Downloaded from http://sp.lyellcollection.org/ at Pennsylvania State University on February 20, 2016 A late Atokan regional encrinite (early late Moscovian, Middle Pennsylvanian) in the Sierra Agua Verde, Sonora state, NW Mexico B. E. B U I T R O N - S , / ~ N C H E Z l, C. G O M E Z - E S P I N O S A 1, E. A L M A Z A N - V J k Z Q U E Z 2 & D. V A C H A R D 3 ' Universidad Nacional Aut6noma de MOxico, Instituto de Geologia, Departamento de Paleontologia, Ciudad Universitaria, Delegaci6n Coyoacdn, 14510 MOxico, D. F., Mexico (e-mail: blancab@servidor, unam. mx) 2Universidad de Sonora, Departamento de Geologia, Boulevard Luis Encinas y Rosales 83000 Hermosillo, Sonora, Mexico 3UniversitO de Lille 1, Sciences de la Terre, UMR 8014 du CNRS, Laboratoire LP3, Bdtiment SN5, 59655 Villeneuve d'Ascq COdex, France (e-mail: daniel, [email protected]) Abstract: In the Sierra Agua Verde, central Sonora state, NW Mexico, the La Joya Formation exhibits an alternation (100 m thick) of calcareous siltstone and fossiliferous limestone with nodular cherts. This latter contains an abundant and diverse late Atokan (i.e. Podolskian =early late Moscovian, Middle Pennsylvanian) fossil assemblage composed of phylloid algae, fusulinids, chaetetids, tabulate corals, gastropods, fenestellid bryozoans, spiriferid and productid brachiopods, crinoids and conodonts. The crinoidal beds constitute a good example of a regional encrinite. They include several species of the parataxonomic stem form-genera Cyclocaudex, Cyclocrista, Heterosteleschus, Mooreanteris, Pentagonopterix, Preptopremnum, Cycloscapus and Pentaridica. Their preservation indicates the combination of preburial decay on the sea floor and post-burial decay within the sediment. The high degree of silicification of the crinoids indicates that they were possibly associated with siliceous organisms (Porifera?), not preserved in the assemblages. The studied thanatocoenosis is typical of tropical shallow seas, and reveals strong biogeographical affinities with the assemblages of the midcontinental and southern regions of the USA. Particularly, the Atokan crinoids of central Sonora are similar to those from Kansas and Texas, confirming the close palaeogeographic connection of southern USA and northern Mexico during the Middle Pennsylvanian. The Sierra Agua Verde is located 120 km eastward of Hermosillo City, Sonora state, in the 'Sierras and Valles del Norte' Mexican subprovince. This range covers a surface area of 255 km 2 and its centre is at 109~176 and 29~176 (Fig. 1). It is composed of sedimentary, igneous and metamorphic rocks of Palaeozoic, Mesozoic and Cenozoic age. Geology and palaeontology of the Sierra Agua Verde were described by Poole et al. (1984), Stewart et al. (1988, 1999), Minjarez-Sosa et al. (1993), Ochoa-Granillo & Sosa-Leon (1993), Stewart & Poole (2002), Mendoza-Madera et al. (2004) and Buitr6n-Sfinchez et al. (2005a, b). The Palaeozoic series includes the following formations: Puerto Blanco, Cuarcita Proveedora, Buelna and Arrojos (Cambrian); E1 Boquinete (Ordovician); E1 Pollo (Devonian); Santiago (Mississippian); La Joya (Pennsylvanian); and Tuntunud6 (Permian). Material and methods The La Joya Formation was logged and sampled bed by bed. All the beds marked as 'crinoids' in Figure 2 were studied, as well as those of other Pennsylvanian outcrops from Sonora in order to investigate several 'regional encrinites' in the sense of Ausich (1997) and Ginsburg (2005). Our study of sequence stratigraphy has determined that the La Joya Formation appears as a transgressive systems tract (TST), whose main lithologies are Chaetetes reef mounds (Almaz/mV~zquen et al. 2007), fusulinellid floatstones, phylloid algae framestones and crinoidal rudstones. These crinoidal rudstones are particularly abundant and well known in the Mississippian and Pennsylvanian of the USA, but only a very few have been studied in Mexico. Our investigation is essentially based on the extraction of the best-preserved specimens of crinoids for ~kLVARO,J. J., ARETZ,M., BOULVAIN,F., MUNNECKE,A., VACHARD,D. & VENNIN,E. (eds) 2007. Palaeozoic Reefs and Bioaccumulations: Climaticand Evolutionary Controls. Geological Society, London, Special From: Publications, 275, 201-209.0305-8719/07l$15.00 9 The Geological Society of London. Downloaded from http://sp.lyellcollection.org/ at Pennsylvania State University on February 20, 2016 202 B.E. BUITRON-S_ANCHEZ E T AL. Fig. 1. Locations of (a) Sonora in Mexico and (b) Sierra Agua Verde in Sonora. taxonomic purposes and a microfacies analysis from a taphonomic point of view, i.e. the taphofacies method of Speyer & Brett (1988) and C6zar (2003). Quantitative analyses based on the techniques summarized by Holterhoff (1996) concerning the crinoid biofacies will be applied to the material (G6mez-Espinosa's PhD 2007); this preliminary study presents only the systematic and taphonomic data obtained in hand specimens and thin sections. Regional encrinites Regional encrinites were defined as crinoidal grainstones and packstones composed of more than 50% by volume of pelmatozoan debris that are at least 5-10 m thick and 500 km 2 in extension (Ausich 1997; Ginsburg 2005). They are an example of taphonomic feedback (Kidwell & Jablonski 1983) by the predominance of one special skeletal grain type. Regional encrinites Downloaded from http://sp.lyellcollection.org/ at Pennsylvania State University on February 20, 2016 ATOKAN REGIONAL ENCRINITE OF NW MEXICO Metres ~ l t / ~ I J I Brachiopods 10o_.~[ ~_": =1= =. _=j_== Fusulinids II , I , IjlCrinoids I L ' .... , " ~Solitary corals / F---i--i-.==- t 8o.-ll I . ~ ,o-lL ',,' I/ ', Synngopora ','J | Jr- ['_~_T_--_:.~Fus~,in'ds / eO--i ~ / ~ 2 - c ; -i - i C.noJ , l e,~ I IChaetetid reef mound !~-_-=--=~_ "_-_~-_-__~-= Solitary corals so-I ,, I w l ~ i ".T-'i--iT l l.,.. I 1 I I I Fusulinids t l '--'-~ --'~'l 40i ";U--'-----L4 ' i ,~ 3o- ~ I z /Crinoids :--- Synngopora ,-i Brachiopods Solitary corals Chaetetids Syringopora Brachiopods 203 occur from the Ordovician to the Jurassic, but the acme of their development in North America is in the early Mississippian (Osagean) (Ausich 1997; Ginsburg 2005). At this time and during the Pennsylvanian individual regional encrinites occur with other carbonate lithofacies in different formations of the USA and northern Mexico. Three main factors related to the evolutionary taphonomical and diagenetical history of crinoids and their skeletal remains are important for the formation of regional encrinites: (1) the long geological history of crinoids is marked by episodes of exceptional abundance, which produce encrinites (Ausich et al. 1999); (2) the stalks and calyces of crinoids consist of separate plates and segments held together with ligamental tissue in a labyrinthine cavity system, the stereome; and (3) the post-mortem decay of this tissue provides plates of the calyx and more numerous segments of the stem, which are highly porous (Ginsburg 2005). These hydraulically light fragments can be entrained, transported and redeposited by currents and orbital wave velocities as low as 16 cm s-1 using the results from flume experiments with crinoid segments from living specimens (Ginsburg 2005). As a result of this ease of movement, bottom sediments of abundant crinoidal debris were probably frequently mobile resulting in the common occurrence of current bedding and graded bedding of encrinites (Ausich 1997; Ginsburg 2005). Furthermore, as highly porous crinoid ossicles are entrained at lower current velocities than solid carbonate particle of the same size (e.g. Fliigel 2004), accumulation rates in the encrinite are generally very high (more than 50% or even 60% of the rock volume), and the regional encrinites are generally deposited below normal wave base but within storm wave base (Fliige12004). Fossil assemblage in Sierra Agua Verde Crinoids Fusulinids ~ Covered base Limestonewith chert nodules -~ Siltstone Fig. 2. Lithological column of the La Joya Formation in Sierra Agua Verde (Sonora, Mexico). The lower contact is not exposed in the field. The upper contact is tectonic (normal fault). Fifteen field samples were collected in the three levels mentioned as 'Crinoids'. Each one contains about 100 crinoids, taxonomically studied. The studied crinoid fauna has been sampled from marine fossiliferous limestone of the La Joya Formation (100 m thick), which is intercalated with nodular cherts and calcareous siltstone (Fig. 2). The associated biota includes: calcareous algae (Eugonophyllum sp., Zidella? sp., Kamaena? sp., Komia eganensis); problematic foraminifers (Pachysphaerina pachysphaerica, Eotuberitina reitlingerae, Insolentitheca horrida); smaller foraminifers (Endothyra ex gr. bowmani, Climacammina ex gr. moelleri, Globivalvulina bulloides); fusulinids (Eostaffella grozdilovae (=E. acuta of the authors), Millerella sp., Pseudostaffella sp., Staffella powwowensis, Downloaded from http://sp.lyellcollection.org/ at Pennsylvania State University on February 20, 2016 204 B.E. BUITR6N-SANCHEZ ETAL. Eoschubertella texana, Fusulinella thompsoni, F. llanoensis, Nipperella? sp.); chaetetids; tabulate corals (Michelinia sp.) and zaphrentid solitary corals (Lophophyllidium sp.); gastropods (Euomphalus sp., cf. Donaldina robusta); brachiopods (Spirifer sp.); and conodonts. These fossils indicate an Atokan age (=Podolskian=early late Moscovian = Middle Pennsylvanian). The crinoid fauna (Fig. 3) includes 11 parataxonomic stem species: Cyclocaudex insaturatus Moore & Jeffords, Cyclocrista martini Miller, Heterosteleschus keithi Miller, Lamprosterigma erathense Moore & Jeffords, Lamprosterigma mirificum Moore & Jeffords, Mooreanteris waylandensis Miller, Pentagonopterix insculptus Moore & Jeffords, Preptopremnum laeve Moore & Jeffords, Preptopremnum rugosum Moore & Jeffords, Cycloscapus laevis Moore & Jeffords and Pentaridica simplicis Moore & Jeffords. This fauna appears similar to those described in the USA by Moore & Jeffords (1968). The thanatocoenosis is typical of tropical shallow seas and shows a strong affinity with the characteristic species of the midcontinental and southern regions of the USA. In particular, the Sonora Pennsylvanian crinoids show similarities to the assemblages of Kansas and Texas (Buitr6nSfinchez et al. 2004, 2005a, b). All these regions are located in tropical and subtropical latitudes during the Pennsylvanian epoch (Heckel 2002). They also have much in common with Eurasiatic-Arctic faunas corresponding to the same climatic zones during this epoch (see, for example, the reconstructions of Scotese & Langford 1995; Ross 1995; Vachard et al. 1997, 2000b, c). Similarly, fusulinids, calcareous algae and smaller foraminifers represented in the La Joya Formation show affinities with Arizona, New Mexico, Texas and California microfaunas. The Permian fusulinids (P6rez-Ramos 1992) share the same affinities. These new data confirm the palaeogeographical reconstructions of Vachard et al. (1997, 2000a-c) and Buitr6n-Sfinchez et al. (2004) in Mexico and Guatemala. Taphonomical implications The taphonomical study of crinoids was not initiated until the 1960s (Ausich et al. 1999). Crinoids are rarely represented as complete fossils, commonly they are disarticulated after death and are preserved as crown parts, cups, arm ossicles, fragments and stem fragments. A few days after death the muscles and ligaments decay, which leads to the disintegration of the skeleton into isolated elements. In order to be preserved completely, a crinoid must be deeply and quickly buried to prevent the reexcavation by currents or disruption by scavengers and burrowers (Donovan 1991; Ausich et al. 1999). In normal marine conditions the echinoderms usually disarticulate into individual ossicles within a period of 1-2 weeks, depending on their construction and environmental factors, but specifically the arms and cirri of recent crinoids begin to disarticulate within the 3 first days after their death, whereas calyx and certain segments of the arms may be disarticulated after 6 days (Dornbos & Bottjer 2001). In crinoids the articulations between multicolumnal segments of the column consist only of intercolumnal ligament, which are zones of structural weakness after death. This was demonstrated in experiments on recent isocrinids. A similar ligamentary organization of the column was observed in crinoids of the Early Mississippian. When a column begins to disarticulate the articulation between segments breaks first, producing fragments of column of almost equal length. Finally, when all the through-going ligaments decay, the whole column disarticulates into individual columnals (Baumiller & Ausich 1992; Ausich & Baumiller 1993). The effects of physical disturbance on echinoderms were studied by Kidwell & Baumiller (1990). The results obtained from laboratory experiments with living crinoids indicate that recently dead crinoids remain articulated through hours of physical disturbance, whereas dead echinoids disarticulate quickly after physical disturbance. In the La Joya Formation the majority of crinoids are represented by fragments of columns and a very low percentage by isolated plates (three specimens in 10 representative samples, each containing approximately 100 characteristic elements of skeleton). Thus, it can be inferred that decay allowed disarticulation of the arms and calyces, and a breaking of the intercolumnar ligaments, but not complete disarticulation of the columns, are evidenced by columns fragments. The preservation of crinoids in La Joya Formation probably shows a combination of decay of the crinoids preburial on the sea floor and decay of crinoids post-burial within the sediments. The limestones rich in crinoids exhibit various taphonomic processes and difference in the depositional environment. Coarse-grained crinoidal limestones are occasionally interpreted as having originated as a result of gravity flows (Cook & Mullins 1983; Martin 1999; Fltigel 2004) but more frequently as tempestites (e.g. Fliige12004). This interpretation is confirmed by the Sierra Agua Verde deposits, owing to the absence of turbidite figures of turbidites and microfacies Downloaded from http://sp.lyellcollection.org/ at Pennsylvania State University on February 20, 2016 ATOKAN REGIONAL ENCRINITE OF NW MEXICO 205 Fig. 3. Late Atokan crinoid fauna from Sierra Agua Verde (Sonora, Mexico). All scale bars are 0.700 mm. (a) Cyclocaudex insaturatus Moore & Jeffords. (b) Cyclocrista martini Miller. (e) Heterosteleschus keithi Miller. (d) Lamprosterigma mirificum Moore & Jeffords. (e) Lamprosterigma erathense Moore & Jeffords. (f) Cycloscapus laevis Moore & Jeffords. (g) Preptopremnum laeve Moore & Jeffords. (h) Preptopremnum rugosum Moore & Jeffords. (i) Mooreanteris waylandensis Miller. (j) Pentagonopterix insculptus Moore & Jeffords and Pentaridica simplicis Moore & Jeffords. (k) Field picture of the crinoidal rudstones in the Sierra Agua Verde. Downloaded from http://sp.lyellcollection.org/ at Pennsylvania State University on February 20, 2016 206 B.E. BUITR6N-SANCHEZ ETAL. analyses that infer tempestites, and probably distal tempestites (according to the criteria summarized by Flfigel 2004, p. 596). The problem is to understand how and when favourable weak currents for suspension feeders like crinoids (see detailed data in Holterhoff 1997) become strong storm currents, i.e. the interaction of organisms with moving fluids. An average accumulation consists of 50% columnal plates that are totally disarticulated, 5% Fusulinella and very rare smaller foraminifers, 5% fenestellid bryozoa (and rare other remains of metazoans) and 40% syntaxial sparite cement. Nevertheless, the associated fusulinellids are poorly preserved, with many truncation facets indicating their allochthonous origin (e.g. Fernfindez-L6pez 2000). The cementation of the distal tempestites with crinoids is very rapid and took place very early, as the ossicles are not affected by compaction or other threedimensional finite strain (compare with Rowan 1991). The character of distal tempestite is also obvious in associated deposits, namely fusulinellid wackestone-packstone, because of the homogeneous micritic matrix and the absence of corrosion features on the fusulinellids, their oligotipy and the absence of associated photophile green algae, all characteristics indicating a rapid burial after transport and sorting. Ginsburg (2005) has underlined that fusulinids offer many opportunities for taphonomic feedbacks that are comparable to that of crinoidal debris. Chaetetid reef mounds, which developed in coeval beds, are located in the lower part of the photic zone, just below wave base (Almaz~mVfi.zquez et al. 2007) - thus supporting the interpretation of the depositional environments of the crinoidal beds. The absence of encrusting algae or foraminifers (e.g. Claracrusta, Calcivertella) on the fusulinellids corroborates the rapid burial of the biota within the microfacies, whereas the absence of ichnofossils in the outcrop indicates an absence of conditions for the establishment of an infrabenthic community. Consequently, in the Atokan Sierra Agua Verde carbonate platform, the crinoid biotopes have been very productive in supplying carbonate fragments for the distal tempestites, first by the ossicles and, secondly, because the stalked crinoids probably acted as substrates for the fusulinellids, another important component of the tempestites (Fig. 4). The chaetetid reef mounds were located in a lower part of the platform and trapped the sediments transported by the storm currents, i.e. the fusulinellids floatstones and the crinoid rudstones. Between the crinoid biotopes and these chaetetid mounds, infaunal organisms were probably inhabiting the micritic substrate (Fig. 4). Fig. 4. Reconstruction of the biotopes of crinoids (1) and fusulinellids (2), with their first accumulations (1.1 and 2.1)and the final deposits (2.1.1 and 1.1.1) accumulated around the chaetetid constructions (3) in the Sierra Agua Verde (Sonora, Mexico). Not to scale. The accumulation of crinoids may also correspond to colder water inputs, similar to those reconstructed in the Permian deposits of Oman (Weidlich 2007). Nevertheless, as indicated above, these crinoids are interpreted as tropical, and the possible mixing of cold and warm waters warrants further discussion in various outcrops and/or areas (see also the 'Auernig paradox' of Samankassou 2002 in the Carnic Alps). Other arguments for possible cold episodes are: (a) that the skeletal composition of the benthic fauna in these levels of the Sierra Agua Verde was principally calcium carbonate (Table 1), and poorly aragonitic, with a dominance of the 'heterozoan association' on the 'photozoan association' (e.g. Samankassou 2002); and (b) the existence of biogenic chert formation (see Beauchamp & Baud 2002) during the Atokan of Sonora, where most of microfossils are preserved by silicification and show internal diagenesis as molds with ornamental features. The source of silica is organic in origin because there is no evidence of volcanism or evaporation zones in the studied area, and because this type of silicification is common in all the investigated encrinites, although their provenances and deposit environments vary. The high degree of silicification suggests the presence of siliceous organisms (Porifera?); consequently, the apparently crinoid-dominated accumulations might correspond to crinoid-sponge associations. The physico-chemical conditions caused the dissolution of these skeletons of associated sponges, and this dissolved silica penetrated and preserved the dead organisms. In the microfacies the first silicifications appear as small points within the crinoid network. The lithostatic compression can be inferred preferentially to the sedimentation, and the silica could also have been mobilized from the interbedded siltstone from burial Downloaded from http://sp.lyellcollection.org/ at Pennsylvania State University on February 20, 2016 ATOKAN REGIONAL ENCRINITE OF NW MEXICO 207 Table 1. Original mineralogy composition of the benthicfauna present in Agua Verde range C, common; X, less common (after Martin 1999) Taxon Gastropods Brachiopods Sponges Bryozoans Echinoderms Benthic foraminifers Aragonite Calcite low Mg Calcite high Mg C C X C C C X Aragonite + Calcite Silica C C C X diagenesis. Therefore, more geochemical investigations are necessary in order to elucidate the fossil diagenesis of these crinoid accumulations. Conclusions 9 The crinoid assemblage from the La Joya Formation is diversified and composed of 11 stem-based species. 9 The crinoids are typical forms of Pennsylvanian age; more precisely, they are late Atokan in age, as indicated by the associated Fusulinella. 9 Chaetetids, corals, fusulinellids, calcareous algae and brachiopods indicate tropical seas with normal salinity. Therefore, these conditions can be assumed for the crinoids; nevertheless, their possible value as local proxies of colder waters input is debatable. 9 The biotopes of each group are probably located in different environments along a carbonate platform and largely depending on the current system. 9 Crinoids indicate the presence of a hard substrate where they colonized, and were probably in turn the substrate for many epifaunal organisms, the fusulinellids for instance. 9 The crinoids of the La Joya Formation, represented by fragments of columns and a very low percentage of isolated plates, indicate that decay allowed disarticulation of the arms and calyx and the breaking of the intercolumnar ligaments, but not the through-going ligament of columnal plates. These crinoids show a typical combination of a preburial decay on the sea bottom and a post-burial decay within the sediments. 9 Regional encrinites of the La Joya Formation are the result of in situ accumulations rapidly reworked by storm waves and resedimented as distal tempestites. 9 These distal tempestites were finally accumulated near chaetetid bioconstructions developed in another part of the platform. Between the crinoid biotopes and these constructions some infaunal organisms probably inhabited the micritic substrate. The high degree of silicification of the crinoids indicates that they were possibly associated with siliceous organisms (Porifera?), not preserved in the assemblages. The thanatocoenoses suggest a strong affinity with mid-continental and southern regions in the USA, especially with Kansas and Texas. The seaways of Sonora, southern USA and the midcontinent were apparently well connected during the Atokan, as the ecological conditions were remarkably similar. The Evaluation-orientation de la Coop6ration Scientifique, Asociaci6n Nacional de Instituciones de Educaci6n Superior, Consejo Nacional de Ciencia y Tecnologia, Universidad Nacional Autonoma de Mexico project No. MOOU01 'Un estudio sedimentol6gico, micropalentol6gico y geoquimico del Paleozoico de M6xico', and UNAM-PAPIIT project No. IN104103-3 'Bioestratigrafia de rocas de plataforma del Pensilvfinico y P~rmico de Sonora, Mdxico', financially supported this research. We are grateful to L. Pille for her technical help. We thank the reviewers T. Kammer, E. Gluchowski and M. Aretz for their constructive remarks and corrections. References ALMAZAN-VAZQUEZ, E., BUITRON-SANCHEZ, B. E., VACHARD, D., MENDOZA-MADERA, C. ~ GOMEZESPINOSA, C. 2007. The late Atokan (Moscovian, Pennsylvanian) chaetetid accumulations of Sierra Agua Verde, Sonora (NW Mexico): composition, facies and palaeoenvironmental signals. In: ALVARO, J. J., ARETZ, M., BOULVAIN,F., MUNNECKE, A., VACHARD,D. & VENNIN, E. 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