a zonal basis. It is probably for the same reason that the age of the Itaituba Formation remains controversial despite the presence of abundant and varied macro- and microfaunas (including smaller foraminifers, fusulinaceans, and conodonts) and numerous previous biostratigraphic investigations (e.g., Petri, 1952, 1956; Fulfaro, 1965; Daemon and Contreiras, 1971; Tengan et al., 1976; Caputo and Crowell, 1985; Lemos and Medeiros, 1989, 1996; Altiner and Savini, 1995). Currently, several opinions exist on the age of the Itaituba faunas (Table 1). The possible Mississippian affinity of the lowest fusulinid assemblage has been commented upon by Petri (1952) who, nevertheless, correlated the entire Itaituba Formation with Millerella and Fusulinella zones with the Desmoinesian of the Middle Pennsylvanian. This was largely followed by Mendes (1956a, 1956b, 1957, 1959, 1961), who studied the brachiopod faunas of the formation. Conodonts were later recovered from the formation, but they produced apparently three different ages, with Tengan et al. (1976) suggesting a late Mississippian to middle Pennsylvanian age and Lemos and Medeiros (1996) constraining a late Chesterian (late Mississippian) age, in contrast to Lemos and Medeiros (1989), who proposed a Morrowan to Atokan age (Table 1). Adding to the debate, Daemon and Contreiras (1971) and Caputo and Crowell (1985) argued for a much younger Westphalian D to Stephanian age (corresponding to late Moscovian to Gzhelian) on account of palynological data from boreholes. More recently, Altiner and Savini (1995) constrained the Itaituba Formation as Bashkirian (Morrowan to early Atokan) based on fusulinaceans and smaller foraminifers from borehole samples. The uncertainties on the age of the Itaituba Formation have considerably hindered not only the improvement of stratigraphic models for hydrocarbon exploration within the basin, but also our understanding of the biogeographic evolution of northern South

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ECENT GEOLOGICAL

America during the Carboniferous. With this in mind, in this paper we attempt to provide a new constraint on the age of the formation using a new brachiopod collection from the basal Itaituba Formation from its type area. In addition, the biogeographic affinity of the Itaituba brachiopods is also addressed. This brachiopod collection was made by one of us (NSM) from the lower Itaituba Formation exposed at Caima Quarry 1 section in the Caima area, Itaituba, Amazon Basin (Fig. 1), during his field work with a joint Japanese-Brazilian geological survey team in 2000 and 2001. STRATIGRAPHY AND AGE OF BRACHIOPOD FAUNA

The Itaituba Formation rests conformably on the nearshore siliciclastic deposits of the Monte Alegre Formation, and is overlain by the Upper Carboniferous evaporites (salt and anhydrite) of the Nova Olinda Formation (Rocha Campos, 1985). Lithologically, the Itaituba Formation is composed of epeiric argillaceous limestone interbedded with numerous shale and sandstone beds in the lower part, and thin-bedded dolomitic limestone in the upper part (Rocha Campos, 1985). At Caima, the lower part of the Itaituba Formation is exposed at two quarries (Fig. 1). Of these, Quarry 1 is located about 1 km north of Caima village, where a 16.2 m thick succession of the Itaituba Formation is exposed. This succession can be lithologically correlated with that of the basal Itaituba Formation, typically cropping out at the riverbank of the Tapajos River at Itaituba, about 20 km east of Caima. The unit at both localities is marked by a carbonate-dominated sequence and massive yellow quartz sandstone at its base (Rocha Campos, 1985). A diverse brachiopod fauna from the Caima Quarry 1 section is described below. Quarry 2 is about 2.5 km north of Quarry 1 (Fig. 1), but there is a gap in vertical thickness of 6.2 m between the sections of the two quarries in view of their respective correlation to the succession of the same formation from a borehole made between these two quarries (Fig. 2). In this correlation, the base of the section at Caima Quarry 2 is about 6.2 m stratigraphically above the top of the Caima Quarry 1 section (Fig. 2). At Caima Quarry 2, the sequence is about 16.6 m thick and comprises argillaceous limestone. The brachiopods from this quarry are characterized by species of Buxtonioides Mendes, 1959, Linoproductus Chao, 1927b, Anthracospirifer Lane, 1963, Neospirifer Fredericks, 1923, and Cleiothyridina Buckman, 1906, coexisting with late Morrowan conodonts: Adetognathodus lautus (Gunnel, 1933), Idiognathodus sinuosis Ellison and Graves, 1941, I. incurvus Dunn, 1966, Idiognathoides sinuatus Harris and Hollingsworth, 1933, Declinognathodus noduliferus (Ellison and Graves, 1941), and Streptognathodus parvus Dunn, 1966 (Chen et al., 2004).

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JOURNAL OF PALEONTOLOGY, V. 79, NO. 5, 2005

TABLE 1—Historic reviews of the chronostratigraphic division of the Itaituba Formation, Amazon Basin.

FIGURE 1—Simplified geological map of the Amazon Basin, Brazil, showing location of the Caima section.

CHEN ET AL.—UPPERMOST MISSISSIPPIAN BRACHIOPODS OF THE AMAZON BASIN

FIGURE 2—Composite columnar sections of the lower Itaituba Formation at Caima of the Amazon Basin.

The lower part of the Itaituba Formation exposed at the Caima Quarry 1 section comprises gray thin-bedded argillaceous limestone with intercalations of brown and gray shale and siltstone. Brachiopods from the Caima Quarry 1 section consist of 19 taxa: Inflatia cf. gracilis Gordon, Henry, and Treworgy, 1993, Eomarginifera oddonei (Mendes, 1959), Kozlowskia petrii Mendes, 1959, Brasilioproductus chandlessi (Derby, 1874), Marginovatia cf. catinulus Gordon and Henry, 1990, Streptorhynchus hallianus Derby, 1874, Tapajotia tapajotensis (Derby, 1874), Rhipidomella penniana (Derby, 1874), Schizophoria (Schizophoria) sp., Orthotichia morganiana (Derby, 1874), Anthracospirifer oliveirai (Mendes, 1956b), Anthracospirifer sp., Spirifer sp., Mesochorispira dresseri (Mendes, 1956b), Phricodothyris mourai Mendes, 1956b, Spiriferidae gen. and sp. undet., Punctospirifer sp., Composita sp., and C. caimaensis n. sp. Among them, 10 species, Eomarginifera oddonei, Kozlowskia petrii, Brasilioproductus chandlessi, Streptorhynchus hallianus, Tapajotia tapajotensis, Rhipidomella penniana, Anthracospirifer oliveirai, Mesochorispira dresseri, Phricodothyris mourai, and C. caimaensis, are known only from the Amazon Basin. Of the remaining taxa, Inflatia cf. gracilis, Marginovatia

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cf. catinulus, and Anthracospirifer sp. are morphologically comparable with Inflatia gracilis, Marginovatia catinulus, and Anthracospirifer increbescens (Hall, 1858), respectively, and the latter three species are restricted to the upper Chesterian in the North American midcontinent (Weller, 1914; Gordon and Henry, 1990; Gordon et al., 1993); Orthotichia morganiana ranges from the lower Pennsylvanian to Permian (Hall and Clarke, 1892; Chao, 1927a); and the remaining five are identified merely to genus or generically undetermined because of a lack of sufficient specimens and/or poor preservation. Apart from the above three forms comparable with the certain Mississippian species, Mesochorispira Carter, 1992 is also generally limited to the Lower Carboniferous in North America (Carter, 1992), Asia (Chen and Archbold, 2000), and Europe (Sartenaer and Plodowski, 2002). In addition, Anthracospirifer oliveirai may be synonymous with A. leidyi (Norwood and Pratten, 1855) of the Chesterian in Illinois (for more detailed discussions, see below). Langenheim (1991, p. 342) considered that Rhipidomella nevadensis (Meek, 1877) is synonymous to R. penniana; the former is a prominent, widespread zonal species occurring in latest Chesterian sequences at many localities in the southwestern US (Langenheim, 1991; Morrow and Webster, 1992). Additionally, Kozlowskia Fredericks, 1933 is very common in the Pennsylvanian and younger strata. Brasilioproductus Mendes, 1959 has been placed in synonymy with the lower Pennsylvanian Sandia Sutherland and Harlow, 1973 by Langenheim (1991) and Gordon et al. (1993), suggestive of a remarkable morphologic resemblance between the two. However, there are no taxa specific to the Pennsylvanian in age. As a result, the described brachiopod fauna from the basal Itaituba Formation at Caima Quarry 1 appears to range from latest Mississippian and early Pennsylvanian in age, but is herein considered most likely to be of Chesterian age, consistent with the associated fusulinacean and conodont faunas. At Itaituba, the fusulinid Millerella cf. marblensis Thompson, 1942, which also extends to the upper part of the formation, was found in association with the basal Itaituba Formation brachiopod fauna; while Tetrataxis sp. and Fusulinella silvai Petri, 1952 appear only in higher beds of the formation (Rocha Campos, 1985, p. 222). Although Petri (1952) dated all the Itaituba Formation fusulinids as Desmoinesian, he also stated that the lowest Itaituba Formation fusulinids appear to have a Mississippian affinity. In fact, species of Millerella Thompson, 1942 are also common in the Serpukhovian (equal to the Chesterian) fusulinacean faunas in northwest China (e.g., Zhao et al., 1984) and Japan (e.g., Ueno and Nakazawa, 1993). Accordingly, fusulinids from the Itaituba Formation may range from Chesterian to Desmoinesian in age. Conodonts have also been reported from both outcrop sections and subsurface samples of the Itaituba Formation. Fulfaro (1965) described a conodont fauna from some outcrops of the Itaituba Formation exposed along the Tapajos River. This conodont fauna, which was later revised in taxonomy by Tengan et al. (1976), includes some Pennsylvanian species, Cavusgnathus gigantus (Gunnell, 1933), Idiognathodus magnificus Stauffer and Plummer, 1932, and I. delicatus Gunnell, 1931, as well as several species diagnostic of the Mississippian: Gnathodus bilineatus (Roundy, 1926), Ozarkodina delicatula (Stauffer and Plummer, 1932), Rachistognathus muricatus (Dunn, 1965), Geniculatus claviger (Roundy, 1926), and Gnathodus girtyi Hass, 1953 (Dunn, 1965, 1970; Lane and Straka, 1974; Rocha Campos, 1985; Conil et al., 1990; Webster and Groessens, 1990). Lemos and Medeiros (1996) described a Chesterian (late Mississippian) conodont fauna from the lower Itaituba Formation exposed along the Tapajos River in the Caima areas. Tengan et al. (1976) listed a conodont fauna from some subsurface samples of the Itaituba Formation from one

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well in the Upper Amazon area, including Cavusgnathus gigantus, Idiognathodus sinuatus Harris and Hollingsworth, 1933, I. delicatus, Oxinagnathus corrugatus (Harris and Hollingsworth, 1933), Streptognathodus anteeccentricus Dunn, 1966, S. parvus, and Ozarkodina delicatula. Tengan et al. (1976) considered the fauna to be of Morrowan age. In addition, Gnathodus bassleri (Harris and Hollingsworth, 1933), Streptognathodus expansus Igo and Koike, 1964, and Rachistognathus muricatus were identified by Tengan et al. (1976) from an assemblage reported originally by Araujo and Rocha Campos (1969) from a well in the Upper Amazon area (see Rocha Campos, 1985, p. 222). Similarly, Lemos and Medeiros (1989) proposed a Morrowan to Atokan age for the conodonts from the subsurface successions of the Itaituba Formation of the Amazon Basin (Table 1). In view of the evidence from the brachiopods, fusulinaceans, and conodonts from the outcrop successions, the basal part of the Itaituba Formation is most likely latest Chesterian [equating late Serpukhovian in accordance with Groves et al. (1999) and Lane et al. (1999)] in age, although it may span across the Mississippian–Pennsylvanian boundary. As such, the entire Itaituba Formation would range from Chesterian to Desmoinesian in age (Table 1), if the Morrowan to Atokan (Bashkirian–Moscovian) faunas from the upper part of the lower Itaituba Formation (Altiner and Savini, 1995; Chen et al., 2004) are taken into account. In this case, the possible mid–Carboniferous boundary may be positioned within the upper part of the Caima Quarry 1 succession. FAUNAL AFFINITY

As discussed above, the majority of the identified species (about 71%) of the Itaituba brachiopod fauna are endemic to the Amazon Basin. However, at the generic level, other than both Brasilioproductus and Tapajotia Dresser, 1954, all genera (87.5%) are also very common in the uppermost Mississippian (or Lower Carboniferous) and lower Pennsylvanian (or Upper Carboniferous) in the North American midcontinent (e.g., Sturgeon and Hoare, 1968; Sutherland and Harlow, 1973; Gordon, 1975; Langenheim, 1991; Henry and Gordon, 1992). In particular, the presence of the comparable elements of several Mississippian species (such as Inflatia gracilis, Marginovatia catinulus, and Anthracospirifer increbescens), which are characteristic of the Mississippian brachiopod fauna of the western United States (Arrow Canyon, Clarke County, Nevada), in the Amazon Basin is indicative of close linkage between these two faunas, therefore confirming Langenheim’s (1991) observation that there was a continuity of fauna and environment from the Amazon to the southwestern United States. In sharp contrast to the strong affinities with the southern North American faunas, the Amazon fauna, curiously, shares no species with other South American faunas outside the Amazon. This is in part attributable to the fact that Chesterian successions are missing from many areas of South America. SYSTEMATIC PALEONTOLOGY

Horizon, locality, and repository.All described specimens were collected from the basal Itaituba Formation of the Caima Quarry 1 section in the Amazon Basin and are housed in the Department of Geology, Niigata University, Japan, with prefix NUB. Terms and classification.All morphologic terms in current use follow those of Brunton et al. (1996) and Williams and Brunton (1997). The supraordinal classification follows Williams et al. (1996). The classifications of the Productida, Orthotetidina, Orthida, Spiriferida, and Athyridida are after Brunton et al. (2000), Williams and Brunton (2000), Williams and Harper (2000), Carter et al. (1994), and Alvarez and Rong (2002), respectively.

Phylum BRACHIOPODA Dume´ril, 1806 Subphylum RHYNCHONELLIFORMEA Williams, Carlson, Brunton, Holmer, and Popov, 1996 Class STROPHOMENATA Williams, Carlson, Brunton, Holmer, and Popov, 1996 Order PRODUCTIDA Sarycheva and Sokolskaya, 1959 Suborder PRODUCTIDINA Waagen, 1883 Superfamily PRODUCTOIDEA Gray, 1840 Family PRODUCTELLIDAE Schuchert, 1929 Subfamily PLICATIFERINAE Muir-Wood and Cooper, 1960 Tribe YAKOVLEVIINI Waterhouse, 1975 Genus INFLATIA Muir-Wood and Cooper, 1960 Discussion.The diagnostic characteristics of the genus are: outline elongate subquadrate, lateral slopes steep, median sulcus shallow, corpus cavity fairly deep, trails moderately long; costae or costellae regularly spaced; rugae weak, confined to posterior part of ventral disk, and over dorsal disc; a single spine row along hinge and extending across ears; spines scattered on costae of ventral disk; dorsal spines absent. The descriptive characteristics given above incorporate those given by Muir-Wood and Cooper (1960) and Gordon et al. (1993). In 1993, Gordon et al. also compared Inflatia with its allies, notably Antiquatonia Miloradovich, 1945, Keokukia Carter, 1990, Rugoclostus Easton, 1962, Brasilioproductus, and Sandia. In the same paper, they also reviewed the stratigraphic distribution of Inflatia species, and concluded that Inflatia is restricted to the Mississippian in North America. We agree with Gordon et al. (1993) that some Inflatia species previously reported outside the United States require further revision. For instance, McKellar (1965) created Inflatia dekielis to include several specimens named Dictyoclostus cf. simplex Campbell by Maxwell (1964, p. 42, pl. 7, figs. 23–27) from the Westphalian of Queensland, Australia. This Queensland species is much larger than typical Inflatia illustrated by Gordon et al. (1993), and sculptured with more irregular costae and narrower rugae, which form a well-defined reticulation over the disks. In addition, a row of four spines is arranged across the flanks (see McKellar, 1965, pl. 3, figs. 2, 6); two rows of coarse spines are present along the hinge. Several coarse spine bases are scattered on the trail (see McKellar, 1965, pl. 3, fig. 10a). These morphologic features suggest the Queensland species belongs in Antiquatonia rather than Inflatia. INFLATIA

CF. GRACILIS

Gordon, Henry, and Treworgy, 1993 Figure 3.1–3.12

cf. Inflatia gracilis GORDON, HENRY, 8.6–8.11, 8.14, 8.31–8.33.

AND

TREWORGY, 1993, p. 17, fig.

Description.Medium to large Inflatia (measurements see Table 2), with elongate oval to subquadrate outline; widest at hinge line; profile strongly curved; both valves geniculate; ears large, slightly inflated. Ventral beak stout, incurved, overhanging hinge line; umbo moderately broad, strongly curved posteriorly, with steep lateral slopes; median sulcus narrow, shallow, originating anterior to beak, more pronounced at start of protrail (sensu Brunton and Lazarev, 1997), indistinct on trail; trail long, subquadrate in shape, with subparallel lateral margins. Dorsal valve gently concave; corpus cavity moderately deep. External costae round in cross section, originating at beak, bifurcating anteriorly, 17–18 in 10 mm at start of protrail; rugae fine, confined to disk, forming regular reticulation with costae; spines coarse, scatter on costae of disk, ears, and trail; a row of halteroid spines along hinge, about five to seven spines on each side of beak, projecting laterally; those scattered on ventral disk fine; those on trail coarser, over one costa in diameter. Dorsal ornaments and interiors not observed.


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FIGURE 3—1–12, Inflatia cf. gracilis Gordon, Henry, and Treworgy, 1993. 1, 2, NUB-491, a crushed ventral valve in anteroventral and posterior views; 3–5, NUB-492, a ventral valve in later, anteroventral, and posterior views; 6, 8, NUB-493, a ventral valve in anteroventral and posterior views; 7, NUB-494, a specimen in posterior view; 9, 12, NUB-495, NUB-497, two specimens in anteroventral views; 10, 11, NUB-496, a specimen in posterior and anteroventral views. 13–20, Brasilioproductus chandlessi (Derby, 1874). 13, 16–18, NUB-506, a specimen in posterior, lateral, anteroventral, and posteroventral views; 14, 15, 19, NUB-507, a specimen in anteroventral, lateral, and posterior views; 20, NUB-508, a specimen in ventral view. All from the lower Itaituba Formation of the Caima section, Itaituba, Amazon Basin; all 31.5.

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TABLE 2—Measurements (in mm) of Inflatia cf. gracilis. Specimens

Width

Length

C-length

V-height

N. of costae**

NUB-491* NUB-492 NUB-493 NUB-494* NUB-495 NUB-496* NUB-497* NUB-498 NUB-499

28.20 25.51 21.57 23.96 18.25 25.18 23.37 17.72 21.09

28.00 26.31 24.67 23.35 23.24 19.40 19.45 17.85 21.88

42.00 48.04 47.45 47.43 39.85 39.36 37.95 29.81 42.57

13.50 14.60 14.84 14.38 12.50 12.63 11.50 10.54 13.72

15 17 17 16 18 15 15 17 16

* Specimens have been crushed. ** Number of costae in 10 mm at the midlength of the shell; C-length: Curvature length; V-height: Ventral valve height (hereafter).

Material examined.NUB-491, NUB-492, NUB-493, NUB494, NUB-495, NUB-496, NUB-497, NUB-498, NUB-499. Discussion.Except for being considerably larger than the midcontinent materials, the specimens examined fit almost within the concept of Inflatia gracilis (see Gordon et al., 1993) from the uppermost Chesterian of the American midcontinent. Externally, the type species I. inflata (McChesney, 1860, p. 40) is distinct from the examined materials in having a row of spines along the hinge extending onto ears and relatively coarser ribs. Another species, Inflatia cherokeensis, created by Gordon et al. (1993), approaches the Amazon materials in possessing the same total number of ribs on the ventral valve. The described materials differ, however, from the former in their more slender shape, finer and more closely spaced costae and costellae, and smaller ears. Two species of Inflatia have been described from the Viseán of eastern Australia by Roberts (1964, 1976), but neither are morphologically close to the Amazon specimens: I. elegans Roberts (1964, p. 202–204, pl. 2, figs. 1–18) is smaller, has finer but more numerous ribs, and lacks a ventral sulcus; I. engeli Roberts (1976, p. 61–64, pl. 12, figs. 11–28) is more globular, possesses a greater number of costae, and lacks a ventral sulcus. Two Kazakhstan species, I. patria Litvinovich (in Litvinovich et al., 1969, pl. 52, figs. 1–3) from the Viseán and I. uschkarensis Litvinovich (Litvinovich et al., 1969, pl. 53, figs. 1–3) from the Serpukhovian, are also similar in overall appearance to the described materials, from which both central Asian species are distinct in being significantly larger, having a greater numbers of costae and spines, and lacking a ventral median sulcus. Family PRODUCTIDAE Gray, 1840 Subfamily PRODUCTINAE Gray, 1840 Tribe KOZLOWSKIINI Brunton, Lazarev, and Grant, 1995 Genus EOMARGINIFERA Muir-Wood and Cooper, 1960 EOMARGINIFERA ODDONEI (Mendes, 1959) Figure 4.3–4.5 Marginifera oddonei MENDES, 1959, p. 63, pl. 5, figs. 1, 2; ROCHA CAMPOS AND ARCHANGELSKY, 1985, pl. 6, fig. 11.

Description.Medium-sized for genus (about 19.60–25.00 mm wide); subovate in outline; ventral valve evenly convex; beak small, incurved but not turned over hinge line; umbo convex and rounded posteriorly, geniculate anteriorly; lateral slopes steep; median sulcus shallow, indistinct, originating anterior to umbo, increasing in width at start of protrail, becoming prominent on trail. External costae regularly deeply spaced, bifurcating, usually eight to nine in 5 mm at midlength of shell; concentric rugae confined to protrail; five to six coarse spine bases arranged anterior portion of shell. Material examined.NUB-500, NUB-501, NUB-502. Discussion.Although no dorsal material is available for

study, all features observed from the ventral valve agree well with the concept of Mendes’s species. The fine, distinctive costae and a smaller number of coarse spines suggest Mendes’s species is more suitable for Eomarginifera than Marginifera Waagen, 1883. From the type species E. longispinus (s. Brunton, 1966), E. oddonei is differentiated in having a larger size, a shallower ventral median sulcus, and less regularly spaced costae. Genus KOZLOWSKIA Fredericks, 1933 KOZLOWSKIA PETRII Mendes, 1959 Figure 4.8–4.12 Kozlowskia petrii MENDES, 1959, p. 65–69, pl. 5, figs. 3a–c, 4a–c, 5.

Description.Small species for genus (approximately 16.2– 16.9 mm wide, 14.7–15.5 mm long); outline subquadrate to suboval; widest slightly anterior to midlength of shell; both valves geniculate anteriorly; ears small. Ventral disk subquadrate-shaped; beak small, pointed, overhanging hinge line; umbo rounded posteriorly, flattening anteriorly at start of protrail; median sulcus absent; protrail short, nearly perpendicular to disk. Dorsal valve concave, subquadrate in shape; corpus cavity moderately deep; fold absent. Ventral costae weak, confined to disk and protrail; spines coarse, halteroid, about 1.1 mm in diameter, three on protrail, two on each flank; dorsal ornaments and interiors not observed. Material examined.NUB-503, NUB-504, NUB-505. Discussion.The new materials examined are identical to Mendes’s (1959) species in all observed characteristics. The most similar species to Kozlowskia petrii is the type species K. capaci (d’Orbigny, 1842; see also Kozlowski, 1914, pl. 2, figs. 1–15; pl. 5, fig. 13; Muir-Wood and Cooper, 1960, pl. 63, figs. 13–19) from the Lower Permian of Capinota, Bolivia, both having a small size and weakly defined ribs, and lacking a sulcus and fold. However, the type species is distinct from K. petrii in being less consistently rectangular and more oval in outline, and having a less anteriorly flattened ventral umbo and rugae on the disks. K. petrii also approaches two widespread North American species: K. haydenensis (Girty, 1903; see also Dunbar and Condra, 1932; Hoare, 1961; Sutherland and Harlow, 1967, 1973; Sturgeon and Hoare, 1968) and K. splendens (Norwood and Pratten, 1855; see also Girty, 1915; Dunbar and Condra, 1932; Sturgeon and Hoare, 1968; Carter and Poletaev, 1998). The latter two are characteristic of the Pennsylvanian faunas in North America. However, the presence of a ventral median sulcus, pair of larger ears, and more consistent ribs distinguishes K. haydenensis from K. petrii. Unlike K. petrii, K. splendens has a larger size, a relatively more transverse outline, larger ears, and a more strongly ribbed ventral umbo. None of Kozlowskia species described from Spain by Winkler Prins (1968) and Martıńez Chacoń (1982) is particularly close morphologically to the Amazon species. Subfamily BUXTONIINAE Muir-Wood and Cooper, 1960 Tribe TOLMATCHOFFIINI Sarycheva in Sarycheva et al., 1963 Genus BRASILIOPRODUCTUS Mendes, 1959 BRASILIOPRODUCTUS CHANDLESSI (Derby, 1874) Figures 3.13–3.20, 4.6, 4.7 Productus chandlessi DERBY, 1874, p. 51–53, pl. 4, figs. 1–4, 9–11, 13, 16; pl. 6, fig. 1; KATZER, 1903, p. 164, pl. 7, fig. 3a–c. Productus (Dictyoclustus) chandlessi DERBY; REED, 1933, p. 520–521. Productus semireticulatus chandlessi DERBY; DUARTE, 1938, p. 22–23, pl. 1, fig. 4; pl. 4, figs. 6–9. Brasilioproductus chandlessi (DERBY); MENDES, 1959, p. 49–52, pl. 1, figs. 1–4; MENDES, 1961, p. 12–13, figs. 11–13; ROCHA CAMPOS AND ARCHANGELSKY, 1985, pl. 6, fig. 16.

Description.Small tolmatchoffid shell (measurement see Table 3), with subquadrate outline; widest at hinge line; profile

CHEN ET AL.—UPPERMOST MISSISSIPPIAN BRACHIOPODS OF THE AMAZON BASIN TABLE 3—Measurements (in mm) of Brasilioproductus chandlessi. Specimens

Width

Length

C-length

V-height

N. of costae**

NUB-506 NUB-507 NUB-508

25.93 38.44 18.02

21.43 24.18 18.58

35.52 47.10 24.76

12.82 13.91 8.87

13 13 13

** Number of costae in 10 mm at the midlength of the shell.

strongly curved; ears large, slightly inflated. Ventral valve geniculate anteriorly; beak stout, incurved, overhanging hinge line; umbo evenly convex, with moderately steep lateral slopes; median sulcus narrow, originating anterior to beak, becoming wider and deeper at start of protrail; trail short, curved. Dorsal valve gently concave, with moderately deep corpus cavity. Ventral costae and costellae rounded, with narrower interspaces, bifurcating anteriorly; about 10–11 costae and costellae in 10 mm at start of protrail; rugae variously developed, confined to disk when present, forming reticulation with costae; spines coarse, a row along hinge, three to four spines on ears, numerous spines scattered on ribs of disk and trail; those on disk usually erect, nearly as wide as one rib in diameter, those on trail often occupying more than one rib in diameter. Dorsal ornamentation and interiors not observed. Material examined.NUB-506, NUB-507, NUB-508. Discussion.Another Itaituba Formation species Brasilioproductus chronici Mendes (1959, p. 52–54, pl. 1, fig. 5; pl. 7, figs. 1–3) was created for the productid shells that are separated from B. chandlessi by its smaller size, more elongate outline, and regularly spaced ribs. B. chandlessi also approaches externally Sandia, which has been treated as a junior synonym of Brasilioproductus by Langenheim (1991, p. 342) and Gordon et al. (1993, p. 6). However, taking into account their different internal features, Brunton et al. (2000) referred Sandia to the Productellidae and Brasilioproductus to the Productidae. Apart from the internal distinctions, the type species Sandia brevis Sutherland and Harlow (1973, p. 41, pl. 7, figs. 14–23; pl. 8, figs. 1–3) also differs from B. chandlessi in having longitudinal ridges on the trail and lacking dorsal spines. Superfamily LINOPRODUCTOIDEA Stehli, 1954 Family LINOPRODUCTIDAE Stehli, 1954 Subfamily LINOPRODUCTINAE Stehli, 1954 Genus MARGINOVATIA Gordon and Henry, 1990 MARGINOVATIA CF. CATINULUS Gordon and Henry, 1990 Figure 4.13 cf. Marginovatia catinulus GORDON 5.30.

AND

HENRY, 1990, p. 541, fig. 5.1–

Material examined.NUB-509. Discussion.A ventral valve (NUB-509), about 19 mm wide, 18.5 mm long, and 10 mm thick, is similar to M. catinulus. The diagnostic features of the species are: outline subcircular to elongate suboval; profile broadly arched; ventral beak small, slightly incurved, but not overhanging hinge line; umbo evenly convex, decreasing gradually toward anterior margin; rugae widely spaced and steplike on disk and flanks; costae beginning at umbo, intercalating anteriorly; about 13–18 costae and costellae in 5 mm at midlength of shell; numerous spines scattered on disk and ears. Externally, this species is closely allied to the type species Marginovatia minor (Snider, 1915), from which M. catinulus differs by its larger size and slightly coarser, more widely spaced ribs. In addition, the broadly arched shells, less pronounced rugae, and larger ears of M. catinulus are also noteworthy. M. eurekensis Gordon and Henry (1990, p. 543–544, fig. 6.1–6.11) from the

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Eureka Formation of Nevada is a shell intermediate in size between M. minor and M. catinulus, having 18–22 costae and costellae in 5 mm on the trail, and more numerous spines on the ears. M. aureocollis Gordon and Henry (1990, p. 544–546, fig. 7.1–7.33) from Utah is smaller than M. catinulus and has a longer trail and a narrower and more highly arched ventral umbo. M. pumila (Sutherland and Harlow, 1973, p. 59, pl. 13, figs. 7–12; Gordon and Henry, 1990, p. 547–548, fig. 9.1–9.25) from the Morrowan cannot include the described specimens as it is a small, globular species having finer ribs, and averages 19 costae and costellae in 5 mm at midlength of the ventral disk. Order ORTHOTETIDA Waagen, 1884 Suborder ORTHOTETIDINA Waagen, 1884 Superfamily ORTHOTETOIDEA Waagen, 1884 Family SCHUCHERTELLIDAE Williams, 1953 Subfamily STREPTORHYNCHINAE Stehli, 1954 Genus STREPTORHYNCHUS King, 1850 STREPTORHYNCHUS HALLIANUS Derby, 1874 Figure 4.2 Streptorhynchus hallianus DERBY, 1874, p. 35–37, pl. 5, figs. 1, 2, 5, 8, 12, 14, 16, 18; pl. 8, fig. 3; KATZER, 1903, pl. 6, fig. 6; DUARTE, 1938, p. 19–20, pl. 5, figs. 3, 4; DRESSER, 1954, p. 27–29, pl. 1, fig. 5; pl. 8, figs. 2, 3, 5, 6. Kiangsiella hallianus (DERBY); MENDES, 1956a, p. 15–18, pl. 1, figs. 1–3.

Material examined.NUB-510. Discussion.Streptorhynchus hallianus is represented by a medium-sized ventral valve (NUB-510), about 18 mm wide and 19 mm long, in the Amazon collections. The shell has a small, pointed beak settling on the moderately high, conical umbo. The costae and costellae are regularly spaced with much narrower interspaces, increasing in number by intercalation; 11 costellae are visible in the space of 5 mm at the umbo. Mendes (1956a) altered Derby’s species to Kiangsiella Grabau in Chao, 1927a. However, the described specimen and Derby’s illustrated materials possess evenly spaced costae and costellae, and lack alternative costation and a well-developed dorsal median ridge; the latter two features are diagnostic of Kiangsiella. Accordingly, S. hallianus seems not to be suitable for Kiangsiella. Family ORTHOTETIDAE Waagen, 1884 Genus TAPAJOTIA Dresser, 1954 TAPAJOTIA TAPAJOTENSIS (Derby, 1874) Figure 6.1 Streptorhynchus tapajotensis DERBY, 1874, p. 37–41, pl. 5, figs. 3, 6, 7, 9, 10; pl. 8, fig. 9. Derbyia tapajotensis (DERBY); DUARTE, 1938, p. 17, 18, pl. 4, figs. 2–4. Tapajotia tapajotensis (DERBY); DRESSER, 1954, p. 34–38, pl. 3, figs. 1– 6; pl. 4, figs. 8, 9, 11. Derbyoides tapajotensis (DERBY); MENDES, 1956a, p. 18–21, pl. 2, figs. 1–6; pl. 4, fig. 1.

Material examined.NUB-511. Discussion.An incomplete ventral valve is assigned to Derby’s species because it agrees fully with the original definition (Derby, 1874) and the illustrations (Dresser, 1954) of the species in all observed aspects. According to these authors, this Amazon species is characterized externally by weakly resupinate shells and differentiated ribs usually having two to three finer costellae intercalated between two coarser ribs, and internally by possessing a weak myophragm in the ventral valve and losses of adductor scars and myophragm in the dorsal valve. The latter features suggest that Derby’s species is not suitable for Derbyoides Dunbar and Condra, 1932.

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CHEN ET AL.—UPPERMOST MISSISSIPPIAN BRACHIOPODS OF THE AMAZON BASIN Class RHYNCHONELLATA Williams, Carlson, Brunton, Holmer, and Popov, 1996 Order ORTHIDA Schuchert and Cooper, 1932 Suborder DALMANELLIDINA Moore, 1952 Superfamily DALMANELLOIDEA Schuchert, 1913 Family RHIPIDOMELLIDAE Schuchert, 1913 Subfamily RHIPIDOMELLINAE Schuchert, 1913 Genus RHIPIDOMELLA Oehlert, 1890 RHIPIDOMELLA PENNIANA (Derby, 1874) Figures 4.14–4.32, 5.1, 5.2 Orthis penniana DERBY, 1874, p. 23, pl. 5, figs. 13, 15, 17, 19–22; pl. 8, fig. 2. Rhipidomella penniana (DERBY); KATZER, 1903, p. 172, pl. 6, fig. 9a–c; KATZER, 1933, p. 153, pl. 6, fig. 9; DUARTE, 1938, p. 13, pl. 2, figs. 2, 3; DRESSER, 1954, p. 83–85, pl. 1, figs. 1–4, 6, 7; ROCHA CAMPOS AND ARCHANGELSKY, 1985, pl. 1, figs. 20–22; pl. 6, fig. 12.

Description.Medium-sized for genus (measurements see Table 4); elongate ovate in outline; dorsibiconvex in profile; shell width at midlength of shell; anterior commissure rectimarginate. Ventral beak small, slightly incurved, extending posteriorly on interarea; umbo convex posteriorly, flattening to concave anteriorly; interarea broad, concave, apsacline, occupied medially by an open, triangular delthyrium; median sulcus variously developed, commencing anterior to beak when present. Dorsal beak small, pointed, incurved over interarea; interarea low, apsacline; notothyrium triangular, filled by cardinal process (Fig. 5.2); median sulcus shallow, distinct at umbo, becoming more prominent near anterior margin. Costellae fine, with narrower interspaces, intercalating anteriorly, with three to five costae and costellae in 2 mm near anterior margins; growth lines conspicuous, lamellose near anterior margins. Teeth prominent, bladelike; dental plates short, reaching valve floor, diverging laterally; ventral muscle field slightly elevated, forming platform; diductor scars large, flabellate, well impressed, and completely enclosing elliptical adductor scars; a myophragm narrow, low when separating adductor scars, abruptly broadening and thickening anterior to adductor scars, and then dividing diductor scars, terminating at anterior margins of muscle field; numerous genital pits lying in front of muscle field (see Fig. 5.1). Cardinal process large with numerous lobes; brachiophores strong, anteroventrally directed, diverging at 808–908, fused to valve floor by secondary deposits and posteriorly forming inner edge to sockets; sockets triangular, deep; myophragm usually secondarily thickened, tapering posteriorly, forming a low ridge separating faint posterior pair of adductor scars; anterior pair of adductor scars elliptical, indistinct (Fig. 5.2). Material examined.NUB-512, NUB-513, NUB-514, NUB515, NUB-516, NUB-517, NUB-518, NUB-519. Discussion.Although Dresser (1954, p. 85) placed Rhipidomella cora (d’Orbigny, 1842; see also Kozlowski, 1914, p. 64– 67, pl. 9, figs. 35–60) in synonymy with Derby’s species, the Bolivian species has a more circular outline and a consistently thick dorsal myophragm (see Kozlowski, 1914, fig. 15), and lacks

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TABLE 4—Measurements (in mm) of Rhipidomella penniana. Specimens

Width

Length

Thickness

N. of costae**

NUB-512† NUB-513‡ NUB-514† NUB-515 NUB-516 NUB-517 NUB-518 NUB-519

18.10 15.66 18.12 16.24 14.11 14.37 17.38 17.94

19.10 19.26 18.68 16.58 14.73 14.79 17.97 19.06

3.57 5.03 3.72 5.06 5.29 5.52 5.82 7.50

25 22 23 24 23 23 24 25

† Dorsal valve only. ‡ Ventral valve only. ** Number of costae in 10 mm at the midlength of the shell.

a prominent dorsal median sulcus; whereas R. penniana is relatively more elongate and dorsally sulcate; its dorsal myophragm thickens posteriorly and tapers anteriorly. Derby’s species also resembles superficially R. nevadensis from the Chesterian of the southwestern United States with the reflection that Langenheim (1991, p. 342) placed the latter in the synonym with R. penniana. However, Meek’s (1877) original illustrations and the topotypes of R. nevadensis figured by Morrow and Webster (1992, pl. 2, figs. 1–3) are distinguishable from R. penniana in having a more circular outline, a nearly biconvex lateral profile, and a broader median sulcus at the ventral anterior margin. Like Rhipidomella penniana, the type species R. michelini (Le´veille´, 1835) is also a dorsibiconvex species, but the latter lacks a pronounced dorsal median sulcus and has smaller muscle fields that are confined posterior to the umbo. By contrast, the muscle fields usually extend anteriorly to the midlength in the shell of R. penniana. In addition, R. penniana clearly differs from R. australis (McCoy, 1844; see also Campbell, 1957, p. 52–54, pl. 12, figs. 13–20) as the latter possesses a smaller size, a subpentagonal outline, and subparallel lateral margins; from R. burlingtonensis (Hall, 1859) (see also Weller, 1914, p. 149–151, pl. 21, figs. 5–7; pl. 83, figs. 7, 8) by its relatively narrower ventral umbo, less-developed ventral sulcus, better-developed dorsal sulcus, coarser ribs, and more pronounced lamellae near the anterior margins; from R. huntingdonensis Girty, 1928 (Carter and Kammer, 1990, p. 84, fig. 4.3–4.7) as the latter is relatively more transverse, has a narrower dorsal median sulcus and a thinner myophragm in the ventral valve, and lacks a ventral sulcus; and from R. diminutiva Rowley, 1900 (Carter, 1999, p. 112–113, fig. 6oo–ss) as the latter is proportionately more transverse and possesses a nearly biconvex profile and a weaker sulcus in both valves. Rhipidomella radiate Chen and Shi, 2003 (p. 162, pl. 9, figs. 6–12) from the Viseán and Serpukhovian of the Tarim Basin also approaches the described materials in overall appearance, but the Tarim species is significantly smaller and relatively more circular in outline, and thus cannot include the Amazon materials.

← FIGURE 4—1, Schizophoria (Schizophoria) sp. NUB-520, a ventral valve in ventral view. 2, Streptorhynchus hallianus Derby, 1874. NUB-510, a ventral valve in ventral view; 3–5, Eomarginifera oddonei (Mendes, 1959), NUB-500, NUB-501, NUB-502, three ventral valves in ventral views. 6, 7, Brasilioproductus chandlessi. NUB-498, NUB-499, two ventral valves in ventral views. 8–12, Kozlowskia petrii Mendes, 1959. 8–10, NUB503, a specimen in ventral, lateral, and dorsal views; 11, 12, NUB-504, NUB-505, two ventral valves in ventral views. 13, Marginovatia cf. catinulus Gordon and Henry, 1990. NUB-509, a ventral valve in posteroventral view. 14–32, Rhipidomella penniana (Derby, 1874). 14, 15, NUB512, a ventral valve in external and interior views; 16, 17, NUB-513, a dorsal valve in interior and external views; 18, NUB-514, a ventral valve in interior view; 19, 20, NUB-515, a specimen in ventral and dorsal views; 21, NUB-516, a specimen in dorsal view; 22–26, NUB-517, a specimen in ventral, dorsal, anterior, lateral, and posterior views; 27, 28, NUB-518, a specimen in dorsal and ventral views; 29–32, NUB-519, a specimen in ventral, dorsal, anterior and later views. All from the basal Itaituba Formation of the Caima section, Itaituba, Amazon Basin; all 31.5.

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JOURNAL OF PALEONTOLOGY, V. 79, NO. 5, 2005 Genus ORTHOTICHIA Hall and Clarke, 1892 ORTHOTICHIA MORGANIANA (Derby, 1874) Figure 6.7, 6.8 Orthis? morganiana DERBY, 1874, p. 29, pl. 3, figs. 1–7, 9, 11, 34; pl. 4, figs. 6, 14, 15. Orthotichia morganiana (DERBY); HALL AND CLARKE, 1892, p. 213, pl. 7, figs. 11–15; KATZER, 1903, p. 164, pl. 5, fig. 6a–f; KOZLOWSKI, 1914, p. 62, pl. 3, figs. 11, 12; DUARTE, 1938, p. 11, pl. 1, figs. 1, 2; DRESSER, 1954, p. 86, pl. 1, figs. 8–11, 13.

FIGURE 5—Interiors of Rhipidomella penniana. Top, Ventral valve (based on specimen NUB-512); bottom, dorsal valve (based on specimen NUB-513). t: tooth; di: diductor scar; ad: adductor scar; vm: ventral myophragm; gp: gentical pit; cp: cardinal process; s: socket; b: brachiophore; dm: dorsal myophragm; aad: anterior adductor scar; pad: posterior adductor scar. Scale bars represent 6 mm.

Superfamily ENTELETOIDEA Waagen, 1884 Family SCHIZOPHORIIDAE Schuchert, 1929 Genus SCHIZOPHORIA King, 1850 Subgenus SCHIZOPHORIA (SCHIZOPHORIA) King, 1850 SCHIZOPHORIA (SCHIZOPHORIA) sp. Figure 4.1 Discussion.A ventral valve (NUB-520), about 24.2 mm wide and 23.5 mm long, has a suboval outline and a moderately convex umbo, the margins gradually curving laterally and anteriorly; about 28 costellae are visible in 5 mm at the midlength of the shell. The shell possesses a stout beak and is slightly concave anteriorly but lacks a median sulcus; the external surface is multicostellate. These external features agree fully with S. (Schizophoria), but the inadequate material hinders the definition of a certain species.

Description.Medium-sized shells, about 16 mm wide, 13 mm long, and 7 mm thick; transversely ovate in outline; slightly ventrobiconvex in profile; hinge line narrow, about half of shell width; shell widest at midlength; cardinal extremities rounded; ventral beak low, slightly incurved, not overhanging hinge line; interarea moderately high, triangular, median sulcus broad, shallow. Dorsal valve gently convex in profile; interarea low, indistinct; median fold absent; surface finely costellate; costellae pitted and with intercalation anteriorly, about 20 in 5 mm anterior to midlength of shell. Ventral interior with dental plates thin, long, slightly diverging; median septum long, low. Dorsal interior not observed. Material examined.NUB-521. Discussion.The new material examined further affirms Dresser’s (1954) statement that the Amazon species possesses a broad, shallow ventral median sulcus, unlike the observation of Derby (1874). Chao (1927a) applied this species to several small specimens from the Lower Permian Chihsia Formation in Nanjing, South China. However, Wang et al. (1982) altered these Nanjing materials to Orthotichia indica (Waagen, 1883) described originally from the Middle Permian Wargal Formation of the Salt Range and reviewed by Grant (1976) and Chen (2004). Like O. morganiana, O. indica is also transversely ovate, but it differs from the Amazon species in having a much more inflated, dorsibiconvex profile and a rather deep ventral median sulcus. O. chekiangensis Chao (1927a), also from the Chihsia Formation of South China, differs from the Amazon species in having a significantly larger size, a more transverse outline, and a broader, deeper ventral median sulcus. Order SPIRIFERIDA Waagen, 1883 Suborder SPIRIFERIDINA Waagen, 1883 Superfamily SPIRIFEROIDEA King, 1846 Family SPIRIFERIDAE King, 1846 Subfamily SERGOSPIRIFERINAE Carter in Carter et al., 1994 Genus ANTHRACOSPIRIFER Lane, 1963 ANTHRACOSPIRIFER OLIVEIRAI (Mendes, 1956b) Figure 7.1–7.4, 7.9–7.13 Brachythyris opimus HALL; DUARTE, 1938, p. 29, pl. 1, fig. 16. Spirifer oliveirai MENDES, 1956b, p. 48–51, pl. 1, fig. 3a–d; pl. 2, figs. 1, 2.

Material examined.NUB-522, NUB-523, NUB-524, NUB525.

→ FIGURE 6—1, Tapajotia tapajotensis (Derby, 1874). NUB-511, a ventral valve in ventral view, 32; 2, 4, Phricodothyris mourai Mendes, 1956b. NUB-533, NUB-534, two dorsal valves in dorsal views, respectively, 31.5; 3, 5, 6, Composita sp. 3, NUB-537, a specimen in ventral view, 5, 6, NUB-538, a specimen in dorsal and ventral views, both 31.5. 7, 8, Orthotichia morganiana (Derby, 1874). NUB-521, a specimen in ventral and dorsal views, 31.5. 9–25, Composita caimaensis n. sp. 9, 10, NUB-539, a dorsal valve in dorsal and internal views; 11–14, NUB-540, a specimen in dorsal, anterior, ventral, and posterior views; 15, 16, NUB-541, NUB-542, two ventral valves in ventral views; 17, 18, paratype, NUB-543, a ventral valve in ventral and internal views; 19, 20, NUB-544, a specimen in ventral and dorsal views; 21–25, holotype, NUB-545, a specimen in dorsal, posterior, anterior, lateral, and ventral views, all 31.5. All from the basal Itaituba Formation of the Caima section, Itaituba, Amazon Basin.


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Discussion.Mendes (1956b) proposed Spirifer oliveirai to include spiriferid shells with ribs which rarely branched and which include three costae in the ventral sulcus, four on the dorsal fold, and seven to eleven on each lateral flank. These Amazon specimens agree perfectly with the description and illustrations of the Chesterian species Spirifer leidyi Norwood and Pratten (1855, p. 72, pl. 9, fig. 2a–c; see also Weller, 1914, p. 345–346, pl. 47, figs. 17–31), which should be assigned to Anthracospirifer (see Sutherland and Harlow, 1973, p. 77; Carter, 1990, p. 240). Accordingly, Mendes’s species may be synonymous with Anthracospirifer leidyi. However, the internal characteristics of A. leidyi remain unknown. It is hard to compare fully the Chesterian species with the Amazon specimens. For these reasons, A. oliveirai is tentatively retained for the Amazon materials, but the external resemblance between A. leidyi and Mendes’s species is noteworthy. In addition, Mendes (1956b) also referred the previously described Itaituba Formation specimens possessing five costae in the sulcus and six on the fold (e.g., Derby, 1874; Katzer, 1903; Duarte, 1938; Dresser, 1954) to A. oliveirai, but the greater number of costae in the sulcus or on the fold are diagnostic of another Chesterian species, A. increbescens (Hall, 1858; Weller, 1914; Sutherland and Harlow, 1973). These specimens therefore should be excluded from A. oliveirai. Another ally to Mendes’s species is A. brenckelei Carter (1990, p. 238–241, fig. 9.1–9.25) from the Late Osagean of the Upper Mississippi Valley, but Carter’s species differs from A. oliveirai in possessing relatively finer, more angular ribs, and slightly rounded to subangular extremities. The type species A. birdspringensis Lane (1963, p. 388–390, pl. 44, figs. 4, 7, 10–18; pl. 45, fig. 3) has an overall appearance close to that of the Amazon species, from which Lane’s species is differentiated by the possession of a less incurved ventral beak, a more strongly grooved ventral interarea, three to eight lateral sulcal costae, and common lateral bifurcations. Another Pennsylvanian species, A. opimus (Hall, 1858; see also Dunbar and Condra, 1932; Hoare, 1961; Spencer, 1967; Sturgeon and Hoare, 1968), also has the same total number of ribs in the sulcus and on the fold, and thus may be confused with A. oliveirai, from which Hall’s species differs in having a smaller size, a more rectangular outline, a higher ventral interarea, and a more strongly arched ventral beak. ANTHRACOSPIRIFER sp. Figure 7.14, 7.15 Discussion.One specimen (NUB-526), about 29 mm wide, 27.5 mm long, and 16.5 mm thick, is present in the Amazon collections. The important characteristics of the specimen are: outline subrectangular; lateral profile nearly biconvex; widest at hinge line; ventral beak pointed, incurved, overhanging interarea; interarea moderately high, triangular, concave, vertically grooved, with a median triangular, open delthyrium; median sulcus broad, deep, commencing at beak, defined by two sharp sulcus-bounding costae from lateral areas; dorsal valve evenly convex, with a median fold that broadens and elevates highly anteriorly; costae subangular to rounded in cross section; one simple median sulcal costa with two lateral sulcal costae on each side; lateral sulcal

costae derived from sulcus-bounding costae; six costae on fold; about 12 costae on each lateral area; pair of costae next to sulcus (or fold) bifurcating once, others simple. Comparable specimens from the Itaituba Formation have been previously referred to A. opimus (see Derby, 1874, p. 15–16, pl. 1, fig. 4; pl. 2, fig. 7; pl. 4, fig. 12) and A. rockymontanus (Marcou, 1858) (see Katzer, 1903; Duarte, 1938; Dresser, 1954). However, according to Dunbar and Condra (1932), the true A. opimus possesses three costae in the ventral median sulcus and four on the fold, and thus cannot include the Amazon specimens. Although A. rockymontanus (see also Dunbar and Condra, 1932; Hoare, 1961; Sturgeon and Hoare, 1968; Sutherland and Harlow, 1973) also possesses five costae in the sulcus and six on the fold, Marcou’s species clearly differs from the Amazon materials by its significantly smaller size [lectotype selected by Girty (1903, p. 384) is about 16 mm wide and 15.8 mm long], higher ventral interarea, less incurved ventral beak, and a lingual extension at the anterior margin. The appearance of these Amazon specimens, overall, approaches the Chesterian Spirier increbescens Hall (1858, p. 706, pl. 27, fig. 6a–i; Weller, 1914, p. 343–344, pl. 46, figs. 1–12), which was transferred to Anthracospirifer by Sutherland and Harlow (1973, p. 77). Like the Chesterian species, the Amazon specimens also possess a similar size, a subrectangular outline, a nearly biconvex lateral profile, a moderately high, triangular, concave interarea, and comparable ribbings. However, absence of cardinal extremities hampers a comparison with the distinctive, slightly mucronate extremities of A. increbescens and the rib count on the lateral slopes. Only 12 ribs are visible on the each broken lateral slope of the described shell, whereas 14–18 ribs are present on each lateral slope of the Chesterian species. Therefore, the inadequate and poorly preserved material prevents us from assigning a certain species name, although this specimen is comparable with A. increbescens in many respects. Subfamily SPIRIFERINAE King, 1846 Genus MESOCHORISPIRA Carter, 1992 MESOCHORISPIRA DRESSERI (Mendes, 1956b) Figure 7.18–7.22 Spirifer camerata MORTON; DERBY, 1874, p. 12, pl. 1, figs. 1, 3, 6, 9, 14; pl. 2, figs. 2, 15; pl. 4, fig. 4; pl. 5, fig. 11; KATZER, 1903, p. 164, pl. 4, fig. 1. Spirifer (Neospirifer) cameratus (MORTON) variant; DRESSER, 1954, p. 61, pl. 5, fig. 11; pl. 8, figs. 1, 4. Neospirifer dresseri MENDES, 1956b, p. 52, pl. 2, figs. 3–5; pl. 3, figs. 1–5; pl. 4, fig. 2a–b; ROCHA CAMPOS AND ARCHANGELSKY, 1985, pl. 6, fig. 9.

Material examined.NUB-528, NUB-529, NUB-530, NUB531, NUB-532. Discussion.The materials examined agree perfectly with Mendes’s (1956b) description of the species. The diagnostic features are: medium to large shells, with transversely ovate outline; cardinal extremities subangular to slightly rounded; costae rounded posteriorly, flattening anteriorly, with narrower interspaces; →

FIGURE 7—1–4, 9-13, Anthracospirifer oliverirai (Mendes, 1956b). 1, 2, NUB-522, a ventral valve in ventral and posterior views; 3, 4, NUB-523, NUB-524, two ventral valves in ventral views; all 32. 9–13, NUB-525, a specimen in dorsal, posterior, anterior, ventral, and lateral views; all 32. 5–7, Spiriferidae genus and species undetermined. NUB-535, a ventral valve in ventral, internal, and posterior views, 31.5. 8, Punctospirifer sp. NUB-536, a dorsal valve in dorsal view, 31.5. 14, 15, Anthracospirifer sp. NUB-526, a specimen in dorsal and ventral views, both 31.8. 16, 17, Spirifer sp. NUB-527, a broken specimen in posterior and ventral views, 31.5. 18–22, Mesochorispira dresseri (Mendes, 1956b). 18, 19, NUB528, a ventral valve in ventral and internal views; 20–22, NUB-532, a ventral valve in ventral, interior, and lateral views; all 32. All from the basal Itaituba Formation of the Caima section, Itaituba, Amazon Basin.


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three bundles of costae close to median sulcus bifurcate three to four times, but not fasciculate; costae on each flank bifurcate once or simple; ventral interarea high, broad, concave, marked by vertical grooves and horizontal striation, occupied medially by a large, open, triangular delthyrium; dental plates well developed, diverging anterolaterally, consisting of dental flanges and ventral adminicula. We agree with Mendes (1956b) that these Amazon specimens represent a distinct species independent from Spirifer camerata, but disagree with him that his new species is suitable for Neospirifer. According to Poletaev (1997), the true Neospirifer species possesses distinctive fasciculation of the costae; whereas the neotype of Spirifer striatus (Martin, 1793), type species of Spirifer, has incipient fascicles next to the sulcus or fold (Archbold and Thomas, 1984; see also Chen and Archbold, 2000). The Amazon specimens often possess three bundles of costae next to the fold or sulcus on each flank, its costae continue to branch some 6 mm from the umbo, and are not bifurcated symmetrically and fasciculated. As such, these Amazon specimens are more suitable for Spirifer than Neospirifer. However, Carter (1992) created the subgenus Spirifer (Mesochorispira) to include those species possessing a double reversal in growth form, from rounded cardinal extremities in juveniles, to slightly transverse mucronate extremities at midsize, and back to rounded cardinal extremities in adults. Those Spirifer species that have a transversely subtrigonal to subsemicircular outline and alate, mucronate, or angular cardinal extremities in all growth stages are retained to the subgenus S. (Spirifer). When compared with Spirifer striatus (s. Archbold and Thomas, 1984), ‘‘N.’’ dresseri clearly differs in its relatively less inflated ribs and the cardinal extremities that are subangular, not alate or mucronate. Mendes’s species therefore is not suitable for S. (Spirifer). In contrast, these features are closely allied to that of Spirifer grimesi Hall, 1858, type species of S. (Mesochorispira), from the Tournaisian of the United States (see also Weller, 1914; Carter, 1992, 1999) in all observed aspects. As a result, Mendes’s species is assigned to S. (Mesochorispira), which has been upgraded to a full genus by Poletaev (1999) and Sartenaer and Plodowski (2002). The major difference between M. grimesi (Hall, 1858) and M. dresseri is that Hall’s species is very large, has a lower ventral interarea, a more strongly incurved ventral beak, and a more pronounced, angularly floored ventral median sulcus. Another Lower Carboniferous species, M. konincki (Dewalque, 1895), well illustrated and redescribed by Sartenaer and Plodowski (2002, pls. 1–5) from Belgium, is also similar in overall appearance to M. dresseri, from which the Belgian species clearly differs in having a proportionately more elongate outline, more rounded extremities, and less distinctive sulcus-bounding costae. Genus SPIRIFER Sowerby in J. Sowerby and J. de Sowerby, 1818–1821 SPIRIFER sp. Figure 7.16, 7.17 Description.Medium-sized for genus; outline appears to be transversely ovate; ventral beak pointed, incurved, and close to dorsal umbo; ventral umbo evenly convex, with gentle slopes to lateral margins; interarea low, broad, subrectangular, concave, marked by numerous vertical and horizontal striations; delthyrium wide, open; median sulcus deep, well defined by a pair of distinct sulcus-bounding costae. Dorsal beak small, incurved; median fold distinct, originating at beak. External costae rounded in cross section; median sulcal costa simple; lateral sulcal costae bifurcating; three primary costae on fold, of these, lateral costae bifurcating twice; three bundles of costae close to fold or sulcus at each flank, with primary costae originating at beaks, and bifurcating two to

three times; seven pairs of costae close to extremities, simple. Concentric growth lines fine, dense. Discussion.As discussed above, Carter (1992) retained Spirifer to include these species having a transversely subtrigonal to subsemicircular outline and alate, mucronate, or angular cardinal extremities in all growth stages. The incipient fasciculation of the costae suggests the described specimen is suitable for Spirifer rather than Neospirifer. Mesochorispira is another possible genus to include the present materials, however, unlike M. grimesi, the Amazon material appears to have more frequently bifurcating costae next to the sulcus or fold and a more transverse outline; the cardinal extremities extend laterally and somewhat appear slightly alate, although they are broken. Nevertheless, a broken specimen (NUB-527) is insufficient to define fully a certain species. Mesochorispira dresseri, described above, cannot include the present specimen as it has a less transverse outline and flatter ribs that are less pronounced and fasciculated. SPIRIFERIDAE genus and species undetermined Figure 7.5–7.7 Discussion.A broken ventral valve (NUB-535) possesses a small, pointed, slightly incurved beak, and a large, slightly apsacline interarea that is occupied medially by a triangular delthyrium. The delthyrium, with delthyrial angle of 458–508, extends about 5 mm from the beak. The umbo is medially convex and has an apical angle of about 1258–1298. Lateral slopes are weakly costate; the costae are diminished and almost invisible. Internally, the dental plates are rather thick, diverge, and comprise dental flanges and outward-curved, short ventral adminicula that partially encompass the muscle field. The median septum is absent. The weak ribbings, indistinctive median sulcus, and slightly apsacline interarea distinguish the described specimen from any known genera and species of the Spiriferidae. These features also recall Pseudosyrinx sp. of Isaacson and Dutro [1999, p. 631, fig. 3.8 (left)] from the upper Zorritas Formation (Tournaisian), northern Chile and Pseudosyrinx cf. missouriensis Weller, 1914 of Carter (1999, p. 133, fig. 21a–d) from the Tournaisian of Oklahoma. However, the Amazon specimen is not a Pseudosyrinx species due to its slightly apsacline interarea, incipient delthyrial structure, and lack of punctuate shell. Whereas Pseudosyrinx Weller, 1914 is usually punctated and possesses a high, procline, flat ventral interarea and a distinctive delthyrial plate. These distinctions suggest the present ventral valve cannot be assigned to any known genera. However, the generic and specific assignment remains undetermined due to insufficient material. Suborder DELTHYRIDINA Ivanova, 1972 Superfamily RETICULARIOIDEA Waagen, 1883 Family ELYTHIDAE Fredericks, 1924 Subfamily PHRICODOTHYRIDINAE Caster, 1939 Genus PHRICODOTHYRIS George, 1932 PHRICODOTHYRIS MOURAI Mendes, 1956b Figure 6.2, 6.4 Spirifer (Martinia) perplexa (MCCHESNEY); DERBY, 1874, p. 16, pl. 3, figs. 27, 39, 40, 45, 50; pl. 8, fig. 13. Reticularia perplexa (MCCHESNEY); KATZER, 1903, p. 172, pl. 6, fig. 11a–b. Squamularia perplexa (MCCHESNEY); DUARTE, 1938, p. 31, pl. 5, figs. 10–12. Phricodothyris perplexa (MCCHESNEY); DRESSER, 1954, p. 50, pl. 6, figs. 1–4, 6. Phricodothyris mourai MENDES, 1956b, p. 44, pl. 1, figs. 1, 2; pl. 5, fig. 4.

Material examined.NUB-533, NUB-534. Discussion.A full description of the species can be found in Mendes (1956b). The described specimens are nearly equidimensional to slightly transverse in outline and moderately biconvex


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FIGURE 8—Serial sections of Composita caimaensis n. sp. (based on specimen NUB-546). The numbers are distance from the ventral beak; the direction of the sections is with ventral valve downward. hp: hinge plate.

in profile. Their growth lamellae are regularly spaced, about seven lamellae in 5 mm at the midlength of the shell; the edges of lamellae bear elongate impressions of the double-barrelled spines. P. mourai resembles superficially Phricodothyris perplexa (McChesney, 1860), a widespread species of the Pennsylvanian in the US (e.g., Girty, 1915; Dunbar and Condra, 1932; Hoare, 1961; Sturgeon and Hoare, 1968; Sutherland and Harlow, 1973), but the latter is differentiated from Mendes’s species by its ventrobiconvex profile, more transverse dorsal valve with a lower umbo, and narrower lamellae. Order SPIRIFERINIDA Ivanova, 1972 Suborder SPIRIFERINIDINA Ivanova, 1972 Superfamily PENNOSPIRIFERINOIDEA Dagys, 1972 Family PUNCTOSPIRIFERIDAE Waterhouse, 1975 Genus PUNCTOSPIRIFER North, 1920 PUNCTOSPIRIFER sp. Figure 7.8 Discussion.One incomplete dorsal valve (NUB-536) is assigned to Punctospirifer as it is medium-sized, transversely triangular in outline; the dorsal valve possesses a highly elevated median fold, originating at the beak, becoming more pronounced anteriorly; the lateral slopes are gentle. External plicae are high, rounded in cross section, with rounded interspaces, about eight plicae on each flank; the concentric lamellae are regularly imbricate, densely arranged near the anterior margins, and widely spaced on the anterior part of the fold; numerous tiny conelike spine bases contrasting with hairlike spines are visible at front of lamellae, with punctae scattered on the shell. The observed features agree substantially with P. leinzi Mendes (1956b, p. 62–66, pl. 2, fig. 6; pl. 5, figs. 1–3) from the Itaituba Formation and P.? TABLE 5—Measurements (in mm) of Composita caimaensis n. sp. Specimens

Width

Length

Thickness

NUB-539† NUB-540 NUB-541† NUB-542† NUB-543 NUB-544 NUB-545‡ NUB-546

26 28.2 26 28 26.5 28.5 32 31.4

.20 20.1 .18 24 20.5 26 26.5 26.8

5 18 4.5 5.5 5.1 14 14.5 14.4

† Ventral valve only. ‡ Holotype.

subtexta (White, 1862) of Carter (1999, p. 135, figs. 21e–x, 22, 23) from the Tournaisian of Oklahoma. However, a more meaningful comparison with Mendes’s species and Carter’s specimens requires further collections. Order ATHYRIDIDA Boucot, Johnson, and Staton, 1964 Suborder ATHYRIDIDINA Boucot, Johnson, and Staton, 1964 Superfamily ATHYRIDOIDEA Davidson, 1881 Family ATHYRIDIDAE Davidson, 1881 Subfamily SPIRIGERELLINAE Grunt, 1965 Genus COMPOSITA Brown, 1845 COMPOSITA sp. Figure 6.3, 6.5, 6.6 Material examined.NUB-537, NUB-538. Discussion.The Amazon material examined is elongate-ovate in outline and has a high, narrow ventral umbo and rounded cardinal extremities, with the greatest width at the midlength of the shell. The external surfaces bear prominent, densely concentrically arranged growth lines and lamellae. The overall appearance agrees substantially with Derby’s (1874) original description of ‘‘Athyris subtilita Hall,’’ which was altered to his new species, Composita reedi, by Mendes (1957, 1961). However, the fold and sulcus of our specimens are distorted because of compaction. As such, these poorly preserved specimens do not allow specific assignment. COMPOSITA CAIMAENSIS new species Figures 6.9–6.25, 8 Athyris subtilita (HALL); DERBY, 1874, pl. 6, fig. 2.

Diagnosis.Subpentagonal to transversely ovate Composita, with ventrobiconvex profile, indistinct sulcus, and fold. Description.Medium-sized for genus (measurements see Table 5); outline subpentagonal to transversely ovate, slightly wider than long; widest at midlength of shell; ventribiconvex in profile; anterior commissure moderately uniplicate; cardinal extremities rounded. Ventral valve evenly convex; beak stout, rounded, strongly incurved, overhanging dorsal beak; foramen proportionately large, subcircular, usually interrupted by wide delthyrium; umbo evenly convex, gradually sloping to lateral margins; palintrope narrow, moderately high, triangular, concave; delthyrium occluded by dorsal beak; median sulcus indistinct at umbo, becoming conspicuous near anterior margin where it rapidly deepens and widens, forming a short lingual extension. Dorsal valve

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moderately inflated; beak small, pointed, incurved; umbo gently convex; lateral slopes evenly convex; median fold low, indistinct, produced in posterior half portion of the shell. External surfaces with fine growth lines and widely spaced lamellae; shell substance moderately thick. Ventral interior with strong dental plates, supporting elongate but thin teeth; dorsal interior sockets deep with thin socket plates; hinge plates united, subquadrate in shape (Fig. 8). Etymology.Named for the Caima village, Itaituba, Amazon Basin. Types.NUB-545 is selected as holotype; NUB-543 is selected as paratype. From the basal Itaituba Formation of the Caima Quarry 1 section. Other material examined.NUB-539, NUB-540, NUB-541, NUB-542, NUB-544, NUB-546, and 12 unregistered specimens. Of these, NUB-546 is sectioned. Occurrence.The basal Itaituba Formation of the Chesterian (late Mississippian) age of the Caima Quarry 1 section of the Itaituba area, the Amazon Basin, Brazil. Discussion.Of the specimens ascribed to ‘‘Athyris subtilita’’ by Derby (1874), most of them were collected from Bomjardim (see Derby, 1874, pl. 1, figs. 5, 7, 8; pl. 3, figs. 8, 16, 19; pl. 9, fig. 4) and agree with his description that they are longer than wide (Derby, 1874, p. 7). However, a ventral mold from the Itaituba Beach (see Derby, 1874, pl. 6, fig. 2) is transversely ovate in outline, and thus does not fit Derby’s description. Therefore, both Bomjardim and Itaituba materials seem not to be conspecific. We agree with Mendes (1957) that these Amazon specimens represent a new species C. reedi as the true C. subtilita, figured by Boucot et al. (1965, fig. 537.2a–c), possesses a more elongate outline and pronounced concentric growth lines. However, C. reedi is retained only for these Bomjardim specimens, while the Itaituba specimen of Derby (1874) should be excluded from Mendes’s species due to its transverse outline. In contrast, this Itaituba specimen is closely allied to our specimens in terms of size, shell outline, and other respects, and thus is assignable to the new species. Except for the different outline, C. reedi is also distinct from the new species in having a relatively deeper tongue and a higher fold at the anterior margins. The latter features also distinguish the Pennsylvanian C. ohioense Sturgeon and Hoare (1968, pl. 19, figs. 4–14) from the new species, although they share comparable size, shell outline, and ornamentation. The subpentagonal to ovate outline and more weakly developed sulcus and fold differentiate the new species from the type species C. ambiguus (Sowerby, 1822) redescribed by Grunt (1980), Brunton (1980), and Chen et al. (2003). Of the Composita species described by Carter (1987) from the Tournaisian Banff Formation of Alberta, Canada, C. athabaskensis Warren (in Allan et al., 1932) of Carter (1987, p. 61, pl. 21, figs. 17– 36) is the most similar species to C. caimaensis, both possessing a subpentagonal to ovate outline and comparable ornamentation; however, this Canadian species can not be confused with our specimens as it has a significantly smaller size, a nearly biconvex profile, and less pronounced lamellae. C. matutina Carter (1988, p. 53, figs. 15.1–15.20, 16) from the Glen Park Formation of Illinois and Missouri, USA, also possesses a subpentagonal outline and somewhat approaches the new species, from which the Carter’s species is distinct in being smaller and having a more pronounced sulcus and fold. Another Mississippian ally, Composita prolixa Carter (1987, p. 66, pl. 20, figs. 13–28; see also Chen and Shi, 2003, p. 171, pl. 11, figs. 27–29, 34, 35), also cannot be confused with the described species as it is significantly smaller and more elongate. ACKNOWLEDGMENTS

We are grateful to R. Matsumoto from The University of Tokyo, Japan, and A. K. Scomazzon from Universidade Federal do

Rio Grande do Sul, Brazil, for their kind help in the field and discussing the sedimentary sequences of the Amazon Basin. A. K. Scomazzon is particularly acknowledged for providing the unpublished conodont data. Our thanks also extend to the following companies (or institutions): Companhia Agro-Industrial Monte Alegre-CAIMA, The University of Tokyo, and PETROBRAS S. A. (Brazilian Oil Company) for their permission to access the studied outcrops and for providing financial and logistic support for the field work. P. R. Teixeira of the PETROBRAS S. A. is also thanked for his discussion on the regional geology of the Caima area. The constructive suggestions from both reviewers, C. H. C. Brunton of The Natural History Museum, London, and J. L. Carter of the Carnegie Museum, Pittsburgh, have greatly improved this paper. This study was supported by an aid grant from the Japanese Society for Promotion of Sciences (JSPS P01103 to ZQC) and a discovery grant from the Australian Research Council (DP0452296 to ZQC). REFERENCES

ALLAN, J. A., P. S. WARREN, AND R. L. RUTHERFORD. 1932. A preliminary study of the eastern ranges of the Rocky Mountains in Jasper Park, Alberta. Transactions of the Royal Society of Canada, 26, series 3, section, 4:225–249. ALTINER, D., AND R. SAVINI. 1995. Pennsylvanian foraminifera and biostratigraphy of the Amazonas and Solimoes basins (North Brazil). Revue de Paleóbiologie, 14(2):417–453. ALVAREZ, F., AND JIA-YU RONG. 2002. Athyridida, p. 1475–1604. In A. Williams, C. H. C. Brunton, and S. J. Carlson (eds.), Treatise on Invertebrate Paleontology, Pt. H, Brachiopoda. Volumes 2–3. Geological Society of America and The University of Kansas, Lawrence. ARAUJO, J. S., AND A. C. ROCHA CAMPOS. 1969. Conodonts from the Itaituba Formation (Pennsylvanian) at well Pest-2-Am, Amazon Basin. Ciencia Cultura, 21(2):221–222. ARCHBOLD, N. W., AND G. A. THOMAS. 1984. Neospirifer Fredericks, 1924 (Spiriferida, Brachiopoda): A review. Journal of Paleontology, 58:626–635, 4 pls. BOUCOT, A. J., J. G. JOHNSON, AND R. D. STATON. 1964. On some atrypoid, retzioid, and athyridoid Brachiopoda. Journal of Paleontology, 38(5):805–822, pls. 125–127. BOUCOT, A. J., J. G. JOHNSON, AND R. D. STATON. 1965. Suborders Retziidina and Athyrididina, p. H649–H667. In R. C. Moore (ed.), Treatise on Invertebrate Paleontology, Pt. H, Brachiopoda. Volume 2. Geological Society of America and The University of Kansas Press, Lawrence. BROWN, T. 1845. Illustrations of the Fossil Conchology of Great Britain and Ireland, With Descriptions and Localities of All Species. London, p. 117–136. BRUNTON, C. H. C. 1966. Silicified productids from the Viseán of County Fermanagh. Bulletin of the British Museum (Natural History) Geology, 12(5):173–243. BRUNTON, C. H. C. 1980. Type specimens of some Upper Palaeozoic Athyrididae brachiopods. Bulletin of the British Museum, London, 34: 219–234. BRUNTON, C. H. C., AND S. S. LAZAREV. 1997. Evolution and classification of the Productellidae (Productida), Upper Paleozoic brachiopods. Journal of Paleontology, 71:381–394. BRUNTON, C. H. C., F. ALVAREZ, AND D. I. MACKINNON. 1996. Morphological terms used to describe the cardinalia of articulate brachiopods: Homologies and recommendations. Historical Biology, 11:9–41. BRUNTON, C. H. C., S. S. LAZAREV, AND R. E. GRANT. 1995. A review and new classification of the brachiopod order Productida. Palaeontology, 38:915–936. BRUNTON, C. H. C., S. S. LAZAREV, R. E. GRANT, AND J. G. JIN. 2000. Productidina, p. 424–609. In A. Williams, C. H. C. Brunton, and S. J. Carlson (eds.), Treatise on Invertebrate Palaeontology, Pt. H, Brachiopoda (revised). Volume 3. Linguliformea, Craniiformea and Rhynchonelliformea (part). Geological Society of America and The University of Kansas Press, Lawrence. BUCKMAN, S. S. 1906. Brachiopod Nomenclature: Epithyris, Hypothyris,

CHEN ET AL.—UPPERMOST MISSISSIPPIAN BRACHIOPODS OF THE AMAZON BASIN Cleiothyris Phillips, 1841. Annals and Magazine of Natural History, series 7, 18:321–327. CAMPBELL, K. S. W. 1957. A Lower Carboniferous brachiopod-coral fauna from New South Wales. Journal of Paleontology, 31(1):34–98. CAPUTO, M. V., AND J. L. CROWELL. 1985. Migration of glacial centers across Gondwana during Paleozoic Era. Geological Society of America Bulletin, 96:1030–1036. CARTER, J. L. 1987. Lower Carboniferous brachiopods from the Banff Formation of Western Alberta. Geological Survey of Canada Bulletin, 53(378), 99 p., 29 pls. CARTER, J. L. 1988. Early Mississippian Brachiopods from the Glen Park Formation of Illinois and Missouri. Bulletin of the Carnegie Museum, Number 27, 82 p. CARTER, J. L. 1990. New brachiopods (Brachiopoda: Articulata) from the Late Osagean of the Upper Mississippi Valley. Annals of Carnegie Museum, 59(3):219–247. CARTER, J. L. 1992. New genera of Lower Carboniferous spiriferid brachiopods (Brachiopoda: Spiriferida). Annals of Carnegie Museum, 61(4):327–338. CARTER, J. L. 1999. Early Osagean (Upper Tournaisian) brachiopods from a bioherm in the St. Joe Formation near Kenwood, Oklahoma. Annals of Carnegie Museum, 68(2):91–149. CARTER, J. L., AND T. W. KAMMER. 1990. Late Devonian and Early Carboniferous brachiopods (Brachiopoda, Articulata) from the Price Formation of West Virginia and adjacent areas of Pennsylvania and Maryland. Annals of Carnegie Museum, 59:77–103. CARTER, J. L., AND V. I. POLETAEV. 1998. Atokan (Late Bashkirian or early Moscovian) brachiopods from the Hare Fiord Formation of Ellesmere Island, Canadian Arctic Archipelago. Annals of Carnegie Museum, 67:105–180. CARTER, J. L., J. G. JOHNSON, R. GOURVENNEC, AND H. F. HOU. 1994. A revised classification of the spiriferid brachiopods. Annals of Carnegie Museum, 63(4):327–374. CASTER, K. E. 1939. A Devonian fauna from Colombia. Bulletins of American Paleontology, 24(83), 218, 14 pls. CHAO, Y. T. 1927a. Brachiopod fauna of the Chihsia Limestone. Geological Society of China Bulletin, 6(2):83–120, 2 pls. CHAO, Y. T. 1927b. Productidae of China. Pt. I: Productinae. Palaeontologica Sinica, series B, 5(2):1–192. CHEN, Z. Q. 2004. Lower Permian reef-dwelling brachiopod faunas from the Tarim Basin, Northwest China: Biostratigraphy, palaeoecology and biogeography. Palaeontolographica Abteilung A, 272:1–96, 12 pls. CHEN, Z. Q., AND N. W. ARCHBOLD. 2000. Tournaisian–Viseán brachiopods from the Gancaohu area of the Tienshan Mountains, Xinjiang, Northwest China. Geobios, 32(2):183–199. CHEN, Z. Q., AND G. R. SHI. 2003. Lower Carboniferous brachiopods from the Heshilafu Formation of western Kunlun Mountains, NW China. Palaeontographica Abteilung A, 268:103–187. CHEN, Z. Q., G. R. SHI, AND L. P. ZHAN. 2003. Early Carboniferous brachiopods from the Qaidam Basin, northwest China. Journal of Paleontology, 77(5):844–862. CHEN, Z. Q., J. TAZAWA, G. R. SHI, AND N. S. MATSUDA. 2004. Pennsylvanian (Carboniferous) brachiopods from the Itaituba Formation of the Amazon Basin, Brazil. Alcheringa, 28(2):441–468. CONIL, R., E. GROESSENS, M. LALOUX, E. POTY, AND F. TOURNEUR. 1990. Carboniferous guide foraminifera, corals and conodonts in the Franco-Belgian and Campine Basins: Their potential for widespread correlation. Courier Forschungsinstitut Senckenberg, 130:15–30. DAEMON, R. F., AND C. J. A. CONTREIRAS. 1971. Zoneamento palinolo´gica da Bacia do Amazonas. Anais XXV Congress Brasileira Geologia, Sao Paulo. Sociedade Brasileira de Geologia, 3:79–88. DAGYS, A. S. 1972. Morfologiia i sistematika mezozoiskikh retsioidnykh brakhiopod. In Morfologicheskie i filogeneticheskie voprosy Paleontologii. Akademiia Nauk SSSR, Sibirskoe Otdelenie, Institut Geologii i Geofiziki Trudy, 112:94–105. (In Russian) DAVIDSON, T. 1881. On genera and species of spiral-bearing Brachiopoda, from specimens developed by the Rev. Norman Glass. Geological Magazine, 8:1–13. DERBY, O. A. 1874. On the Carboniferous Brachiopoda of Itaituba, Rio Tapajo´s, Province of Para´, Brazil. Cornell University Science Bulletin, 1(2):1–63, 9 pls.

923

DERBY, O. A. 1894. The Amazonian Upper Carboniferous fauna. Journal of Geology, 2(5):480–501. DEWALQUE, G. 1895. Sur Spirifer mosquensis auct. Annales de la Socie´te´ geólogique de Belgique Bulletin, 22(1894–1895):56–57. D’ORBIGNY, A. 1836–1843. Voyages dans l’Ame ´ rique Me´ridionale. Paleóntologie 3(4). P. Bertrand, Paris, 188 p. (In French) D’ORBIGNY, A. 1842. Voyages dans l’Ame ´ riques Me´ridionale. Geólogie, Paleóntologie; Foraminife`res, 3:50–56. Pitois-Levrault, Paris. (In French) DRESSER, H. 1954. Notes on some Brachiopods from the Itaituba Formation (Pennsylvanian) of the Tapajos River, Brazil. In K. E. Caster and H. Dresser (eds.), Contributions to Knowledge of the Brazilian Paleozoic. No. 1B. Bulletins of American Paleontology, 35(149):15– 84, 8 pls. DUARTE, A. G. 1938. Brachiopodos do rio Parauary. Servico Geologico e Mineralogico do Brasil Boletim, 84:1–38, pls. 1–6. DUME´RIL, A. M. C. 1806. Zoologie Analytique ou Methode Naturelle de Classı´fication des Animaux. Allais, Paris, 344 p. DUNBAR, C. O., AND G. E. CONDRA. 1932. Brachiopoda of the Pennsylvanian System in Nebraska. Nebraska Geological Survey Bulletin Series, 2(5), 377 p., 44 pls. DUNN, D. L. 1965. Late Mississippian conodonts from the Bird Spring Formation in Nevada. Journal of Paleontology, 39:1145–1150, pl. 140. DUNN, D. L. 1966. New Pennsylvanian platform conodonts from southwestern United States. Journal of Paleontology, 40:1294–1303, pls. 157, 158. DUNN, D. L. 1970. Middle Carboniferous conodonts from western United States and phylogeny of the platform group. Journal of Paleontology, 44:312–342, pls. 61–64. EASTON, W. H. 1962. Carboniferous formations and faunas of central Montana. United States Geological Survey Professional Papers, 348: 1–126, 13 pls. ELLISON, S., AND R. W. GRAVES JR. 1941. Lower Pennsylvanian (Dimple Limestone) conodonts of the Marathon region, Texas. Missouri University, School of Mines and Metallurgy Bulletin, Technology Series, 14(3):1–21, 3 pls. FREDERICKS, G. N. 1924. Aravanskaia kamennougol’naia fauna. Izvestiia Geologicheskogo Komiteta, 42(5–9):183–196, pl. 5. (In Russian) FREDERICKS, G. N. 1923. Paleontologicheskie etiudy. 2. O verkhne-kamennougol’nykh spiriferidakh Urala. Izvestiia Geologicheskogo Komiteta, 38(3):295–324. (In Russian) FREDERICKS, G. N. 1933. Paleontologicheskie etiudy, 4. O nekotorykh verkhne-paleozoiskikh brakhiopodakh Evrazii. Materialy Tsentral’nogo Nauchno-Issledovatel’skogo Geologo-Razvedochnogo Instituta, Paleontologiia i Stratigrafiia Sbornik, 2:24–33. (In Russian) FULFARO, V. J. 1965. Conodontes do calcario Itaituba do Carbonifero do Rio Tapajos, Estado do Para. Geologia Sociedade Brasileira de Geologia Boletim, 14(1–2):29–40. GEORGE, T. N. 1932. The British Carboniferous Reticulate Spiriferidae. Quarterly Journal of the Geological Society of London, 88(4):516–575, pls. 31–35. GIRTY, G. H. 1903. The Carboniferous formation and faunas of Colorado. United States Geological Survey Professional Paper, 16, 546 p., 10 pls. GIRTY, G. H. 1915. The fauna of the Batesville sandstone of northern Arkansas. U.S. Geological Survey Bulletin, 593, 170 p. GIRTY, G. H. 1928. The Pocono fauna of the Broad Top Coal Field, Pennsylvania. U.S. Geological Survey Bulletin, 51, 355 p. GORDON JR., M. 1975. Brachiopods of the Amsden Formation (Mississippian and Pennsylvanian) of Wyoming. U.S. Geological Survey Professional Paper, 848:1–86. GORDON JR., M., AND T. W. HENRY. 1990. Marginovatia, a mid–Carboniferous genus of Linoproductid brachiopods. Journal of Paleontology, 64(4):532–551. GORDON JR., M., T. W. HENRY, AND J. D. TREWORGY. 1993. Late Mississippian productoid brachiopods Inflatia, Keokukia, and Adairia, Ozark region of Oklahoma and Arkansas. Journal of Paleontology Memoir, 30:1–29. GRANT, R. E. 1976. Permian brachiopods from southern Thailand. Journal of Paleontology Memoir, 9:1–269, 71 pls. GRAY, J. E. 1840. Synopsis of the Contents of the British Museum (fortysecond edition). London, 370 p.

924


GROVES, J. R., T. I. NEMYROVSKA, AND A. S. ALEKSSEEV. 1999. Correlation of the type Bashkirian Stage (Middle Carboniferous, South Ural) with the Morrowan and Atokan Series of the midcontinental and western United States. Journal of Paleontology, 73(3):529–539. GRUNT, T. A. 1965. Nadsemeistvo Athyridacea, p. 237–253. In V. E. Ruzhencev and T. G. Sarycheva (eds.), Razvitic i smena morskikh organizmov na rubezhe paleozoia i mesozoia. Akademiia Nauk SSSR Trudy Paleontologicheskogo Institut, 108, 430 p. GRUNT, T. A. 1980. Atirididy Russkoi platformy. (Athyridides of Russian Platform.) Akademiia Nauk SSSR, Trudy Paleontologicheskogo Institut, 182, 164 p. (In Russian) GUNNELL, F. H. 1931. Conodonts from the Fort Scott Limestone of Missouri. Journal of Paleontology, 5:244–252, pl. 29. GUNNELL, F. H. 1933. Conodonts and fish remains from the Cherokee, Kansas City, and Wabaunsee groups of Missouri and Kansas. Journal of Paleontology, 7:261–297, pls. 31–33. HALL, J. 1858. Palaeontology of Iowa, p. 473–724, 29 pls. In J. Hall and J. D. Whitney (eds.), Report of the Geological Survey of the State of Iowa: Embracing the Results of Investigations Made During Portions of the Years 1855–57. Volume 1 (Pt. 2). C. van Benthuysen, Albany. HALL, J. 1859. Palaeontology of New York. Volume 3. Containing Descriptions and Figures of the Organic Remains of the Lower Helderberg Group and the Oriskany Sandstone. C. van Benthuysen, Albany, 532 p., 120 pls. HALL, J., AND J. M. CLARKE. 1892. An introduction to the study of the Brachiopoda, intended as a handbook for the use of students, Pt. I, New York State Geologist Annual Report, 11:128–300, pls. 1–22; New York State Museum Annual Report, 45:449–616. HARRIS, R. W., AND R. V. HOLLINGSWORTH. 1933. New Pennsylvanian conodonts from Oklahoma. American Journal of Science, 25:193–204, 1 pl. HASS, W. H. 1953. Conodonts of the Barnett Formation of Texas. U.S. Geological Survey Professional Paper, 243-F:69–94, pls. 14–16. HENRY, T. W., AND M. GORDON JR. 1992. Middle and upper Chesterian brachiopod biostratigraphy, eastern Appalachians, Virginia and West Virginia. In P. K. Sutherland and W. I. Manger (eds.), Recent advances in Middle Carboniferous biostratigraphy—a symposium. Oklahoma Geological Survey Circular, 94:1–21. HOARE, R. D. 1961. Desmoinesian Brachiopoda and Mollusca from southwest Missouri. Missouri University Studies, 36, 262 p., 23 pls. IGO, H., AND T. KOIKE. 1964. Carboniferous conodonts from the Omi Limestone, Niigata Prefecture, central Japan (Studies of Asian conodonts, Pt. 1). Paleontological Society of Japan, Transactions and Proceedings, n.s., 53:179–193, pls. 27, 28. ISAACSON, P. E., AND J. T. DUTRO JR. 1999. Lower Carboniferous brachiopods from Sierra de Almeida, northern Chile. Journal of Paleontology, 73(4):625–633. IVANOVA, E. A. 1972. Osnovnye zakonomernosti evoliutsii spiriferid (Brachiopoda). Paleontologicheskii Zhurnal, 1972(3):28–42. (In Russian) KATZER, F. 1903. Grundzu¨ge der Geologie des uteren Amazonasgebietes (des Staates para in Brasilien). Leipzig, 296 p., 15 pls. KATZER, F. 1933. Geologia do Estado do Para´ (Brasil). Museu Paraense Emilio Goeldi Boletim, 9:1–288, 16 pls. KING, W. 1846. Remarks on certain genera belonging to the class Palliobranchiata. Annals and Magazine of Natural History, series 1, 18: 26–42, 83–94. KING, W. 1850. A monograph of the Permian fossils of England. Palaeontographical Society Monograph, 3(1), 258 p., 28 pls. KOZLOWSKI, R. 1914. Les brachipodes du carbonifere supe´rieur de Bolivie. Annales de Paleontology, 9, 100 p. LANE, H. R., AND J. J. STRAKA. 1974. Late Mississippian and Early Pennsylvanian conodonts, Arkansas and Oklahoma. Geological Society of America Special Paper, 152:1–144. LANE, H. R., P. L. BRENCKLE, J. F. BAESEMENN, AND B. C. RICHARDS. 1999. IUGS Carboniferous in the middle of the Carboniferous: Arrow Canyon, Nevada, USA. Episodes 22(4):272–283. LANE, N. G. 1963. A silicified Morrowan brachiopod faunule from the Bird Spring Formation, southern Nevada. Journal of Paleontology, 37(2):379–392, 3 pls. LANGENHEIM JR., R. L. 1991. Paleogeographic significance of Late Carboniferous Amazonian and Arctic brachiopods on the southwestern

continental margin of North America, p. 341–345. In D. I. MacKinnon, D. E. Lee, and J. D. Campbell (eds.), Brachiopods Through Time. Dunedin, New Zealand. LEMOS, V. S., AND R. A. MEDEIROS. 1989. Transgresso˜es e regresso˜es cıćlicas e correˆncia de conodontes no Morrowano e Atokano da Bacia do Amazonas. In Congresso Brasileiro de Paleontologia, 11 Curitiba, 1989. Sociedade Brasileira de Paleontologia, Curitiba, Anais, 2:961– 969. LEMOS, V. S., AND R. A. MEDEIROS. 1996. Conodontes do Carbonı´fero Inferior da Bacia do Amazonas. B. Geoc. PETROBRAS, Rio do Janeiro, 10(1/4):55–60. LE´VEILLE´, C. 1835. Apercu geologique de quelques localites tres riches en coquilles sur les frontieres de France et de Belgique. Societe geologique de France, Memories, series 1, (2):29–40, 2 pls. (In French) LITVINOVICH, N. V., G. G. AKSENOVA, AND T. P. RAZINA. 1969. Stratigrafiia i litologiia otlozhenii nizhnego karbona zapadnoi chasti Tsentral’nogo Kazakhstana. Nedra, Moscow, 448 p., 72 pls. (In Russian) MARCOU, J. 1858. Geology of North America, with two reports on the prairies of Arkansas and Texas, the Rocky Mountains of New Mexico, and the Sierra Nevada of California, originally made for the United States Government. Zurich, 144 p. MARTIN, W. 1793 (1794?). Figures and descriptions of Petrifactions collected in Derbyshire, to which are added a systematical list of the minerals which have been found constituting the substance of extraneous fossils in that county, and a brief introduction to the knowledge of Petrifactions in general. Lyon and Atkinson, London, 29 pls. MARTIŃEZ CHACOŃ, M. L. 1982. El geńero Kozlowskia (Productidina, Brachiopoda), en el Carbonı´fero de la Cordillera Canta´brica (NO de Espanã). Trabajos de Geologıá, Universidad de Oviedo, 12:73–82. MAXWELL, W. G. H. 1964. The Geology of the Yarrol Region, Pt. I, Biostratigraphy. University of Queensland, Department of Geology, Papers, 5(9):1–79, 14 pls. MCCHESNEY, J. H. 1860. Descriptions of new species of fossils from the Palaeozoic rocks of the western States. Chicago Academy of Sciences Transactions, 1(1):1–87, pls. 1–9. MCCOY, F. 1844. A synopsis of the characters of the Carboniferous limestone fossils of Ireland. London, 207 p., 29 pls. MCKELLAR, R. G. 1965. An Upper Carboniferous brachiopod fauna from the Monto district, Queensland. Geological Survey of Queensland Publications, 328:1–15, 5 pls. MEEK, F. B. 1877. Palaeontology. United States Geological Exploration of the 40th Parallel by Clarence King, 4(1):1–197, pls. 1–17. MENDES, J. C. 1952. Faunula Permo–Carbonı´fera marinha de Capivari. Sao Paulo Universidade, Faculdade de Filosofia, Ciencias e Letras, Boletim (Geologia 7), 134(7):1–20, 1 pl. (In Portuguese) MENDES, J. C. 1956a. Orhtotetacea e Dalmanellacea do Carbonı´fero Superior do Rio Tapajo´s (se´rie Itaituba). Sociedade Brasileira de Geologia, Boletim, 5(1):11–38, 4 pls. (In Portuguese) MENDES, J. C. 1956b. Spirifercea Carbonifero do Rio Tapajo´s (se´rie Itaituba), Estado do Para´, Brasil. Sao Paulo Universidade, Faculdade de Filosofia, Ciencias e Letras, Boletim (Geologia 13), 193:23–81, 5 pls. (In Portuguese) MENDES, J. C. 1957. Rhynchonellacea, Rostropiracea e Terebratulacea do Carbonı´fero do Rio Tapajo´s, Brasil. Sociedae Brasileira de Geologia, Boletim, 6(1):15–35, 3 pls. (In Portuguese) MENDES, J. C. 1959. Chonetacea e Productacea Carbonı´feros da Amazoˆnia. Sao Paulo Universidade, Faculdade de Filosofia, Ciencias e Letras, Boletim (Geologia 17), 236:1–83, 7 pls. (In Portuguese) MENDES, J. C. 1961. Notas Suplementares sobre os Braquio´podes Carbonı´feros da Amazońia. Sociedae Brasileira de Geologia, Boletim, 10(1):5–24. (In Portuguese) MENDES, J. C. 1972. Braquiopodes e Moluscos neocarbonı´feros da Amazoˆnia (Formacaõ Itaituba). Academia Brasileira de Ciencias Anais, 44: 243–246. (In Portuguese) MILORADOVICH, B. V. 1945. Nekotorye dannye po morfologii rakovin produktid. (Some data on the morphology of the shells of Productidae.) Akademiia Nauk SSSR, Izvestiia, Seriia Biologicheskaia, 4:485–500. (In Russian) MOORE, R. C. 1952. Brachiopods, p. 197–267. In R. C. Moore, C. G. Lalicker, and A. G. Fischer (eds.), Invertebrate Fossils. McGraw-Hill, New York.

CHEN ET AL.—UPPERMOST MISSISSIPPIAN BRACHIOPODS OF THE AMAZON BASIN MORROW, J. R., AND G. D. WEBSTER. 1992. New stratigraphic, petrographic, and biostratigraphic data on the proposed Mississippian–Pennsylvanian boundary stratotype, Granite Mountain, west-central Utah. Oklahoma Geological Survey Circular, 94:55–67. MUIR-WOOD, H. M., AND G. A. COOPER. 1960. Morphology, classification and life habits of the Productoidea (Brachiopoda). Memoir of the Geological Society of America, 81, 447 p., 135 pls. NORTH, F. J. 1920. On Syringothyris Winchell, and certain Carboniferous Brachiopoda referred to Spiriferina d’ Orbigny. Geological Society of London Quarterly Journal, 76(2):162–227, pls. 11–13. NORWOOD, J. G., AND H. PRATTEN. 1855. Notice of Producti found in the western States and Territories, with descriptions of twelve new species. Academy of Natural Sciences of Philadelphia Journal, series 2, 3:5–22. OEHLERT, D. P. 1890. Note sur diffe´rents groupes e´tablis dans le genre Orthis et en particulier sur Rhipidomella Oehlert (5 Rhipidomys Oehlert, olim). Journal de Conchyliologie, 30:366–374. (In French) PETRI, S. 1952. Fusulinidae do Carbonı´fero do rio Tapajo´s, Estado do Par. Sociedae Brasileira de Geologia Boletim, 1(1):30–45, 2 pls. (In Portuguese) PETRI, S. 1956. Foraminı´feros do Carbonı´fero do Amazońia. Sociedae Brasileira de Geologia Boletim, 5(1):17–30. (In Portuguese) PINTO, I. D. 1977. Corais Carbonı´feros da Bacia Amazoˆnica. Pesquisas, Universidade Federal do Rio Grande do Sul, 8:59–131. (In Portuguese) POLETAEV, S. V. 1997. A revision of the genus Neospirifer Fredericks, 1924 and a description of Lutuginia, a new genus in the subfamily Neospiriferinae. Paleontological Journal, 31(3):302–311. POLETAEV, S. V. 1999. Revision of Palaeochoristes and description of a new Early Carboniferous spiriferid genus Phragmobrachythyris. Paleontological Journal, 33(3):254–262. REED, F. R. C. 1933. Some Upper Carboniferous brachiopods from Brazil. Annals and Magazine of Natural History, series 10, 11(65):519–537, 1 pl. ROBERTS, J. 1964. Lower Carboniferous faunas from Wiragulla and Dungog, New South Wales. Royal Society of New South Wales Journal and Proceedings, 97(6A):193–215, 5 pls. ROBERTS, J. 1976. Carboniferous Chonetacean and Productacean brachiopods from eastern Australia. Palaeontology, 19(1):17–77, pls. 3– 13. ROCHA CAMPOS, A. C. 1985. Paraguay and Uruguay, South America, p. 216–226. In C. Martinez Diaz, R. H. Wagner, C. F. Winkler Prins, and L. F. Granados (eds.), The Carboniferous of the World. II. Australia, Indian Subcontinent, South Africa, South America and North Africa. IUGS Publication, n. 20, 447 p., Instituto Geologico y Minero de Espana. ROCHA CAMPOS, A. C., AND S. ARCHANGELSKY. 1985. Biostratigraphy and correlations, South America, p. 226–245, 13 pls. In C. Martinez Diaz, R. H. Wagner, C. F. Winkler Prins, and L. F. Granados (eds.), The Carboniferous of the World. II. Australia, Indian Subcontinent, South Africa, South America and North Africa. IUGS Publication, n. 20, 447 p., Instituto Geologico y Minero de Espana. ROUNDY, P. V. 1926. The micro-fauna in Mississppian formations of San Saba County, Texas. U.S. Geological Survey Professional Paper, 146: 1–63, pls. 1–4. ROWLEY, R. R. 1900. Descriptions of new species of fossils from the Devonian and sub-Carboniferous rocks of Missouri. American Geologist, 25:261–273. SARTENAER, P., AND G. PLODOWSKI. 2002. Description of Mesochorispira konincki (Dewalque, 1895), a spiriferid brachiopod from the Belgian Upper Tournaisian. Bulletin de L’Institut Royal des Sciences Naturelles de Belgique, Sciences de la Terre, 72:69–84. SARYCHEVA, T. G., AND A. V. N. SOKOLSKAYA. 1959. O klassifikatsin lozhnoporistykh brakhiopod. Akademiia Nauk SSSR Doklady, 125: 181–184. (In Russian) SARYCHEVA, T. G., A. V. N. SOKOLSKAYA, G. A BESNOSOVA, AND S. V. MAXIMOVA. 1963. Brakhiopody i paleogeografiia karbona Kuznetskoi kotloviny. Akademiia Nauk SSSR, Paleontologicheskii Institut Trudy, 95, 547 p., 64 pls. (In Russian) SCHUCHERT, C. 1913. Brachiopoda, p. 355–420. In K. A. Von Zittel (ed.), Paleontology (second edition). Volume 1. London.

925

SCHUCHERT, C. 1929. Classification of brachiopod genera, fossil and recent, p. 10–25. In C. Schuchert and C. M. LeVene (eds.), Brachiopoda (generum et genotyporum index et bibliographia), Fossilium Catalogus I: Animalia, Pars 42. W. Junk, Berlin. SCHUCHERT, C., AND C. M. COOPER. 1932. Brachiopod genera of the suborders Orthoidea and Pentameroidea. Peabody Museum of Natural History Memoirs, 4(1):1–270, 29 pls. SOWERBY, J. 1822. The Mineral Conchology of Great Britain, 4:17–114, pls. 319–383. London. SOWERBY, J., AND J. DE C. SOWERBY. 1818–21. The Mineral Conchology of Great Britain, 3:1–184, pls. 204–306. London. SNIDER, L. C. 1915. Paleontology of the Chester Group in Oklahoma. Oklahoma Geological Survey Bulletin, 24(2):67–122, pls. 3–7. SPENCER, R. S. 1967. Pennsylvanian Spiriferacea and Spiriferinacea of Kansas. University of Kansas Paleontological Contributions Paper, 14: 1–35. STAUFFER, C. R., AND H. J. PLUMMER. 1932. Texas Pennsylvanian conodonts and their stratigraphic relations. Texas University, Bureau of Economy Geology Publication, 3201:13–50, pls. 1–4. STEHLI, F. G. 1954. Lower Leonardian Brachiopoda of the Sierra Diablo. American Museum of Natural History Bulletin, 105:257–358, pls. 17– 27. STURGEON, M. T., AND R. D. HOARE. 1968. Pennsylvanian brachiopods of Ohio. Geological Survey of Ohio Bulletin, 63:1–95, 22 pls. SUTHERLAND, P. K., AND F. H. HARLOW. 1967. Late Pennsylvanian Brachiopods from north-central New Mexico. Journal of Paleontology, 41(5):1065–1089, 6 pls. SUTHERLAND, P. K., AND F. H. HARLOW. 1973. Pennsylvanian brachiopods and biostratigraphy in southern Sangre de Cristo Mountains, New Mexico. New Mexico Bureau of Mines and Mineral Resources Memoir, 27, 173 p., 18 pls. TENGAN, C., S. SHIMABUKURO, AND A. C. ROCHA CAMPOS. 1976. Conodontes carbonı´ferous do Poco FBst-1-AM, Bacia do Amazonans, Brasil, Anais XXIX Congress Brasileira Geologia, Belo Horizontes, resumos. Sociedade Brasileira de Geologia, 365. THOMPSON, M. L. 1942. New genera of Pennsylvanian fusulinids. American Journal of Sciences, 240:403–420. UENO, K., AND T. NAKAZAWA. 1993. Carboniferous foraminifers from the lowermost part of the Omi Limestone Group, Niigata Prefecture, central Japan. Science Report, Institute of Geosciences, University Tsukuba, series B, 14:1–51. WAAGEN, W. H. 1882–1885. Salt Range fossils: Productus Limestone fossils, Pt. 4(2–4): Brachiopoda. Memoirs of the Geological Survey of India, Palaeontologica Indica, series 13,1:329–770, pls. 25–86. WANG, G. P., Q. Z. LIU, Y. G. JIN, S. Z. HU, W. P. LIANG, AND Z. T. LIAO. 1982. Brachiopoda, p. 186–256, pls. 74–102. In Palaeontological Atlas of East China (Late Paleozoic). Volume 2. Geological Publishing House, Beijing, 495 p., 157 pls. (In Chinese) WATERHOUSE, J. B. 1975. New Permian and Triassic brachiopod taxa. University of Queensland, Department of Geology, Papers, 7(1):1–23, 2 pls. WEBSTER, G. D., AND E. GROESSENS. 1990. Conodont subdivision of the Lower Carboniferous. Courier Forschungsinstitut Senckenberg, 130: 34–40. WELLER, S. 1914. The Mississippian Brachiopoda of the Mississippi Valley Basin. Illinois State Geological Survey Monograph, 1:1–508, 83 pls. WHITE, C. A. 1862. Description of new species of fossils from the Devonian and Carboniferous rocks of the Mississippi Valley. Boston Society of Natural History Proceedings, 9:8–33. WILLIAMS, A. 1953. The classification of the strophomenoid brachiopods. Washington Academy of Sciences Journal, 43(1):1–13. WILLIAMS, A., AND C. H. C. BRUNTON. 1997. Morphological and anatomical terms applied to brachiopods, p. 421–440. In A. Williams, C. H. C. Brunton, and S. J. Carlson (eds.), Treatise on Invertebrate Paleontology, Pt. H, Brachiopoda (revised). Volume 1. Introduction. The Geological Society of America and The University of Kansas, Lawrence. WILLIAMS, A., AND C. H. C. BRUNTON. 2000. Orthotetidina, p. 644–681. In A. Williams, C. H. C. Brunton, and S. J. Carlson (eds.), Treatise on Invertebrate Paleontology, Pt. H, Brachiopoda (revised). Volume 3. The

926


Geological Society of America and The University of Kansas, Lawrence. WILLIAMS, A., AND D. A. HARPER. 2000. Orthida, p. 714–782. In A. Williams, C. H. C. Brunton, and S. J. Carlson (eds.), Treatise on Invertebrate Paleontology, Pt. H, Brachiopoda (revised). Volume 3. The Geological Society of America and The University of Kansas, Lawrence. WILLIAMS, A., S. J. CARLSON, C. H. C. BRUNTON, L. E. HOLMER, AND L. POPOV. 1996. A supra-ordinal classification of the Brachiopoda.

Philosophical Transactions of Royal Society of London, series B, 351: 1171–1193. 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, 9 pls. ZHAO, Z. X., J. X. HAN, AND Z. J. WANG. 1984. The Carboniferous Strata and Its Fauna from Southwestern Margin of Tarim Basin in Xinjiang. Geological Publishing House, Beijing, 87 p., 35 pls. (In Chinese) ACCEPTED 16 SEPTEMBER 2004