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Ichnotaxonomy, Functional Morphology and Paleoenvironmental Context of Mesosauridae Tracks from Permian of Brazil

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Ichnotaxonomy, Functional Morphology and Paleoenvironmental Context of Mesosauridae Tracks from Permian of Brazil Revista Brasileira de Geociências Rafael Costa da Silva et al. 39(4): 705-716, dezembro de 2009 Ichnotaxonomy, functional morphology and paleoenvironmental context of Mesosauridae tracks from Permian of Brazil Rafael Costa da Silva¹, Fernando Antonio Sedor² & Antonio Carlos Sequeira Fernandes³ Abstract Subaqueous vertebrate traces are fairly common in fossil record, but there are few studies on the theme due to the difficulties in classification and interpretation. In Brazil, subaqueous tracks were recorded in rocks of Irati Formation (Lower Permian, Paraná Basin). According to morphological features and interpreta- tion, the ichnofossils were designated as a new ichnotaxon, Mesosaurichnium natans igen. nov. et isp. nov., and attributed to the activities of basal reptiles Mesosauridae. Based on faciology, the ichnofossils possibly were produced in a lower shoreface affected by storm waves and do not correspond to the original sedimentary surface, which was covered by a thin layer of crustacean shelly fragments during the production of the tracks. The tracks were produced by mesosaurids during the swimming next to the bottom with the long tail maintain- ing the propulsion. The mesosaurids were able to use the tail as the main propelling organ for fast swimming under undulatory propulsion, but with the paddle-like pedes acting as important accessory propellant during oscillatory propulsion, conferring greater maneuverability during slow swimming. Keywords: subaqueous traces, swimming, Mesosauridae, Permian, Paraná Basin. Resumo Icnotaxonomia, morfologia funcional e contexto paleoambiental de pegadas de Meso- sauridae do Permiano do Brasil. Pegadas subaquáticas de vertebrados são relativamente comuns no re- gistro fossilífero, mas há poucos estudos sobre o tema devido à dificuldade na classificação e interpretação. No Brasil, pegadas subaquáticas foram registradas na Formação Irati (Permiano Inferior, Bacia do Paraná). De acordo com características morfológicas e interpretação morfofuncional, os icnofósseis foram designados como um novo táxon, Mesosaurichnium natans igen. nov. et isp. nov., e atribuídos aos répteis basais da família Mesosauridae. Com base na faciologia, os icnofósseis foram produzidos na área de transição entre as zonas de supramaré e intermaré e não corresponderiam à superfície original, pois essa encontrava-se coberta por uma fina camada de fragmentos de carapaças de crustáceos na ocasião da formação das pegadas. Os mesossauríde- os teriam produzido essas pegadas durante a natação próxima ao fundo, tendo a cauda como principal órgão propulsor. Eles seriam capazes de utilizar a cauda como o principal órgão de propulsão para natação rápida sob propulsão ondulatória, mas com o pé em forma de nadadeira atuando como um importante propulsor acessório durante a propulsão oscilatória, conferindo maior capacidade de manobra durante a natação lenta. Palavras-chave: pegadas subaquáticas, natação, Mesosauridae, Permiano, Bacia do Paraná. INTRODUCTION Vertebrate tracks produced un- ever, there are doubts on the morphofunctional inter- der water are quite common in fossil record, despite pretation, ichnotaxonomy, preservation and paleoenvi- the necessity of special conditions to their preservation. ronmental context, being the aim of this work the dis- However, there are few studies on the theme due to dif- cussion of these questions. ficulties in classification and interpretation of the tracks and its relationships with the trackmakers. In Brazil, sub- MATERIAL AND METHODS The studied mate- aqueous ichnofossils were previously recorded by Sedor rial consists of 25 rock samples with ichnofossils from & Silva (2004) at Irati Formation (Lower Permian, Passa Irati Formation, collected between 1996 and 2003 by Dois Group, Paraná Basin) and interpreted as drag marks the authors F. A. Sedor and R. C. Silva. The samples of fingers produced by swimming reptiles identified as are housed at the paleontological collection of Museu Mesosauridae Baur 1889 (Amniota, Sauropsida), which de Ciências Naturais, Setor de Ciências Biológicas, skeletal remains are abundant in this geological unit. Universidade Federal do Paraná (MCN-SCB-UFPR), This was the first occurrence of ichnofossils at Curitiba, Paraná State, Brazil. related to these animals and provided the first direct The samples MCN.P.293a, MCN.P.293b, evidence of swimming among the Mesosauridae. How- MCN.P.293c, MCN.P.760, MCN.P.761, MCN.P.762, 1 - CPRM – Companhia de Pesquisa de Recursos Minerais – Serviço Geológico do Brasil, Departamento de Geologia, Divisão de Paleon- tologia, Rio de Janeiro (RJ), Brazil. Email: [email protected], [email protected] 2 - Universidade Federal do Paraná, Setor de Ciências Biológicas, Museu de Ciências Naturais, Curitiba (PR), Brazil. Email: [email protected] 3 - Universidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Geologia e Paleontologia, Rio de Janeiro (RJ), Brazil. Email: [email protected] Arquivo digital disponível on-line no site www.sbgeo.org.br 705 Ichnotaxonomy, functional morphology and paleoenvironmental context of Mesosauridae tracks from Permian of Brazil MCN.P.763a, MCN.P.763b, MCN.P.764, MCN.P.765, angles following techniques by Sarjeant (1975). Inter- MCN.P.766, MCN.P.767, MCN.P.774, MCN.P.851, pretative draws were made based in the original sam- MCN.P.852a, MCN.P.852b, MCN.P.852c, MCN.P.852d, ples, photos, casts and replicas. MCN.P.853 and MCN.P.854 were collected at the Co- The footprints were measured using the fol- dapar limestone quarry, inside the limits of Municipality lowing morphometric parameters: trace length (TL), of Guapirama, Paraná State, Brazil (23º 27’ 09.96” S / trace width (TW) and distance between digits (DD). 50º 02’ 10.92” W, Fig. 1). The quarry, which was deacti- The footprints do not consist in impressions of the au- vated in 1998, is located few kilometers of the interstate topodia but drag marks of digits, so is not possible to highway PR 092 and can be accessed through a second- use conventional parameters such as those described by ary road. The samples MCN.P.262 and MCN.P.266 were Leonardi (1987) in measurement. Only the footprints collected at the Metago (Metais de Goiás S/A) lime- with at least two traces and clear limits were considered stone quarry, Municipality of Portelândia, Goiás State, for measuring because many footprints have gradual Brazil (17º 20’ 18.96” S / 52º 37’ 23.88” W, Fig. 1). limits that cannot be precisely defined. The morpho- The samples MCN.P.426, MCN.P.427 and MCN.P.428 logical concepts used in description of the ichnofossils were collected at the Sucal Mineração Ltda limestone were based in Leonardi (1987). Sixteen samples with quarry, Municipality of Perolândia, Goiás State, Brazil thirty four footprints were measured. The mean, me- (17º 29’ 00.0” S / 52º 03’ 11.60” W, Fig 1). dian, standard deviation and the coefficient of variation Silicon rubber casts were made from samples were calculated for each morphometric parameter. MCN.P.293a, MCN.P.293b and MCN.P.293c following techniques by Raup & Stanley (1971), aiming a better GEOLOGICAL CONTEXT The Paraná Basin con- description and illustration of the ichnofossils. Repli- sists in an intracratonic basin of great extension with rocks cas of plaster of Paris and resin were also made. The deposited between Ordovician and Cretaceous (Schneider samples were photographed with artificial light at low et al. 1974, Milani et al, 1994). It covers an area of ap- Figure 1 - Location of the studied outcrops at Paraná and Goiás States, Brazil. 706 Revista Brasileira de Geociências, volume 39 (4), 2009 Rafael Costa da Silva et al. proximately 1.600.000 km2 and occupies part of the ter- (e.g. Schneider et al. 1974, Cunha & França 1994, Mi- ritories of Brazil, Argentina, Uruguay and Paraguay. The lani et al. 1994), based mainly in Daemon & Quadros Passa Dois Group consists in one of the Permian geo- (1969). The Irati Formation is traditionally correlated logical units and comprises the Irati, Serra Alta, Teresi- to Whitehill Formation from Karoo Basin, in Southern na, Corumbataí and Rio do Rasto Formations (Mendes Africa, based in lithology and fossil record. It is also 1967, Milani et al. 1994). The basal unit, Irati Forma- correlated to Mangrullo Formation in Uruguay and to tion, is characterized by dark shales and mudstones, py- the lower portion of Chacabuco Formation, Chaco- robitumous shales, sandstones and limestones. The main Paraná Basin, Argentina (Lavina 1991). However, the sedimentary structure is horizontal lamination, but ripple Whitehill, Mangrullo and Chacabuco Formations were marks, cross lamination, oolites and intraformational con- dated as Sakmarian-Artinskian (e.g. Oelofsen & Araújo glomerate can occur in the carbonatic strata (Schneider et 1987, Lavina 1991), a younger age than the brazilian al. 1974). This formation was deposited in an extensive one. Recently, new palynological and radiometric stud- epicontinental sea, progressively more saline from base to ies attributed the Artinskian age for the Irati Formation top, in southern Gondwana. Fossils of mesosaurs, crusta- (Premaor et al. 2006, Santos et al. 2006). ceans, insects, sponges, fishes, bivalve mollusks and rare brachiopods are known in rocks from Irati Formation. Description of the studied outcrops The Codapar The faciology shows a complex basin geometry during quarry at Guapirama constitutes a vertical exposition the deposition, with the occurrence of gulfs and bays of of approximately 10 meters situated below the ground varied depths
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