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The Devonian trilobites of Brazil: A summary
Accepted Manuscript The Devonian trilobites of Brazil: A summary Maria da Gloria Pires de Carvalho, Luiza Corral Martins de Oliveira Ponciano PII:
S0895-9811(15)30075-4
DOI:
10.1016/j.jsames.2015.10.010
Reference:
SAMES 1476
To appear in:
Journal of South American Earth Sciences
Received Date: 18 July 2014 Revised Date:
12 October 2015
Accepted Date: 17 October 2015
Please cite this article as: Carvalho, M.d.G.P.d., Ponciano, L.C.M.d.O., The Devonian trilobites of Brazil: A summary, Journal of South American Earth Sciences (2015), doi: 10.1016/j.jsames.2015.10.010. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
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THE DEVONIAN TRILOBITES OF BRAZIL: A SUMMARY
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MARIA DA GLORIA PIRES DE CARVALHO American Museum of Natural History, Division of Paleontology, Research Associate. Central Park West and 79th Street, New York, N.Y. 100-24-5192, USA. [email protected]
ABSTRACT
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LUIZA CORRAL MARTINS DE OLIVEIRA PONCIANO Universidade Federal do Estado do Rio de Janeiro, Departamento de Ciências Naturais, Laboratório de Tafonomia e Paleoecologia Aplicadas – LABTAPHO, Professor. Av. Pasteur, 458, sala 504, 22290-255, Rio de Janeiro, RJ, Brazil. [email protected]
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Devonian trilobites are found in three major Paleozoic intracratonic basins of Brazil (Amazonas, Parnaíba, and Paraná). The trilobites represent the families Homalonotidae, Dalmanitidae, and Calmoniidae. The distribution of these taxa in the Brazilian territory is summarized here because of their remarkable scientific and historical importance, and a revised taxonomy and
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lithostratigraphy of the Devonian (Pragian - Famennian) trilobites from Brazil is presented, based on new research and recent literature. Homalonotids and dalmanitids are relatively cosmopolitan, whereas calmoniids are more endemic and seem to have been restricted to marine cold-waters of
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the southern hemisphere (Malvinokaffric Realm). Although the trilobites within the Brazilian intracratonic basins are approximately contemporaneous, they show various patterns of endemism
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and biogeographical affinities with other Malvinokaffric areas such as Bolivia, South Africa, and the Falkland (Malvinas) Islands. At family level, therefore, trilobite diversity from Brazil is comparatively low, which may indicate biogeographical filtering related to the distance and/or remoteness of the Brazilian basins from more open oceanic waters. Key words: Trilobites, Devonian, Brazil.
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1. INTRODUCTION Three large Brazilian intracratonic basins are discussed in this paper: the Amazonas Basin,
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located in the North; the Parnaíba Basin of the North/Northeast; and the Paraná Basin in the Southeast/South (Fig.1). A fourth major Paleozoic basin (Solimões, in northwestern Brazil) is still poorly studied in terms of its macrofossils and therefore not included in the scope of this
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paper. Each of the basins contains a considerable thickness of Middle Paleozoic clastic
sediments, partly exposed in extensive outcrop belts along their margins. Marine Devonian
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sequences are well represented in these basins and provide evidence of widespread marine transgressions across Brazil, especially during the Pragian - Frasnian. The marine invertebrate fossils discovered in these sequences include trilobites, which are restricted to the families Homalonotidae, Dalmanitidae, and Calmoniidae. The Calmoniidae is considered to be endemic
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to the marine cold-water, southern hemisphere Malvinokaffric Realm. The other two families are more cosmopolitan, whereas their Brazilian representatives seem to be somewhat endemic at genus or species level. The distribution of these trilobites among the Brazilian intracratonic
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basins is shown in Tables 1-4.
The first evidence about Devonian marine invertebrate fossils from Brazil was found during the
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pioneering Morgan expeditions in 1870-71, led by Charles Frederick Hartt, in the Amazonas Basin, and during the Imperial Geological Commission expeditions of 1876-77, directed by Orville Adelbert Derby. A series of publications dealing mainly with the shelly fossils collected on these expeditions were published between 1871 and 1913 (see Melo, 1988 for references). Besides Katzer (1903, 1933) and Melo (1988), only a few additional papers have been published on the Devonian invertebrate fossils from this area during the last years (see Ponciano, 2011 for
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references). Devonian strata of the Parnaíba Basin were first discovered by Small (1914) near to the
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Pimenteira village (nowadays the city of Pimenteiras), in the State of Piauí, but were recognized as Devonian only by Caster (1948). Wilhelm Kegel subsequently collected many marine
invertebrate fossils from this basin in the mid-C20th (Kegel, 1953). Subsequently, Melo (1988),
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Carvalho et al. (1997), Fonseca (2004), Ponciano and Della Favera (2009), Fonseca and Ponciano (2011), Fernandes et al. (2012), Ponciano et al. (2010, 2012a, b), and other few papers have
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accorded attention to Devonian macrofossil assemblages from the Parnaíba Basin. The Devonian of the Paraná Basin has been investigated in several stratigraphic, sedimentological, and paleontological studies. Derby (1878b) first recognized Devonian fossils in this basin. Clarke’s (1913) monograph is a classic publication and still represents one of the most
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important contributions on the Devonian marine invertebrates of the Paraná Basin. Clarke (1913) recognized the distinction between austral and boreal faunas, and also associated the Brazilian fauna with the Falkland (Malvinas) Islands and South Africa, uniting them as an “Austral Fauna”.
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Later, Richter and Richter (1942) erected the terminology “Malvinokaffric Province” for the endemic Devonian trilobites recovered from various localities in the Southern Hemisphere
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(Falkland/Malvinas Islands, South Africa, Brazil - State of Paraná, Ponta Grossa Region). This later became known as the “Malvinokaffric Realm” (Eldredge and Ormiston, 1979). More recent studies have found evidence for an early to early late Emsian decline of the Malvinokaffric fauna in the Paraná Basin (Bosetti et al., 2012), as well as an early Givetian “Lilliput Effect” associated with the collapse of the Malvinokaffric shelly fauna (Bosetti et al., 2010) supposedly with up to 90% size reduction of surviving species, although the extent to which the trilobites were affected
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is unclear. 2. MATERIAL AND METHODS
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The material studied here is represented by internal and external molds, mostly disarticulated; there are some almost complete specimens, but generally they are poorly preserved. Specimens preserved inside concretions (especially those from the Parnaíba Basin) have better preservation.
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Open nomenclature has been retained for certain taxa, due to the incompleteness and poor
preservation of the specimens on which they are based, especially those from the Amazonas
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Basin. Although some specimens were coated with magnesium oxide prior to photography, in many cases this was not possible for conservational reasons.
The trilobites illustrated or cited in this paper are housed in the following institutions: AMNH, American Museum of Natural History, New York.
Facility, Cincinnati.
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CMC IP, Cincinnati Museum Center Invertebrate Paleontology, Geier Collections and Research
DCN/UNIRIO, Departamento de Ciências Naturais da Universidade Federal do Estado do Rio de
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Janeiro, Rio de Janeiro.
DGM, DNPM, Departamento Nacional de Produção Mineral, Rio de Janeiro.
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DZP/UNESP, Departamento de Zoologia da Universidade Estadual de São Paulo, Botucatu. MCT, Museu de Ciências da Terra, Setor de Paleontologia, DNPM, Rio de Janeiro. MNHN, Paris- R, Museum National d’Histoire Naturelle, Paris. NYSM, New York State Museum, Albany. UC, University of Cincinnati, Cincinnati. IGEO/UFRJ, Instituto de Geociências da Universidade Federal do Rio de Janeiro, Rio de Janeiro.
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3. AMAZONAS BASIN
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MN/UFRJ, Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro.
According to Cunha et al. (2007) the Devonian lithostratigraphic units of the Amazonas Basin comprise (in ascending order), the upper part of the Trombetas Group (Manacapuru and Jatapu
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formations), the Urupadi Group (Maecuru and Ererê formations) and the main part of the Curuá Group (Barreirinha and Curiri formations). Trilobites are documented only from the Maecuru
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and Ererê formations. 3.1 Maecuru Formation
The Maecuru Formation (sensu Derby, 1878) consists mainly of a sequence of fluvio-deltaic to neritic sandstones, with few siltstone interbeds. On the northern flank of the Amazonas Basin this
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unit is developed as shallow marine tempestites. These beds contain the highest diversity of shelly faunas in any Brazilian Devonian assemblages (Melo, 1988). The type locality, according to Lange (1967, p. 237), is a poorly delimited outcrop section exposed along the Maecuru River
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bedrock, upstream of the Teuapixuna Rapids (where the overlying Ererê Formation is exposed) and downstream of the Pancada Grande Waterfall (outcrop site of the underlying Trombetas
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Group).
Trilobites (Fig.2) occur in the upper part of the former Lontra Member sensu Lange (1967) [= Maecuru Formation sensu stricto after Derby (1878) and Cunha et al. (2007)], which represents a shallow marine facies of Middle Devonian (latest Emsian-early Eifelian) age (Grahn and Melo, 2004). Homalonotids are represented by a single taxon, Digonus derbyi (Clarke, 1890). Dalmanitids include Amazonaspis maecurua (Clarke, 1890), ?Dalmanites infractus Clarke, 1890,
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and Dalmanitidae gen. and sp. indet. Calmoniids are considerably more diverse, with ?Acastoides menurus (Clarke, 1890), Malvinella australis (Clarke, 1890), ?Malvinella tumiloba (Clarke,
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1890), “Palpebrops” goeldi (Katzer, 1903), Phacopina braziliensis (Clarke, 1890), Vogesina gemellus (Clarke, 1890), plus several gen. indet. species, including galea Clarke, 1890, scirpeus Clarke, 1890, macropyge Clarke, 1890, and acanthurus Clarke, 1890. Another trilobite is of
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uncertain affinity at family level, “Phacops” pullinus Clarke, 1890.
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3.2 Ererê Formation
Derby (1878) introduced the term Ererê Formation for a section of siltstones with interbedded shale and fine-grained sandstones, above the Maecuru Formation and below the Curuá Group. The age of this formation is latest Eifelian to early Givetian (Grahn and Melo, 2004). The type
region, State of Pará.
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area of the Ererê Formation occupies the central part of the Ererê Dome in the Monte Alegre
Marine invertebrate macrofossils in this formation are impoverished, especially the trilobites
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(Fig.2), but they are distributed in a wide variety of lithologies (sandstones to shales). They mostly occur in the lower level of this unit, which is according to microfossil evidence late
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Eifelian (Grahn and Melo, 2004). Trilobites are rare, but are represented by a homalonotid, Burmeisteria oiara (Hartt and Rathbun, 1875) and a calmoniid, Eldredgeia paituna (Hartt and Rathbun, 1875).
4. PARNAÍBA BASIN The Devonian succession of the Parnaíba Basin is represented, in ascending order, by the upper
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Jaicós (Early Devonian pro parte), Itaim, Pimenteira, Cabeças, and lowermost Longá formations (Vaz et al., 2007). During the Devonian, this intracratonic basin had a more restricted access to
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the open ocean than adjacent contemporary basins; hence, its sediments were deposited under shallow marine conditions (Grahn et al., 2006).
No trilobite body fossils have been recovered from the Jaicós, Itaim, and Cabeças formations, but
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trilobite trackways (possible from homalonotids) have been found in Itaim Formation (Santos and Carvalho, 2009). Trilobites body fossils occur mainly in the Pimenteira Formation (Picos and
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Passagem members), and are scarcer in the Longá Formation.
4.1 Picos Member of the Pimenteira Formation
The Picos Member of the Pimenteira Formation (Kegel, 1953) consists of interlayered fine-
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grained hummocky cross-stratified sandstones and siltstones/shales. The base of the Pimenteira Formation marks the first widespread Devonian transgression across the Parnaíba Basin and conformably overlies the Itaim Formation, which has a much sandier character and is probably of
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fluvial to deltaic and shallow marine origin (Melo, 1988). Grahn et al. (2008) assigned a late Eifelian-early Givetian age span to the Pimenteira Formation
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in the eastern outcrop belt of the Parnaíba Basin, based on chitinozoans obtained from shallow drill cores. In the basin’s subsurface and southwestern outcrop belt, however, the upper Pimenteira Formation may be as young as Frasnian or early Famennian (Loboziak et al., 2000; Grahn et al., 2006). Breuer and Grahn (2011a, b) retrieved late early Givetian palynofloras from Picos Member pelites laterally equivalent to the Passagem Member sandstones in the eastern Parnaíba Basin.
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Trilobites from the basal part of the Pimenteira Formation (Picos Member; Fig.3) include a homalonotid, Burmeisteria notica (Clarke, 1913), plus the calmoniids Eldredgeia cf. E. venusta
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(Wolfart, 1968), Metacryphaeus kegeli Carvalho, Edgecombe and Lieberman (1997), and Metacryphaeus tuberculatus (Kozlowski, 1923); figured here for the first time from the
Pimenteira Formation (Fig.3.2). Burmeisteria notica is very abundant and well preserved in
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ironstone concretions, especially near the city of Picos (State of Piauí). More rarely,
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Metacryphaeus tuberculatus and other fossil invertebrates also are found in these concretions.
4.2 Passagem Member of the Pimenteira Formation
The upper part of the Pimenteira Formation comprises moderately fossiliferous, highly micaceous fine-grained sandstones with asymptotic and hummocky cross-stratification, and subordinate
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siltstone interbeds constituting Kegel’s (1953) Passagem Member (attributed by him to the lowest part of the Cabeças Formation). This Member is known only in the eastern outcrop belt of the basin (State of Piauí), where it is currently dated as early to early middle Givetian (Grahn et al.,
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2008; Breuer and Grahn, 2011a, b).
The regional unconformity separating the Passagem sandstones from those of the Cabeças
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Formation sensu stricto (latest Famennian age; Grahn et al., 2006) is observed at Pedro Segundo Ridge and the environs of São José do Piauí village, situated respectively in the northeastern and eastern parts of the State of Piauí (Ponciano et al., 2012a). According to recent interpretations (Ponciano and Della Favera, 2009; Ponciano et al., 2012a), the Passagem Member comprises fine-grained distal mouth-bar deposits, interbedded with delta-front, hummocky cross-stratified, graded sandstone lobes. This entire sequence probably represents a
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flood-dominated fluvio-deltaic system entering shallow marine settings. According to this scenario, turbulent flows during episodic flood events sometimes transcended the limits of the
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littoral zone, thereby reworking littoral and benthic shelf organisms into the resulting shallow marine sandstone bodies.
Trilobites from the Passagem Member (Fig.3) include a calmoniid, Metacryphaeus meloi
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Carvalho, Edgecombe and Lieberman (1997), and a homalonotid, Burmeisteria notica (Clarke,
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1913).
4.3 Longá Formation
The Longá paleoenvironment is generally regarded as a transgressive, storm-dominated, shallow sea (Vaz et al., 2007). The lowest part of this formation is composed of shales with interbedded
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conglomeratic to fine-grained sandstones, and contains an impoverished benthic marine invertebrate assemblage of latest Devonian age (uppermost Famennian), with extremely rare, small and poorly preserved trilobites and brachiopods, together with more abundant mollusks and
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trace fossils. The rest of the formation is of Tournaisian age (Playford et al., 2012), but it has not yielded any shelly fossils to date.
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A small trilobite, only known from a single cephalon and pygidium, was collected by Wilhem Kegel from poorly dated basal Longá Formation outcrops at Barreiras Farm (Valença do Piauí area, State of Piauí, eastern Parnaíba Basin), and referred to “Asteropyge” (Kegel, 1953). It may represent a new form of Metacryphaeus, although the presence of a calmoniid within such a young stratigraphic level (Strunian or perhaps even Tournaisian) is exceptional. It is possible that the fossil may have been reworked from Middle Devonian or older strata (Melo, 1988).
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5. PARANÁ BASIN
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During the Devonian, the Paraná Basin included two sub-basins; the Apucarana Sub-basin in the south, and the Alto Garças Sub-basin in the north. These sub-basins had a different geological evolution and are partially separated from each other by the Três Lagoas and Campo Grande
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structural highs (see Fig.1). The Devonian stratigraphic nomenclature for the Paraná Basin adopted here follows Melo (1988) and Grahn et al. (2010, 2013).
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Within the Apucarana Sub-basin, Melo (1988) and Grahn et al. (2013) regarded Ponta Grossa and São Domingos as separate formations. The Tibaji Member is considered to be a member of the latter. Evans (1894) introduced the term Chapada Group for rocks in the northwestern part of the Paraná Basin (Alto Garças Sub-basin), and the Chapada dos Guimarães is regarded as the type
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area. The lithology differs from stratigraphically equivalent units in the Apucarana Sub-basin mainly in having a higher silt and sand content (the marine environment is shallower in the Alto Garças Sub-basin; Grahn et al., 2013). The Chapada Group still does not have formal
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lithostratigraphic designations. Instead, it is informally divided into Chapada units 1, 2, 3, and 4,
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in ascending order (Melo, 1988).
5.1 Chapada Group
Sediments of the Chapada Group occur in the northern part of the Paraná Basin, within the states of Mato Grosso, Mato Grosso do Sul and Goiás. They also encroach as a thin Paleozoic cover onto the southeastern rim of the adjacent, chiefly Precambrian Parecis Basin to the north (central Mato Grosso State), where Lower Devonian fossils identical to those of Chapada dos Guimarães
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are known to occur (Carvalho and Edgecombe, 1991; Carvalho, 1997; Boucot et al., 2001). Chapada unit 1 (essentially sandy) is largely unfossiliferous (except for trace fossils and land
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plant remains of probable Lochkovian age, in its upper part), and is regarded as an equivalent of the Furnas Formation of the Apucarana Sub-basin, to the south, which has a similar lithology and paucity of fossils. Transitional beds between Chapada units 2 and 3 contain a similar fossil
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assemblage to the upper Ponta Grossa Formation and Tibaji Member of the São Domingos
Formation (Melo, 1988), thus suggesting a late Emsian age for these beds. The lithologies are
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reddish sandstones with conglomeratic levels. The sandstones represent a deltaic environment on a shallow platform, and also contain shale clasts and sporadic channels. The trilobites from Mato Grosso (Figs. 4-6) include the homalonotid Burmeisterella braziliensis Carvalho, 2006, plus the calmoniids Kozlowskiaspis subseciva (Clarke, 1913), ?Calmonia triacantha Carvalho and
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Edgecombe, 1991, and Metacryphaeus australis (Clarke, 1913). Trilobites from Goiás include the calmoniid Metacryphaeus aff. M. australis Carvalho, Melo and Quadros, 1987 and
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Kozlowskiaspis subseciva (Clarke, 1913).
5.2 Ponta Grossa Formation
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The lower part of the Ponta Grossa Formation consists of transgressive, highly fossiliferous gray shales with subordinated siltstones and sandstones. Northeast and southeast of Ponta Grossa city, as well as virtually all over the Apucarana Sub-basin, the formation unconformably overlies coarser, poorer-sorted, and kaolinitic sandstones of the earliest Devonian (Lochkovian) Furnas Formation. These basal Ponta Grossa lithologies are followed by sandy shales with limestones nodules or arenaceous clays. The upper Ponta Grossa section is developed as hard, black, pyritic
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shales. The Ponta Grossa Formation has produced a rich and diverse assemblage of marine
acritarchs, and chitinozoans (Grahn et al., 2013).
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invertebrates including trilobites (Figs. 4-6), as well as latest Pragian-early Emsian miospores,
Two homalonotids are recorded from this unit, Burmeisteria notica (Clarke, 1913) and B.
herscheli (Murchison, 1839) (Simões et al., 2009). There is also a dalmanitid, Dalmanitoides
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accola (Clarke, 1913). Calmoniid trilobites are more diverse, with several genera and numerous species, including Bainella paranaense Carvalho and Edgecombe, 2006, ?Calmonia gonzagana
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(Clarke, 1890), Calmonia michrischia Clarke, 1913, Calmonia signifer Clarke, 1913, Kozlowskiaspis subseciva (Clarke, 1913), Metacryphaeus australis (Clarke, 1913), Metacryphaeus rotundatus (Kozlowski, 1923) (Soares et al., 2008), Paracalmonia cuspidata (Clarke, 1913), Paracalmonia mendesi Popp, Coimbra and Hauch, 1996, Paracalmonia
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paranaensis Popp, Coimbra and Hauch, 1996, Paracalmonia pessula (Clarke, 1913), Paracalmonia salamunii Popp, Coimbra and Hauch, 1996, Paranacaste pontagrossensis Popp, 1989, Paraphacopina polygona Mori and Leme, 2013 (nomem nudum), Pennaia pauliana
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Clarke, 1913, and Pennaia lombardi (Kozlowski, 1913).
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5.3 Tibaji Member of the São Domingos Formation The São Domingos Formation overlies the Ponta Grossa Formation transgressively. The Tibaji Member has a restricted geographical extent, but locally constitutes the lowermost part of the formation. Over much of the Apucarana Sub-basin, the São Domingos Formation is overlain unconformably by the Pennsylvanian-Early Permian Itararé Group and locally (in the subsurface of Paraná State), by the so-called Ortigueira Diamictite, from the late Famennian (Loboziak et al.,
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1995; Milani et al., 2007). The São Domingos Formation consists predominantly of argillaceous shale. Poorly sorted conglomeratic sandstones occur in its basal part, succeeded by shale and
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micaceous siltstone rich in plant debris. The age of the Tibaji Member is early late Emsian, based on chitinozoans (Grahn et al., 2013).
Homalonotids and calmoniids occur in the Tibaji Member (Figs. 4-6), but no dalmanitids have
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been reported. Homalonotids include Burmeisteria notica (Clarke, 1913) and Burmesteria
herscheli (Murchison, 1839). Calmoniids are rarer than in the Ponta Grossa Formation and
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display somewhat lower diversity, but nevertheless include five taxa; Bainella paranaense Carvalho and Edgecombe, 2006, Kozlowskiaspis subseciva (Clarke, 1913), Metacryphaeus australis (Clarke, 1913), Metacryphaeus rotundatus (Kozlowski, 1923), and Tibagya parana (Clarke, 1913).
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6. BIOGEOGRAPHIC OBSERVATIONS Devonian trilobites from the Amazonas, Parnaíba, and Paraná basins of Brazil represent only three families: Homalonitidae, Dalmanitidae, and Calmoniidae. Although homalonotids and
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dalmanitids have a world-wide distribution, calmoniids seem to have been restricted to the coldwater Malvinokaffric Realm of the southern hemisphere. All three families are also well
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represented in Malvinokaffric assemblages from other areas, although those sometimes include members of additional families. For example, proetids occur in Bolivia, South Africa, Argentina, and perhaps in the Falklands (Malvinas) Islands (Eldredge and Ormiston, 1979; Cooper, 1982; Carvalho, 2006; Rustán et al, 2011), and aulacopleurids are known from Bolivia, Argentina and South Africa (Adrain & Edgecombe, 1996; Rustán & Vaccari, 2010). At family level, therefore, trilobite diversity from Brazil is comparatively low, which may indicate biogeographical filtering
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related to the distance and/or remoteness of the Brazilian basins from more open oceanic waters. 6.1 Homalonotids
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Homalonotids are widespread in many shallow-water Malvinokaffric communities, which reduce their value for paleobiogeographical analysis (Eldredge and Ormiston, 1979, p. 148). For
example, Digonus is recorded from Europe (Germany, Belgium, UK, and Czech Republic),
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United States, North and South Africa, and Antarctica, so the occurrence of Digonus derbyi (Clarke, 1890) in the Amazonas Basin is biogeographically uninformative. Similarly,
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Burmeisterella is known from Europe (England, Belgium, and Germany) as well as from the Paraná Basin (Alto Garças Sub-basin). On the other hand, the genus Burmeisteria is currently recognized only from the Devonian of the Malvinokaffric Realm. Burmeisteria oiara occurs only in the Amazonas Basin, B. notica is known from both the Parnaíba and Paraná basins, and B.
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herscheli (Murchison, 1839) is even more widespread, occurring in Brazil (Paraná Basin), the Falkland (Malvinas) Islands, and South Africa. 6.2 Dalmanitids
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Dalmanitids occur in the Lower and Middle Devonian of the Paraná and Amazonas basins, but included taxa differ considerably in their distribution. For example, Amazonaspis maecurua
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(Clarke, 1890) is known only from the Amazonas Basin, but Dalmanitoides Delo, 1935 occurs in several Malvinokaffric assemblages outside Brazil (Rustán & Vaccari, 2012). In Argentina, this genus is represented by D. boehmi (Knod, 1908) and D. drevermanni (Thomas, 1906), in Bolivia by D. scutata Branisa and Vanek, 1973, in Brazil (Paraná Basin) by D. accola (Clarke, 1913), and in South Africa by Dalmanitoides sp. (see Rustán & Vaccari, 2012). Thus the distribution of Dalmanitoides could indicate local endemism.
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6.3 Calmoniids Calmoniids are known from the Amazonas, Parnaíba, and Paraná basins, but no genus has yet
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been documented from all three basins. Moreover, where genera are shared by two of the three basins, they are represented by different species. Thus, all the Brazilian calmoniids appear to be endemic to individual basins at species level (see Table 4).
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Calmoniids restricted to the Amazonas Basin:
?Acastoides menurus, Malvinella australis,? M. tumiloba, “Palpebrops” goeldi, Phacopina
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braziliensis, Vogesina gemellus, and the following gen. indet. species; acanthurus, galea, macropyge, scirpeus, all from the Maecuru Formation, and Eldredgeia paituna from the Ererê Formation.
Calmoniids restricted to the Parnaíba Basin
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Eldredgeia cf. E. venusta and Metacryphaeus kegeli occur only in the Picos Member of the Pimenteira Formation. Metacryphaeus meloi occurs in the overlying Passagem Member of the same formation.
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Calmoniids restricted to the Paraná Basin
Calmoniids are recorded from the Alto Garças and Apucurana sub-basins. One of the three
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calmoniid taxa documented from the Alto Garças sub-basin is known only from Mato Grosso State (?Calmonia triacantha); the other two (Kozlowskiaspis subseciva and Metacryphaeus australis) are more widespread, occurring also in the Apucarana Sub-basin. Several calmoniids are known at present only from the Apucarana Sub-basin. Forms apparently unique to the Ponta Grossa Formation include ?Calmonia gonzagana, C. michrischia, C. signifer, Paracalmonia cuspidata, P. mendesi, P. paranaensis, P. pessula, P. salamunii, Paranacaste
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pontagrossensis, Paraphacopina pontagrossensis (nomem nudum), Pennaia pauliana and P. lombardi. Two other taxa are known only from the Tibaji Member (Tibagya parana and Bainella
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paranaense). Metacryphaeus australis occurs in both sub-basins, as does Kozlowskiaspis subseciva. The distribution of the calmoniids within the three basins is shown in Table 4.
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7. DISCUSSION
The Devonian trilobite assemblages of Brazil share many common features with coeval faunas of
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other Malvinokaffric areas, including low family-level diversity (in fact even lower in Brazil than elsewhere), a mixture of cosmopolitan and endemic taxa at generic level, and an apparently high endemism at species level. When the geographic distribution of Malvinokaffric trilobite assemblages is examined at the genus level, several interesting patterns emerge.
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The homalonotid Burmeisteria is the only trilobite genus known to occur in all three Brazilian basins (although it is poorly represented in the Amazonas Basin). Burmeisteria herscheli occurs only in the Paraná Basin of Brazil, but it is also known from the Falklands (Malvinas) Islands and
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South Africa and was thus quite widespread. On the other hand, B. notica is known only from the Parnaíba and Paraná basins, and B. oiara is restricted to the Amazonas Basin. Another species, B.
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accraensis is known from Ghana (Saul, 1967) but has not been found in other Malvinokaffric assemblages. Thus, endemism is likely present at species level within Burmeisteria. Dalmanitids are represented in Brazil by only two genera, each of which is apparently restricted to a single basin (Dalmanitoides in the Paraná Basin, Amazonaspis in the Amazonas Basin). No calmoniid genus occurs in all three Brazilian basins (at best, some genera are shared by two out of the three; see Table 4). However, many Brazilian calmoniid genera also occur elsewhere,
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although they are still somewhat restricted in their overall distribution. For example, several genera from the Amazonas Basin are known otherwise only from Bolivia, including Malvinella,
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Phacopina, and Vogesina. Genera shared by the Paraná Basin and Bolivia include Calmonia, Kozlowskiaspis, and Pennaia; Calmonia is also documented from Argentina (Vaccari et al., 1994), and a species from South Africa was assigned to Pennaia (Cooper, 1982).
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The most restricted distribution patterns involve genera shared by Bolivia and just one Brazilian basin, for example the calmoniid Bainella, which is known only from the Paraná Basin, but also
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occurs in Bolivia, Argentina, South Africa, and the Falkland (Malvinas) Islands. Another calmoniid, Eldredgeia, has a contrasting distribution that excludes the Paraná Basin but includes the Amazonas and Parnaíba basins as well as Bolivia and South Africa, but it apparently does not occur in the Falkland (Malvinas) Islands. Metacryphaeus is more widespread, occurring in the
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Paraná and Parnaíba basins, Bolivia, South Africa, and the Falkland (Malvinas) Islands although it has not yet been reported from Amazonas Basin. Dalmanitoides is also very widespread, although in Brazil it is known only from the Paraná Basin. Even in cases where genera are
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widespread within the Malvinokaffric Realm, many species have highly restricted distributions (for example, Metacryphaeus rotundatus, from the Paraná Basin and Bolivia; M. tuberculatus,
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from the Parnaíba Basin and Bolivia). Only two trilobite genera (Digonus and Burmeisterella) are known both from Malvinokaffric localities and from farther afield.
8. ACKNOWLEDGMENTS
The authors are very grateful to S. Thurston and J. Denton (AMNH) for making the maps and tables, to L. Cazes and P. Richir (MNHN) for photography and cast-making. We thank V.
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Fonseca and A.C. Fernandes (MN), R. Machado (DNPM/RJ), S. Charbonnier and J-M.Pacaud (MNHN), D. Meyer and C. Brett (UC), B. Hunda (CMC) and E. Landing (NYSM) for permitting
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access to specimens in their institutions. We are grateful to J.G. Maisey (AMNH) for
improvements to the English, and to J. H. Melo (Petrobras) for supplying biostratigraphic data. Finally, we thank to two anonymous reviewers for their careful and helpful revision of the
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manuscript.
Funding for L.C.M.O.Ponciano was provided by Fundação Carlos Chagas Filho de Amparo à
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Pesquisa do Estado do Rio de Janeiro (FAPERJ E-26/110.505/2014).
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FIGURE CAPTIONS
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Fig. 1. Map of Brazil showing the Paleozoic basins and the tectonic structures (I-XII) that delimit them (Modified from Grahn, 1992). I. Iquitos High; II. Caruari High; III. Purus High; IV. Monte Alegre High; V. Gurupá High; VI. Tocantins High; VII. Ferres/Urbano Santos High; VIII. São Francisco High; IX. Upper Xingu High; X. Goiânia High; XI. Campo Grande High; XII. Três Lagoas High.
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Fig. 2. Homalonotid, dalmanitid and calmoniid trilobites from Amazonas Basin. 1. Digonus derbyi (Clarke, 1890). Syntype, MN 3370- I, dorsal view of cast from an external mold of cephalon. 2- 3. Amazonaspis maecurua (Clarke, 1890). 2, Lectotype, MN 3383-I, dorsal view of an incomplete cephalon; 3, Paralectotype, MN 3387-I, dorsal view of interal mold of pygidium (from Carvalho and Fonseca, 2007). 4. Dalmanitidae gen. and sp. indet., dorsal view of an internal mold of an incomplete pygidium. 5. ? Malvinella tumiloba (Clarke, 1890), Syntype, MN 3392-I, internal mold of partial cranidium. 6. Phacopina braziliensis (Clarke, 1890), Holotype, MN 3375-I, internal mold of cephalon. 7. Malvinella australis (Clarke, 1890), Holotype, MN 3389-I, dorsal view of internal mold of cephalon. 8. Gen. ind. acanthurus (Clarke, 1890), Holotype, MN 3374-I, dorsal view of an incomplete pygidium. 9. Eldredgeia paituna (Hartt and Rathbun, 1875), Syntype, MN 3394- I, dorsal view of internal mold of partial cephalon. Specimens 1 to 8 are from the Maecuru Formation, Middle Devonian (latest Emsian-early Eifelian), specimen 9 is from the Ererê Formation, Middle Devonian (latest Eifelian-early Givetian). Scale bar = 10 mm
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Fig. 3. Homalonotid and calmoniid trilobites from Parnaíba Basin 1. Burmeisteria notica (Clarke, 1913). DGM, DNPM 5005- I, dorsal view of an internal mold of a well preserved complete specimen. 2. Metacryphaeus tuberculatus (Kozlowski, 1923). CMC IP 37743, dorsal view of an internal mold of cephalon. 3. Eldredgeia cf. Eldredgeia venusta (Wolfart, 1968). CMC IP 46411 (former UCGM 46411), dorsal view of internal mold of cephalon and anterior part of thorax. 4. Metacryphaeus kegeli (Carvalho et al., 1997). DGM, DNPM 6133- I, Holotype, dorsal view of internal mold of cephalon. 5. Metacryphaeus meloi (Carvalho et al, 1997). DGM, DNPM 6822- I, Holotype, dorsal view of cephalon. Specimens 1 to 4 are from the Picos Member of the Pimenteira Formation, Middle Devonian (late Eifelian- early Givetian); specimen 5 is from the Passagem Member of the Pimenteira Formation, Middle Devonian (early Givetian). Scale bar = 10 mm Fig. 4. Homalonotid trilobites from the Paraná Basin. 1. Burmeisteria notica (Clarke, 1913). Lectotype, DGM, DNPM 55- I, dorsal view of a complete specimen, internal mold. 2. Burmeisterella braziliensis Carvalho, 2006. 2.A, Holotype, MN 7588- I, dorsal view of thorax and pygidium; 2.B, latex cast showing thoracic spines; 2.C, anterior view of cephalon. Specimen 1 is from the Ponta Grossa Formation; specimen 2A-C is from the Chapada Group (State of Mato Grosso). Scale bar: 1 = 30 mm; 2 -3 = 10 mm
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Fig. 5. Calmoniids from Paraná Basin. 1. Calmonia signifer Clarke, 1913. Type species DGM - DNPM 5- I, dorsal view of a complete specimen, internal mold. 2. ?Calmonia gonzagana (Clarke, 1890). Type species DGM - DNPM 19-I, dorsal view of part of thorax and pygidium, internal mold. 3. Bainella paranaense Carvalho and Edgecombe, 2006, Holotype DGM - DNPM 1708- I, dorsal view of partial cephalon and first thoracic segment, internal mold. 4. Metacryphaeus australis (Clarke, 1913). Holotype DGMDNPM 35- I, dorsal view of an individual almost complete, internal mold. 5. Pennaia lombardi (Kozlowski, 1913), Holotype IPM R 50873, dorsal view of a silicone peel of external mold, complete specimen. 6. Paranacaste pontagrossensis Popp, 1989, IGEO-UFRJ 127- Tr, dorsal view of a latex cast of external mold of an almost complete specimen. 7. Tibagya parana (Clarke, 1913). Holotype DGM, DNPM 69-I, dorsal view of incomplete pygidium, internal mold. Scale bar = 10 mm
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Fig. 6. Calmoniid and dalmanitid from Paraná Basin. 1-2. ?Calmonia triacantha Carvalho and Edgecombe, 1991. 1, Paratype, MN 6948-I (former CENPES 696-I), oblique posterolateral view of thoracopygidium; 2, Holotype, MN 6942-I (former CENPES 690-I), latex cast from external mold, dorsal view of cephalon, incompletely preserved. 3 A- B, Paracalmonia cuspidata (Clarke, 1913). Lectotype, DGM, DNPM 78-I, internal and external mold of cephalon. 4. Paracalmonia pessula (Clarke, 1913). Lectotype, DGM, DNPM 85-I, internal mold of cephalon with some thoracic segments. 5. Kozlowskiaspis subseciva (Clarke, 1913). Holotype, DGM, DNPM 25-I, internal mold of a complete individual. 6. Dalmanitoides accola (Clarke, 1913), internal mold of an individual almost complete, Holotype DGM, DNPM 43-I. Scale bar = 10 mm
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TABLE 1. Trilobites from the Amazonas Basin Species
Maecuru Fm.
Ererê Fm.
Homalonotidae Digonus derbyi Burmeisteria oiara
X
Dalmanitidae X X X
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Amazonaspis maecurua ?Dalmanites infractus Dalmanitidae gen. and sp. indet. Calmoniidae
Uncertain Family
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X
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?Acastoides menurus Eldredgeia paituna Malvinella australis ?Malvinella tumiloba “Palpebrops” goeldi Phacopina braziliensis Gen. indet. acanthurus Vogesina gemellus Gen. indet. galea Gen. indet. scirpeus Gen. indet. macropyge
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X
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“Phacops” pullinus Clarke, 1890
X X X X X X X X X
X
X
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TABLE 2. Trilobites from the Parnaíba Basin Species
Pimenteira Fm. Picos Mb.
Pimenteira Fm. Passagem Mb.
X
X
Burmeisteria notica Calmoniidae Eldredgeia cf. E. venusta Metacryphaeus kegeli Metacryphaeus meloi Metacryphaeus tuberculatus
X X
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Homalonotidae
X
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X
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Alto Garças Sub-basin Chapada Group
Apucarana Sub-basin
Mt. Gr.
P. Grossa Fm.
Tibaji Mb.
X X
X X
Goiás
Paraná Group
Homalonotidae Burmeisteria notica Burmeisteria herscheli Burmeisterella braziliensis
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X
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TABLE 3. Trilobites from the Paraná Basin Species
Dalmanitidae
X
Calmoniidae
X X X
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Bainella paranaense ? Calmonia gonzagana Calmonia michrischia Calmonia signifer Kozlowskiaspis subseciva Calmonia triacantha Metacryphaeus australis Metacryphaeus aff. M. australis Metacryphaeus rotundatus Paracalmonia cuspidata P. mendesi P. paranaensis P. pessula P. salamunii Paranacaste pontagrossensis Pennaia lombardi P. pauliana Tibagya parana
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Dalmanitoides accola
X
X
X X X X
X
X
X
X X X X X X X X X
X
X
X
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TABLE 4. Calmoniidae occurrences in the Amazonas, Parnaíba and Paraná basins Species Amazonas Basin Parnaíba Basin Paraná Basin Mae. Ere. Picos Passag. Alto Garças Apucarana Fm. Fm. Mb. Mb. Sub-basin Sub-basin M. Gr. Goiás P. Gro. Tib. ? Acastoides menurus X Bainella paranaense X ?Calmonia gonzagana X Calmonia michrischia X Calmonia signifer X ?Calmonia triacantha X Eldredgeia paituna X Eldredgeia cf. E. venusta X Kozlowskiaspis subseciva X X X X Malvinella australis X ?Malvinella tumiloba X Metacryphaeus australis X X X M. aff. M. australis X Metacryphaeus kegeli X Metacryphaeus meloi X Metacryphaeus X X rotundatus Metacryphaeus X tuberculatus “Palpebrops” goeldi X Paracalmonia cuspidata X Paracalmonia mendesi X Paracalmonia paranaensis X Paracalmonia pessula X Paracalmonia salamunii X Paranacaste X pontagrossensis Pennaia lombardi Pennaia pauliana X Phacopina braziliensis X Gen. idet. acanthurus X Tibagya parana X Vogesina gemellus X
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Highlights 1. Trilobite data are summarized from widely scattered literature.
3. Devonian trilobite family-level diversity in Brazil is low.
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4. Trilobite distributions show various biogeographic patterns.
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2. Stratigraphic and sedimentological data are updated.
S0895-9811(15)30075-4
DOI:
10.1016/j.jsames.2015.10.010
Reference:
SAMES 1476
To appear in:
Journal of South American Earth Sciences
Received Date: 18 July 2014 Revised Date:
12 October 2015
Accepted Date: 17 October 2015
Please cite this article as: Carvalho, M.d.G.P.d., Ponciano, L.C.M.d.O., The Devonian trilobites of Brazil: A summary, Journal of South American Earth Sciences (2015), doi: 10.1016/j.jsames.2015.10.010. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
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THE DEVONIAN TRILOBITES OF BRAZIL: A SUMMARY
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MARIA DA GLORIA PIRES DE CARVALHO American Museum of Natural History, Division of Paleontology, Research Associate. Central Park West and 79th Street, New York, N.Y. 100-24-5192, USA. [email protected]
ABSTRACT
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LUIZA CORRAL MARTINS DE OLIVEIRA PONCIANO Universidade Federal do Estado do Rio de Janeiro, Departamento de Ciências Naturais, Laboratório de Tafonomia e Paleoecologia Aplicadas – LABTAPHO, Professor. Av. Pasteur, 458, sala 504, 22290-255, Rio de Janeiro, RJ, Brazil. [email protected]
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Devonian trilobites are found in three major Paleozoic intracratonic basins of Brazil (Amazonas, Parnaíba, and Paraná). The trilobites represent the families Homalonotidae, Dalmanitidae, and Calmoniidae. The distribution of these taxa in the Brazilian territory is summarized here because of their remarkable scientific and historical importance, and a revised taxonomy and
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lithostratigraphy of the Devonian (Pragian - Famennian) trilobites from Brazil is presented, based on new research and recent literature. Homalonotids and dalmanitids are relatively cosmopolitan, whereas calmoniids are more endemic and seem to have been restricted to marine cold-waters of
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the southern hemisphere (Malvinokaffric Realm). Although the trilobites within the Brazilian intracratonic basins are approximately contemporaneous, they show various patterns of endemism
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and biogeographical affinities with other Malvinokaffric areas such as Bolivia, South Africa, and the Falkland (Malvinas) Islands. At family level, therefore, trilobite diversity from Brazil is comparatively low, which may indicate biogeographical filtering related to the distance and/or remoteness of the Brazilian basins from more open oceanic waters. Key words: Trilobites, Devonian, Brazil.
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1. INTRODUCTION Three large Brazilian intracratonic basins are discussed in this paper: the Amazonas Basin,
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located in the North; the Parnaíba Basin of the North/Northeast; and the Paraná Basin in the Southeast/South (Fig.1). A fourth major Paleozoic basin (Solimões, in northwestern Brazil) is still poorly studied in terms of its macrofossils and therefore not included in the scope of this
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paper. Each of the basins contains a considerable thickness of Middle Paleozoic clastic
sediments, partly exposed in extensive outcrop belts along their margins. Marine Devonian
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sequences are well represented in these basins and provide evidence of widespread marine transgressions across Brazil, especially during the Pragian - Frasnian. The marine invertebrate fossils discovered in these sequences include trilobites, which are restricted to the families Homalonotidae, Dalmanitidae, and Calmoniidae. The Calmoniidae is considered to be endemic
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to the marine cold-water, southern hemisphere Malvinokaffric Realm. The other two families are more cosmopolitan, whereas their Brazilian representatives seem to be somewhat endemic at genus or species level. The distribution of these trilobites among the Brazilian intracratonic
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basins is shown in Tables 1-4.
The first evidence about Devonian marine invertebrate fossils from Brazil was found during the
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pioneering Morgan expeditions in 1870-71, led by Charles Frederick Hartt, in the Amazonas Basin, and during the Imperial Geological Commission expeditions of 1876-77, directed by Orville Adelbert Derby. A series of publications dealing mainly with the shelly fossils collected on these expeditions were published between 1871 and 1913 (see Melo, 1988 for references). Besides Katzer (1903, 1933) and Melo (1988), only a few additional papers have been published on the Devonian invertebrate fossils from this area during the last years (see Ponciano, 2011 for
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references). Devonian strata of the Parnaíba Basin were first discovered by Small (1914) near to the
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Pimenteira village (nowadays the city of Pimenteiras), in the State of Piauí, but were recognized as Devonian only by Caster (1948). Wilhelm Kegel subsequently collected many marine
invertebrate fossils from this basin in the mid-C20th (Kegel, 1953). Subsequently, Melo (1988),
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Carvalho et al. (1997), Fonseca (2004), Ponciano and Della Favera (2009), Fonseca and Ponciano (2011), Fernandes et al. (2012), Ponciano et al. (2010, 2012a, b), and other few papers have
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accorded attention to Devonian macrofossil assemblages from the Parnaíba Basin. The Devonian of the Paraná Basin has been investigated in several stratigraphic, sedimentological, and paleontological studies. Derby (1878b) first recognized Devonian fossils in this basin. Clarke’s (1913) monograph is a classic publication and still represents one of the most
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important contributions on the Devonian marine invertebrates of the Paraná Basin. Clarke (1913) recognized the distinction between austral and boreal faunas, and also associated the Brazilian fauna with the Falkland (Malvinas) Islands and South Africa, uniting them as an “Austral Fauna”.
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Later, Richter and Richter (1942) erected the terminology “Malvinokaffric Province” for the endemic Devonian trilobites recovered from various localities in the Southern Hemisphere
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(Falkland/Malvinas Islands, South Africa, Brazil - State of Paraná, Ponta Grossa Region). This later became known as the “Malvinokaffric Realm” (Eldredge and Ormiston, 1979). More recent studies have found evidence for an early to early late Emsian decline of the Malvinokaffric fauna in the Paraná Basin (Bosetti et al., 2012), as well as an early Givetian “Lilliput Effect” associated with the collapse of the Malvinokaffric shelly fauna (Bosetti et al., 2010) supposedly with up to 90% size reduction of surviving species, although the extent to which the trilobites were affected
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is unclear. 2. MATERIAL AND METHODS
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The material studied here is represented by internal and external molds, mostly disarticulated; there are some almost complete specimens, but generally they are poorly preserved. Specimens preserved inside concretions (especially those from the Parnaíba Basin) have better preservation.
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Open nomenclature has been retained for certain taxa, due to the incompleteness and poor
preservation of the specimens on which they are based, especially those from the Amazonas
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Basin. Although some specimens were coated with magnesium oxide prior to photography, in many cases this was not possible for conservational reasons.
The trilobites illustrated or cited in this paper are housed in the following institutions: AMNH, American Museum of Natural History, New York.
Facility, Cincinnati.
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CMC IP, Cincinnati Museum Center Invertebrate Paleontology, Geier Collections and Research
DCN/UNIRIO, Departamento de Ciências Naturais da Universidade Federal do Estado do Rio de
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Janeiro, Rio de Janeiro.
DGM, DNPM, Departamento Nacional de Produção Mineral, Rio de Janeiro.
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DZP/UNESP, Departamento de Zoologia da Universidade Estadual de São Paulo, Botucatu. MCT, Museu de Ciências da Terra, Setor de Paleontologia, DNPM, Rio de Janeiro. MNHN, Paris- R, Museum National d’Histoire Naturelle, Paris. NYSM, New York State Museum, Albany. UC, University of Cincinnati, Cincinnati. IGEO/UFRJ, Instituto de Geociências da Universidade Federal do Rio de Janeiro, Rio de Janeiro.
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3. AMAZONAS BASIN
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MN/UFRJ, Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro.
According to Cunha et al. (2007) the Devonian lithostratigraphic units of the Amazonas Basin comprise (in ascending order), the upper part of the Trombetas Group (Manacapuru and Jatapu
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formations), the Urupadi Group (Maecuru and Ererê formations) and the main part of the Curuá Group (Barreirinha and Curiri formations). Trilobites are documented only from the Maecuru
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and Ererê formations. 3.1 Maecuru Formation
The Maecuru Formation (sensu Derby, 1878) consists mainly of a sequence of fluvio-deltaic to neritic sandstones, with few siltstone interbeds. On the northern flank of the Amazonas Basin this
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unit is developed as shallow marine tempestites. These beds contain the highest diversity of shelly faunas in any Brazilian Devonian assemblages (Melo, 1988). The type locality, according to Lange (1967, p. 237), is a poorly delimited outcrop section exposed along the Maecuru River
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bedrock, upstream of the Teuapixuna Rapids (where the overlying Ererê Formation is exposed) and downstream of the Pancada Grande Waterfall (outcrop site of the underlying Trombetas
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Group).
Trilobites (Fig.2) occur in the upper part of the former Lontra Member sensu Lange (1967) [= Maecuru Formation sensu stricto after Derby (1878) and Cunha et al. (2007)], which represents a shallow marine facies of Middle Devonian (latest Emsian-early Eifelian) age (Grahn and Melo, 2004). Homalonotids are represented by a single taxon, Digonus derbyi (Clarke, 1890). Dalmanitids include Amazonaspis maecurua (Clarke, 1890), ?Dalmanites infractus Clarke, 1890,
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and Dalmanitidae gen. and sp. indet. Calmoniids are considerably more diverse, with ?Acastoides menurus (Clarke, 1890), Malvinella australis (Clarke, 1890), ?Malvinella tumiloba (Clarke,
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1890), “Palpebrops” goeldi (Katzer, 1903), Phacopina braziliensis (Clarke, 1890), Vogesina gemellus (Clarke, 1890), plus several gen. indet. species, including galea Clarke, 1890, scirpeus Clarke, 1890, macropyge Clarke, 1890, and acanthurus Clarke, 1890. Another trilobite is of
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uncertain affinity at family level, “Phacops” pullinus Clarke, 1890.
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3.2 Ererê Formation
Derby (1878) introduced the term Ererê Formation for a section of siltstones with interbedded shale and fine-grained sandstones, above the Maecuru Formation and below the Curuá Group. The age of this formation is latest Eifelian to early Givetian (Grahn and Melo, 2004). The type
region, State of Pará.
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area of the Ererê Formation occupies the central part of the Ererê Dome in the Monte Alegre
Marine invertebrate macrofossils in this formation are impoverished, especially the trilobites
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(Fig.2), but they are distributed in a wide variety of lithologies (sandstones to shales). They mostly occur in the lower level of this unit, which is according to microfossil evidence late
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Eifelian (Grahn and Melo, 2004). Trilobites are rare, but are represented by a homalonotid, Burmeisteria oiara (Hartt and Rathbun, 1875) and a calmoniid, Eldredgeia paituna (Hartt and Rathbun, 1875).
4. PARNAÍBA BASIN The Devonian succession of the Parnaíba Basin is represented, in ascending order, by the upper
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Jaicós (Early Devonian pro parte), Itaim, Pimenteira, Cabeças, and lowermost Longá formations (Vaz et al., 2007). During the Devonian, this intracratonic basin had a more restricted access to
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the open ocean than adjacent contemporary basins; hence, its sediments were deposited under shallow marine conditions (Grahn et al., 2006).
No trilobite body fossils have been recovered from the Jaicós, Itaim, and Cabeças formations, but
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trilobite trackways (possible from homalonotids) have been found in Itaim Formation (Santos and Carvalho, 2009). Trilobites body fossils occur mainly in the Pimenteira Formation (Picos and
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Passagem members), and are scarcer in the Longá Formation.
4.1 Picos Member of the Pimenteira Formation
The Picos Member of the Pimenteira Formation (Kegel, 1953) consists of interlayered fine-
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grained hummocky cross-stratified sandstones and siltstones/shales. The base of the Pimenteira Formation marks the first widespread Devonian transgression across the Parnaíba Basin and conformably overlies the Itaim Formation, which has a much sandier character and is probably of
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fluvial to deltaic and shallow marine origin (Melo, 1988). Grahn et al. (2008) assigned a late Eifelian-early Givetian age span to the Pimenteira Formation
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in the eastern outcrop belt of the Parnaíba Basin, based on chitinozoans obtained from shallow drill cores. In the basin’s subsurface and southwestern outcrop belt, however, the upper Pimenteira Formation may be as young as Frasnian or early Famennian (Loboziak et al., 2000; Grahn et al., 2006). Breuer and Grahn (2011a, b) retrieved late early Givetian palynofloras from Picos Member pelites laterally equivalent to the Passagem Member sandstones in the eastern Parnaíba Basin.
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Trilobites from the basal part of the Pimenteira Formation (Picos Member; Fig.3) include a homalonotid, Burmeisteria notica (Clarke, 1913), plus the calmoniids Eldredgeia cf. E. venusta
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(Wolfart, 1968), Metacryphaeus kegeli Carvalho, Edgecombe and Lieberman (1997), and Metacryphaeus tuberculatus (Kozlowski, 1923); figured here for the first time from the
Pimenteira Formation (Fig.3.2). Burmeisteria notica is very abundant and well preserved in
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ironstone concretions, especially near the city of Picos (State of Piauí). More rarely,
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Metacryphaeus tuberculatus and other fossil invertebrates also are found in these concretions.
4.2 Passagem Member of the Pimenteira Formation
The upper part of the Pimenteira Formation comprises moderately fossiliferous, highly micaceous fine-grained sandstones with asymptotic and hummocky cross-stratification, and subordinate
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siltstone interbeds constituting Kegel’s (1953) Passagem Member (attributed by him to the lowest part of the Cabeças Formation). This Member is known only in the eastern outcrop belt of the basin (State of Piauí), where it is currently dated as early to early middle Givetian (Grahn et al.,
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2008; Breuer and Grahn, 2011a, b).
The regional unconformity separating the Passagem sandstones from those of the Cabeças
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Formation sensu stricto (latest Famennian age; Grahn et al., 2006) is observed at Pedro Segundo Ridge and the environs of São José do Piauí village, situated respectively in the northeastern and eastern parts of the State of Piauí (Ponciano et al., 2012a). According to recent interpretations (Ponciano and Della Favera, 2009; Ponciano et al., 2012a), the Passagem Member comprises fine-grained distal mouth-bar deposits, interbedded with delta-front, hummocky cross-stratified, graded sandstone lobes. This entire sequence probably represents a
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flood-dominated fluvio-deltaic system entering shallow marine settings. According to this scenario, turbulent flows during episodic flood events sometimes transcended the limits of the
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littoral zone, thereby reworking littoral and benthic shelf organisms into the resulting shallow marine sandstone bodies.
Trilobites from the Passagem Member (Fig.3) include a calmoniid, Metacryphaeus meloi
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Carvalho, Edgecombe and Lieberman (1997), and a homalonotid, Burmeisteria notica (Clarke,
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1913).
4.3 Longá Formation
The Longá paleoenvironment is generally regarded as a transgressive, storm-dominated, shallow sea (Vaz et al., 2007). The lowest part of this formation is composed of shales with interbedded
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conglomeratic to fine-grained sandstones, and contains an impoverished benthic marine invertebrate assemblage of latest Devonian age (uppermost Famennian), with extremely rare, small and poorly preserved trilobites and brachiopods, together with more abundant mollusks and
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trace fossils. The rest of the formation is of Tournaisian age (Playford et al., 2012), but it has not yielded any shelly fossils to date.
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A small trilobite, only known from a single cephalon and pygidium, was collected by Wilhem Kegel from poorly dated basal Longá Formation outcrops at Barreiras Farm (Valença do Piauí area, State of Piauí, eastern Parnaíba Basin), and referred to “Asteropyge” (Kegel, 1953). It may represent a new form of Metacryphaeus, although the presence of a calmoniid within such a young stratigraphic level (Strunian or perhaps even Tournaisian) is exceptional. It is possible that the fossil may have been reworked from Middle Devonian or older strata (Melo, 1988).
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5. PARANÁ BASIN
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During the Devonian, the Paraná Basin included two sub-basins; the Apucarana Sub-basin in the south, and the Alto Garças Sub-basin in the north. These sub-basins had a different geological evolution and are partially separated from each other by the Três Lagoas and Campo Grande
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structural highs (see Fig.1). The Devonian stratigraphic nomenclature for the Paraná Basin adopted here follows Melo (1988) and Grahn et al. (2010, 2013).
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Within the Apucarana Sub-basin, Melo (1988) and Grahn et al. (2013) regarded Ponta Grossa and São Domingos as separate formations. The Tibaji Member is considered to be a member of the latter. Evans (1894) introduced the term Chapada Group for rocks in the northwestern part of the Paraná Basin (Alto Garças Sub-basin), and the Chapada dos Guimarães is regarded as the type
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area. The lithology differs from stratigraphically equivalent units in the Apucarana Sub-basin mainly in having a higher silt and sand content (the marine environment is shallower in the Alto Garças Sub-basin; Grahn et al., 2013). The Chapada Group still does not have formal
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lithostratigraphic designations. Instead, it is informally divided into Chapada units 1, 2, 3, and 4,
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in ascending order (Melo, 1988).
5.1 Chapada Group
Sediments of the Chapada Group occur in the northern part of the Paraná Basin, within the states of Mato Grosso, Mato Grosso do Sul and Goiás. They also encroach as a thin Paleozoic cover onto the southeastern rim of the adjacent, chiefly Precambrian Parecis Basin to the north (central Mato Grosso State), where Lower Devonian fossils identical to those of Chapada dos Guimarães
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are known to occur (Carvalho and Edgecombe, 1991; Carvalho, 1997; Boucot et al., 2001). Chapada unit 1 (essentially sandy) is largely unfossiliferous (except for trace fossils and land
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plant remains of probable Lochkovian age, in its upper part), and is regarded as an equivalent of the Furnas Formation of the Apucarana Sub-basin, to the south, which has a similar lithology and paucity of fossils. Transitional beds between Chapada units 2 and 3 contain a similar fossil
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assemblage to the upper Ponta Grossa Formation and Tibaji Member of the São Domingos
Formation (Melo, 1988), thus suggesting a late Emsian age for these beds. The lithologies are
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reddish sandstones with conglomeratic levels. The sandstones represent a deltaic environment on a shallow platform, and also contain shale clasts and sporadic channels. The trilobites from Mato Grosso (Figs. 4-6) include the homalonotid Burmeisterella braziliensis Carvalho, 2006, plus the calmoniids Kozlowskiaspis subseciva (Clarke, 1913), ?Calmonia triacantha Carvalho and
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Edgecombe, 1991, and Metacryphaeus australis (Clarke, 1913). Trilobites from Goiás include the calmoniid Metacryphaeus aff. M. australis Carvalho, Melo and Quadros, 1987 and
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Kozlowskiaspis subseciva (Clarke, 1913).
5.2 Ponta Grossa Formation
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The lower part of the Ponta Grossa Formation consists of transgressive, highly fossiliferous gray shales with subordinated siltstones and sandstones. Northeast and southeast of Ponta Grossa city, as well as virtually all over the Apucarana Sub-basin, the formation unconformably overlies coarser, poorer-sorted, and kaolinitic sandstones of the earliest Devonian (Lochkovian) Furnas Formation. These basal Ponta Grossa lithologies are followed by sandy shales with limestones nodules or arenaceous clays. The upper Ponta Grossa section is developed as hard, black, pyritic
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shales. The Ponta Grossa Formation has produced a rich and diverse assemblage of marine
acritarchs, and chitinozoans (Grahn et al., 2013).
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invertebrates including trilobites (Figs. 4-6), as well as latest Pragian-early Emsian miospores,
Two homalonotids are recorded from this unit, Burmeisteria notica (Clarke, 1913) and B.
herscheli (Murchison, 1839) (Simões et al., 2009). There is also a dalmanitid, Dalmanitoides
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accola (Clarke, 1913). Calmoniid trilobites are more diverse, with several genera and numerous species, including Bainella paranaense Carvalho and Edgecombe, 2006, ?Calmonia gonzagana
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(Clarke, 1890), Calmonia michrischia Clarke, 1913, Calmonia signifer Clarke, 1913, Kozlowskiaspis subseciva (Clarke, 1913), Metacryphaeus australis (Clarke, 1913), Metacryphaeus rotundatus (Kozlowski, 1923) (Soares et al., 2008), Paracalmonia cuspidata (Clarke, 1913), Paracalmonia mendesi Popp, Coimbra and Hauch, 1996, Paracalmonia
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paranaensis Popp, Coimbra and Hauch, 1996, Paracalmonia pessula (Clarke, 1913), Paracalmonia salamunii Popp, Coimbra and Hauch, 1996, Paranacaste pontagrossensis Popp, 1989, Paraphacopina polygona Mori and Leme, 2013 (nomem nudum), Pennaia pauliana
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Clarke, 1913, and Pennaia lombardi (Kozlowski, 1913).
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5.3 Tibaji Member of the São Domingos Formation The São Domingos Formation overlies the Ponta Grossa Formation transgressively. The Tibaji Member has a restricted geographical extent, but locally constitutes the lowermost part of the formation. Over much of the Apucarana Sub-basin, the São Domingos Formation is overlain unconformably by the Pennsylvanian-Early Permian Itararé Group and locally (in the subsurface of Paraná State), by the so-called Ortigueira Diamictite, from the late Famennian (Loboziak et al.,
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1995; Milani et al., 2007). The São Domingos Formation consists predominantly of argillaceous shale. Poorly sorted conglomeratic sandstones occur in its basal part, succeeded by shale and
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micaceous siltstone rich in plant debris. The age of the Tibaji Member is early late Emsian, based on chitinozoans (Grahn et al., 2013).
Homalonotids and calmoniids occur in the Tibaji Member (Figs. 4-6), but no dalmanitids have
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been reported. Homalonotids include Burmeisteria notica (Clarke, 1913) and Burmesteria
herscheli (Murchison, 1839). Calmoniids are rarer than in the Ponta Grossa Formation and
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display somewhat lower diversity, but nevertheless include five taxa; Bainella paranaense Carvalho and Edgecombe, 2006, Kozlowskiaspis subseciva (Clarke, 1913), Metacryphaeus australis (Clarke, 1913), Metacryphaeus rotundatus (Kozlowski, 1923), and Tibagya parana (Clarke, 1913).
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6. BIOGEOGRAPHIC OBSERVATIONS Devonian trilobites from the Amazonas, Parnaíba, and Paraná basins of Brazil represent only three families: Homalonitidae, Dalmanitidae, and Calmoniidae. Although homalonotids and
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dalmanitids have a world-wide distribution, calmoniids seem to have been restricted to the coldwater Malvinokaffric Realm of the southern hemisphere. All three families are also well
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represented in Malvinokaffric assemblages from other areas, although those sometimes include members of additional families. For example, proetids occur in Bolivia, South Africa, Argentina, and perhaps in the Falklands (Malvinas) Islands (Eldredge and Ormiston, 1979; Cooper, 1982; Carvalho, 2006; Rustán et al, 2011), and aulacopleurids are known from Bolivia, Argentina and South Africa (Adrain & Edgecombe, 1996; Rustán & Vaccari, 2010). At family level, therefore, trilobite diversity from Brazil is comparatively low, which may indicate biogeographical filtering
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related to the distance and/or remoteness of the Brazilian basins from more open oceanic waters. 6.1 Homalonotids
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Homalonotids are widespread in many shallow-water Malvinokaffric communities, which reduce their value for paleobiogeographical analysis (Eldredge and Ormiston, 1979, p. 148). For
example, Digonus is recorded from Europe (Germany, Belgium, UK, and Czech Republic),
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United States, North and South Africa, and Antarctica, so the occurrence of Digonus derbyi (Clarke, 1890) in the Amazonas Basin is biogeographically uninformative. Similarly,
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Burmeisterella is known from Europe (England, Belgium, and Germany) as well as from the Paraná Basin (Alto Garças Sub-basin). On the other hand, the genus Burmeisteria is currently recognized only from the Devonian of the Malvinokaffric Realm. Burmeisteria oiara occurs only in the Amazonas Basin, B. notica is known from both the Parnaíba and Paraná basins, and B.
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herscheli (Murchison, 1839) is even more widespread, occurring in Brazil (Paraná Basin), the Falkland (Malvinas) Islands, and South Africa. 6.2 Dalmanitids
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Dalmanitids occur in the Lower and Middle Devonian of the Paraná and Amazonas basins, but included taxa differ considerably in their distribution. For example, Amazonaspis maecurua
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(Clarke, 1890) is known only from the Amazonas Basin, but Dalmanitoides Delo, 1935 occurs in several Malvinokaffric assemblages outside Brazil (Rustán & Vaccari, 2012). In Argentina, this genus is represented by D. boehmi (Knod, 1908) and D. drevermanni (Thomas, 1906), in Bolivia by D. scutata Branisa and Vanek, 1973, in Brazil (Paraná Basin) by D. accola (Clarke, 1913), and in South Africa by Dalmanitoides sp. (see Rustán & Vaccari, 2012). Thus the distribution of Dalmanitoides could indicate local endemism.
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6.3 Calmoniids Calmoniids are known from the Amazonas, Parnaíba, and Paraná basins, but no genus has yet
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been documented from all three basins. Moreover, where genera are shared by two of the three basins, they are represented by different species. Thus, all the Brazilian calmoniids appear to be endemic to individual basins at species level (see Table 4).
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Calmoniids restricted to the Amazonas Basin:
?Acastoides menurus, Malvinella australis,? M. tumiloba, “Palpebrops” goeldi, Phacopina
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braziliensis, Vogesina gemellus, and the following gen. indet. species; acanthurus, galea, macropyge, scirpeus, all from the Maecuru Formation, and Eldredgeia paituna from the Ererê Formation.
Calmoniids restricted to the Parnaíba Basin
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Eldredgeia cf. E. venusta and Metacryphaeus kegeli occur only in the Picos Member of the Pimenteira Formation. Metacryphaeus meloi occurs in the overlying Passagem Member of the same formation.
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Calmoniids restricted to the Paraná Basin
Calmoniids are recorded from the Alto Garças and Apucurana sub-basins. One of the three
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calmoniid taxa documented from the Alto Garças sub-basin is known only from Mato Grosso State (?Calmonia triacantha); the other two (Kozlowskiaspis subseciva and Metacryphaeus australis) are more widespread, occurring also in the Apucarana Sub-basin. Several calmoniids are known at present only from the Apucarana Sub-basin. Forms apparently unique to the Ponta Grossa Formation include ?Calmonia gonzagana, C. michrischia, C. signifer, Paracalmonia cuspidata, P. mendesi, P. paranaensis, P. pessula, P. salamunii, Paranacaste
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pontagrossensis, Paraphacopina pontagrossensis (nomem nudum), Pennaia pauliana and P. lombardi. Two other taxa are known only from the Tibaji Member (Tibagya parana and Bainella
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paranaense). Metacryphaeus australis occurs in both sub-basins, as does Kozlowskiaspis subseciva. The distribution of the calmoniids within the three basins is shown in Table 4.
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7. DISCUSSION
The Devonian trilobite assemblages of Brazil share many common features with coeval faunas of
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other Malvinokaffric areas, including low family-level diversity (in fact even lower in Brazil than elsewhere), a mixture of cosmopolitan and endemic taxa at generic level, and an apparently high endemism at species level. When the geographic distribution of Malvinokaffric trilobite assemblages is examined at the genus level, several interesting patterns emerge.
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The homalonotid Burmeisteria is the only trilobite genus known to occur in all three Brazilian basins (although it is poorly represented in the Amazonas Basin). Burmeisteria herscheli occurs only in the Paraná Basin of Brazil, but it is also known from the Falklands (Malvinas) Islands and
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South Africa and was thus quite widespread. On the other hand, B. notica is known only from the Parnaíba and Paraná basins, and B. oiara is restricted to the Amazonas Basin. Another species, B.
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accraensis is known from Ghana (Saul, 1967) but has not been found in other Malvinokaffric assemblages. Thus, endemism is likely present at species level within Burmeisteria. Dalmanitids are represented in Brazil by only two genera, each of which is apparently restricted to a single basin (Dalmanitoides in the Paraná Basin, Amazonaspis in the Amazonas Basin). No calmoniid genus occurs in all three Brazilian basins (at best, some genera are shared by two out of the three; see Table 4). However, many Brazilian calmoniid genera also occur elsewhere,
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although they are still somewhat restricted in their overall distribution. For example, several genera from the Amazonas Basin are known otherwise only from Bolivia, including Malvinella,
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Phacopina, and Vogesina. Genera shared by the Paraná Basin and Bolivia include Calmonia, Kozlowskiaspis, and Pennaia; Calmonia is also documented from Argentina (Vaccari et al., 1994), and a species from South Africa was assigned to Pennaia (Cooper, 1982).
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The most restricted distribution patterns involve genera shared by Bolivia and just one Brazilian basin, for example the calmoniid Bainella, which is known only from the Paraná Basin, but also
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occurs in Bolivia, Argentina, South Africa, and the Falkland (Malvinas) Islands. Another calmoniid, Eldredgeia, has a contrasting distribution that excludes the Paraná Basin but includes the Amazonas and Parnaíba basins as well as Bolivia and South Africa, but it apparently does not occur in the Falkland (Malvinas) Islands. Metacryphaeus is more widespread, occurring in the
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Paraná and Parnaíba basins, Bolivia, South Africa, and the Falkland (Malvinas) Islands although it has not yet been reported from Amazonas Basin. Dalmanitoides is also very widespread, although in Brazil it is known only from the Paraná Basin. Even in cases where genera are
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widespread within the Malvinokaffric Realm, many species have highly restricted distributions (for example, Metacryphaeus rotundatus, from the Paraná Basin and Bolivia; M. tuberculatus,
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from the Parnaíba Basin and Bolivia). Only two trilobite genera (Digonus and Burmeisterella) are known both from Malvinokaffric localities and from farther afield.
8. ACKNOWLEDGMENTS
The authors are very grateful to S. Thurston and J. Denton (AMNH) for making the maps and tables, to L. Cazes and P. Richir (MNHN) for photography and cast-making. We thank V.
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Fonseca and A.C. Fernandes (MN), R. Machado (DNPM/RJ), S. Charbonnier and J-M.Pacaud (MNHN), D. Meyer and C. Brett (UC), B. Hunda (CMC) and E. Landing (NYSM) for permitting
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access to specimens in their institutions. We are grateful to J.G. Maisey (AMNH) for
improvements to the English, and to J. H. Melo (Petrobras) for supplying biostratigraphic data. Finally, we thank to two anonymous reviewers for their careful and helpful revision of the
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manuscript.
Funding for L.C.M.O.Ponciano was provided by Fundação Carlos Chagas Filho de Amparo à
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Pesquisa do Estado do Rio de Janeiro (FAPERJ E-26/110.505/2014).
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FIGURE CAPTIONS
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Fig. 1. Map of Brazil showing the Paleozoic basins and the tectonic structures (I-XII) that delimit them (Modified from Grahn, 1992). I. Iquitos High; II. Caruari High; III. Purus High; IV. Monte Alegre High; V. Gurupá High; VI. Tocantins High; VII. Ferres/Urbano Santos High; VIII. São Francisco High; IX. Upper Xingu High; X. Goiânia High; XI. Campo Grande High; XII. Três Lagoas High.
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Fig. 2. Homalonotid, dalmanitid and calmoniid trilobites from Amazonas Basin. 1. Digonus derbyi (Clarke, 1890). Syntype, MN 3370- I, dorsal view of cast from an external mold of cephalon. 2- 3. Amazonaspis maecurua (Clarke, 1890). 2, Lectotype, MN 3383-I, dorsal view of an incomplete cephalon; 3, Paralectotype, MN 3387-I, dorsal view of interal mold of pygidium (from Carvalho and Fonseca, 2007). 4. Dalmanitidae gen. and sp. indet., dorsal view of an internal mold of an incomplete pygidium. 5. ? Malvinella tumiloba (Clarke, 1890), Syntype, MN 3392-I, internal mold of partial cranidium. 6. Phacopina braziliensis (Clarke, 1890), Holotype, MN 3375-I, internal mold of cephalon. 7. Malvinella australis (Clarke, 1890), Holotype, MN 3389-I, dorsal view of internal mold of cephalon. 8. Gen. ind. acanthurus (Clarke, 1890), Holotype, MN 3374-I, dorsal view of an incomplete pygidium. 9. Eldredgeia paituna (Hartt and Rathbun, 1875), Syntype, MN 3394- I, dorsal view of internal mold of partial cephalon. Specimens 1 to 8 are from the Maecuru Formation, Middle Devonian (latest Emsian-early Eifelian), specimen 9 is from the Ererê Formation, Middle Devonian (latest Eifelian-early Givetian). Scale bar = 10 mm
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Fig. 3. Homalonotid and calmoniid trilobites from Parnaíba Basin 1. Burmeisteria notica (Clarke, 1913). DGM, DNPM 5005- I, dorsal view of an internal mold of a well preserved complete specimen. 2. Metacryphaeus tuberculatus (Kozlowski, 1923). CMC IP 37743, dorsal view of an internal mold of cephalon. 3. Eldredgeia cf. Eldredgeia venusta (Wolfart, 1968). CMC IP 46411 (former UCGM 46411), dorsal view of internal mold of cephalon and anterior part of thorax. 4. Metacryphaeus kegeli (Carvalho et al., 1997). DGM, DNPM 6133- I, Holotype, dorsal view of internal mold of cephalon. 5. Metacryphaeus meloi (Carvalho et al, 1997). DGM, DNPM 6822- I, Holotype, dorsal view of cephalon. Specimens 1 to 4 are from the Picos Member of the Pimenteira Formation, Middle Devonian (late Eifelian- early Givetian); specimen 5 is from the Passagem Member of the Pimenteira Formation, Middle Devonian (early Givetian). Scale bar = 10 mm Fig. 4. Homalonotid trilobites from the Paraná Basin. 1. Burmeisteria notica (Clarke, 1913). Lectotype, DGM, DNPM 55- I, dorsal view of a complete specimen, internal mold. 2. Burmeisterella braziliensis Carvalho, 2006. 2.A, Holotype, MN 7588- I, dorsal view of thorax and pygidium; 2.B, latex cast showing thoracic spines; 2.C, anterior view of cephalon. Specimen 1 is from the Ponta Grossa Formation; specimen 2A-C is from the Chapada Group (State of Mato Grosso). Scale bar: 1 = 30 mm; 2 -3 = 10 mm
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Fig. 5. Calmoniids from Paraná Basin. 1. Calmonia signifer Clarke, 1913. Type species DGM - DNPM 5- I, dorsal view of a complete specimen, internal mold. 2. ?Calmonia gonzagana (Clarke, 1890). Type species DGM - DNPM 19-I, dorsal view of part of thorax and pygidium, internal mold. 3. Bainella paranaense Carvalho and Edgecombe, 2006, Holotype DGM - DNPM 1708- I, dorsal view of partial cephalon and first thoracic segment, internal mold. 4. Metacryphaeus australis (Clarke, 1913). Holotype DGMDNPM 35- I, dorsal view of an individual almost complete, internal mold. 5. Pennaia lombardi (Kozlowski, 1913), Holotype IPM R 50873, dorsal view of a silicone peel of external mold, complete specimen. 6. Paranacaste pontagrossensis Popp, 1989, IGEO-UFRJ 127- Tr, dorsal view of a latex cast of external mold of an almost complete specimen. 7. Tibagya parana (Clarke, 1913). Holotype DGM, DNPM 69-I, dorsal view of incomplete pygidium, internal mold. Scale bar = 10 mm
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Fig. 6. Calmoniid and dalmanitid from Paraná Basin. 1-2. ?Calmonia triacantha Carvalho and Edgecombe, 1991. 1, Paratype, MN 6948-I (former CENPES 696-I), oblique posterolateral view of thoracopygidium; 2, Holotype, MN 6942-I (former CENPES 690-I), latex cast from external mold, dorsal view of cephalon, incompletely preserved. 3 A- B, Paracalmonia cuspidata (Clarke, 1913). Lectotype, DGM, DNPM 78-I, internal and external mold of cephalon. 4. Paracalmonia pessula (Clarke, 1913). Lectotype, DGM, DNPM 85-I, internal mold of cephalon with some thoracic segments. 5. Kozlowskiaspis subseciva (Clarke, 1913). Holotype, DGM, DNPM 25-I, internal mold of a complete individual. 6. Dalmanitoides accola (Clarke, 1913), internal mold of an individual almost complete, Holotype DGM, DNPM 43-I. Scale bar = 10 mm
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TABLE 1. Trilobites from the Amazonas Basin Species
Maecuru Fm.
Ererê Fm.
Homalonotidae Digonus derbyi Burmeisteria oiara
X
Dalmanitidae X X X
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Amazonaspis maecurua ?Dalmanites infractus Dalmanitidae gen. and sp. indet. Calmoniidae
Uncertain Family
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?Acastoides menurus Eldredgeia paituna Malvinella australis ?Malvinella tumiloba “Palpebrops” goeldi Phacopina braziliensis Gen. indet. acanthurus Vogesina gemellus Gen. indet. galea Gen. indet. scirpeus Gen. indet. macropyge
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X
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“Phacops” pullinus Clarke, 1890
X X X X X X X X X
X
X
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TABLE 2. Trilobites from the Parnaíba Basin Species
Pimenteira Fm. Picos Mb.
Pimenteira Fm. Passagem Mb.
X
X
Burmeisteria notica Calmoniidae Eldredgeia cf. E. venusta Metacryphaeus kegeli Metacryphaeus meloi Metacryphaeus tuberculatus
X X
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Homalonotidae
X
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X
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Alto Garças Sub-basin Chapada Group
Apucarana Sub-basin
Mt. Gr.
P. Grossa Fm.
Tibaji Mb.
X X
X X
Goiás
Paraná Group
Homalonotidae Burmeisteria notica Burmeisteria herscheli Burmeisterella braziliensis
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X
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TABLE 3. Trilobites from the Paraná Basin Species
Dalmanitidae
X
Calmoniidae
X X X
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Bainella paranaense ? Calmonia gonzagana Calmonia michrischia Calmonia signifer Kozlowskiaspis subseciva Calmonia triacantha Metacryphaeus australis Metacryphaeus aff. M. australis Metacryphaeus rotundatus Paracalmonia cuspidata P. mendesi P. paranaensis P. pessula P. salamunii Paranacaste pontagrossensis Pennaia lombardi P. pauliana Tibagya parana
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Dalmanitoides accola
X
X
X X X X
X
X
X
X X X X X X X X X
X
X
X
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TABLE 4. Calmoniidae occurrences in the Amazonas, Parnaíba and Paraná basins Species Amazonas Basin Parnaíba Basin Paraná Basin Mae. Ere. Picos Passag. Alto Garças Apucarana Fm. Fm. Mb. Mb. Sub-basin Sub-basin M. Gr. Goiás P. Gro. Tib. ? Acastoides menurus X Bainella paranaense X ?Calmonia gonzagana X Calmonia michrischia X Calmonia signifer X ?Calmonia triacantha X Eldredgeia paituna X Eldredgeia cf. E. venusta X Kozlowskiaspis subseciva X X X X Malvinella australis X ?Malvinella tumiloba X Metacryphaeus australis X X X M. aff. M. australis X Metacryphaeus kegeli X Metacryphaeus meloi X Metacryphaeus X X rotundatus Metacryphaeus X tuberculatus “Palpebrops” goeldi X Paracalmonia cuspidata X Paracalmonia mendesi X Paracalmonia paranaensis X Paracalmonia pessula X Paracalmonia salamunii X Paranacaste X pontagrossensis Pennaia lombardi Pennaia pauliana X Phacopina braziliensis X Gen. idet. acanthurus X Tibagya parana X Vogesina gemellus X
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Highlights 1. Trilobite data are summarized from widely scattered literature.
3. Devonian trilobite family-level diversity in Brazil is low.
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4. Trilobite distributions show various biogeographic patterns.
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2. Stratigraphic and sedimentological data are updated.