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Boletín de la Sociedad Geológica Mexicana ISSN: 1405-3322 Sociedad Geológica Mexicana, A.C.

Torres-Martínez, Miguel A.; Villanueva-Olea, Rafael; Sour-Tovar, Francisco Columnar ossicles of Permian crinoids, including two new genera, from the Grupera Formation (Asselian-Sakmarian) of Chiapas, Mexico Boletín de la Sociedad Geológica Mexicana, vol. 72, no. 2, 00005, 2020, May-August Sociedad Geológica Mexicana, A.C.

DOI: https://doi.org/10.18268/BSGM2020v72n2a280819

Available in: https://www.redalyc.org/articulo.oa?id=94366150005

How to cite Complete issue Scientific Information System Redalyc More information about this article Network of Scientific Journals from Latin America and the Caribbean, Spain and Journal's webpage in redalyc.org Portugal Project academic non-profit, developed under the open access initiative This is an open access article under the CCBY-NC-SAThis Universidad NacionalAutónoma de México. Peer Reviewing under the responsibility of Manuscript October29,2019 accepted: Corrected manuscript received: October15,2019 Manuscript received: June 27,2019 BSGM2020v72n2a280819 http://dx.doi.org/10.18268/A280819. de la Sociedad Geológica Mexicana, 72 (2), (Asselian‒ Formation Chiapas, Mexico: Boletín Grupera Sakmarian) of the from Permian crinoids, includingtwo newgenera, Sour-Tovar, F., 2020, Columnar of ossicles Torres-Martínez, M. A., Villanueva-Olea R., How tocitethisarticle: [email protected] Corresponding author: (M.A.Torres-Martínez) * 3000, Coyoacán, 04510,Mexico City, Mexico. ico, Circuito Interior, Avenida Universidad cias. Universidad Nacional Autónoma de Méx 3 ico City, Mexico. Coyoacán, Nacional Mex3000, Universidad04510, da Universidad Ciencias. Autónoma de México, Circuito Interior, Aveni de ultad 2 Mexico City, Mexico. Avenida Universidad 3000, Coyoacán, 04510, México, Circuito de la Investigación Científica, Geología. Universidad Nacional Autónoma de 1 Miguel A. (Asseliano–Sakmariano) deChiapas,México Osículos columnaresdecrinoideospérmicos,incluidosdosgénerosnuevos,laFormaciónGrupera Formation ( Column (https://creativecommons.org/licenses/by-nc-sa/4.0/) Departamento de Paleontología,Departamento Instituto de Museo dePaleontología,Museo Facultad de Cien Departamento de Biología Evolutiva,Departamento Fac ar ossiclesofPermiancrinoids,includingtwonewgenera,fromtheGrupera Torres-Martínez Asselian–Sakmarian) ofChiapas,Mexico 1,* , Rafael - - - - - Sakmarian, Grandian Province. locion, Cyclogrupera Keywords: Parataxa, paleoprovince. same the of that part be should suggested crinoids is it Venezuela), and in Palmarito and Sonora Mexico; in Coahuila, Chiapas USA; in America (Texas andNew Mexico North to the Grandian Province of related been had that unit Permian a Because the specimens were found in region. Chicomuselo the from noids cri Permian on work formal first the inChiapas,is the first Mexico, and all parataxa studied recordThe of the lower Cisuralian (lower Permian). to Carboniferous) (Upper sylvanian insaturatus erathenseLamprosterigma caudiculus regularis range of stratigraphical The ramp. inner the of waters restricted paleoenvironmentdepositional to Preserved elements allow relating the morphospecies. and morphogenera minor grupera Floricyclocion heteromorpha described. are (Asselian‒Sakmarian) lected from the Grupera Formation Permiancol crinoids of ossicles columnar Elevenof morphotypes ABSTRACT Villanueva-Olea Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín is extended is from the Penn are proposed asnew , Heterostelechus keithi, keithi, Heterostelechus 2 and , Francisco and , Asselian‒ , Cyclocaudex Floricyc Cyclo Cyclo Sour-Tovar ------minor Floricyclocion (Asseliano‒Sakmariano). Grupera mación de la Forcolumnares de crinoideos pérmicos Se describen 11 morfotipos de osículos RESUMEN cia Grandiana. Provin Asseliano‒Sakmariano, Floricyclocion Palabras clave: Parataxones, marían partedeestamismapaleoprovincia. for crinoideos Venezuela),los que sugiere se Sonora y Chiapas en México; y Palmarito en (Texas México y en EUA;Nuevo Coahuila, con la Provincia Grandiana de Norteamérica que ha sido relacionada una unidad pérmica en encontrados fueron ejemplares los que a Debido Chicomuselo. de región la de micos sobre crinoideos pér primer trabajo formal el es éste que de además México, Chiapas, por primerataxones se registran vez para Cisuraliano para Todoslos inferior). al (Pérmico inferior Superior) (Carbonífero insaturatus Lamprosterigma caudiculus de estratigráfico alcance El interna. restringidas de la rampacon aguas depósito relacionar el paloambiente de permitieron conservados elementos Los morfoespecies. y se proponen como nuevos morfogéneros se proponen comonuevos 3

e xine e Pensilvánico del extiende se /72(2)2020 regularis

heteromorpha

erathense , , Heterostelechus Cyclogrupera y y Cyclogrupera Cyclocaudex

Cyclo keithi 1 , ------,

Permian crinoids from the Grupera Formation, Chiapas, Mexico ABSTRACT INTRODUCTION / GEOLOGICAL Permian crinoids from the Grupera Formation, Chiapas, Mexico SETTING 1. Introduction 2 2 is common and diverse in Phanerozoic rocks, the rocks,Phanerozoic in diverse and common is crinoids of record fossil the Although shallowPaleozoic. of invertebrates marine environments, especially duringthe of group important most the of one considered been have Crinoids tion of Moore and Jeffords (1968) and given that given and (1968) Jeffords and Moore of tion (Thomka research of area neglected, but important, an as recognized been has material incomplete such of (Hess paleontology crinoid in tasks essential are ossicles disarticulated or isolated about works all context, this In useful. paleobiogeographic or biostratigraphic be could fossils these that possibility the suggesting morph, each of distribution based spatial and is temporal the on scheme a Such records. Paleozoic on mainly working ossicles, the of morphology on the based 2019) system classification Doyle, a proposed and have Donovan 2015; Donovan, ( and Doyle, 2019). essentially ignored (Thomka elements to low taxonomic levels whereby they are these identify accurately to difficult it makes cess isolated pro ossicles.This of finding common the crinoids to occur quickly, resulting disjoining of in processes (Hess various by disarticulated are remains decay, fossilcrinoidal andthe great majority of quickly tissues connective these death Upon ible. flex or rigid either tissue,connective ligamentary of means by adjoined composition ic-magnesian calcit are platesof orossicles aseriesof parts of composed These articulated. are body the pose which the peduncle, calyxandcrown that com thewayin the complete resultis specimens of of or localized lenses (Hess are typically confined to particular bedding planes articulatedare crinoids exceptional, andthey of assemblages particular, In specimens. articulated of reports the than greater is skeletons make their up that ossicles disarticulated of abundance / e.g. Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín osdrn ti polm svrl researchers several problem, this Considering , Moore and Jeffords, 1968; Fearnhead and Fearnhead 1968; Jeffords, and Moore , et al et al , 02. hs ass h post-mortem the causes This 2002). ., ., 2012). According to the classifica the to According 2012). ., et al ., 2002), because the value the because 2002), ., et al. et et al , 2002). The scarcity The 2002). , http://dx.doi.org/10.18268/BSGM2020v72n2a280819 ., 2012; Donovan2012; ., - - - - - /72(2)2020 2. Geologicalsetting pled, is about 15° 43’ 14.2” N ‒ 92° 26’ 46” W, in locality,The where the material studied wassam the material studied of in Chiapas. the Permian of biostratigraphic and paleogeographic implications possible the discuss to is aim second A crinoids. Paleozoic of elements isolated of distribution the ian). This information expands the current data on (Asselian‒Sakmar age Cisuralian early of unit a from columnarossicles the Grupera Formation, of series a describe to aims paper this Chiapas,from there are no previous studies on Permian crinoids óe-ao, 99. h Plooc succession Paleozoic The 1956; 1979). López-Ramos, (Gutiérrez-Gil, Formation Hondo Paso the of base the be could Limestone Vainilla La the that suggested been has and unclear remains position stratigraphic however,actual mation; its limestone andargillaceous limestone. of continues with LaVainilla Limestone, composed succession The 1956). (Gutiérrez-Gil, shale and limestone gray dark of alternations of m 110 by shale andsandstone strata, gray followed 117 mof calated with dark gray limestone; above, there are inter shale silicified of m 300 of up made is base gerina gruperaensis Miller (1944) the byforaminifera means of and Thompson by assigned age (Cisuralian), ian the Asselian‒Sakmar the is from Formation Formation Grupera Rosa Santa the Overlaying 1956). (Gutiérrez-Gil, slate and phyllite shale, composed of mainly is unit This Pennsylvanian). Mississippian‒Middle (Upper Formation Rosa to the Guatemala frontier (Gutiérrez-Gil, 1956). Chiapas extending from LomaAlta andLaPinta iswell-exposed formation insoutheastern This region. Chicomuselo the from unit stratigraphic Formation, considered the oldest Permian litho Grupera the in grain fine of sandstones careous cal in deposited were specimens The 1). (Figure the Chicomuselo municipality, Chiapas, Mexico This unit apparentlyThis overlays the Grupera For The Paleozoic succession begins with the Santa and Schwagerina chiapasensis Schwagerina Article A280819 Schwa Its . ------Article A280819 3. Systematicpaleontology al. Figure 1 (2005). cases rubber castswere made. in calcareous sandstone as authigenic molds. In all by the prefix FCMP.The specimens are preserved aretype specimens designated in the descriptions sidad Nacional Autónoma de México. Figured and the Facultad de Ciencias, Univer Paleontología of de Museo at deposited is described material The 2019) (Figure 2). Torres-Martínez1973; Hernández-García, 1956; (Gutiérrez-Gil, Artinskian‒Roadianage of limestones gray dark to light com of m 600 of posed mainly Guadalupian) Cisuralian‒lower ends with the Paso HondoFormation (upper Geologicalposition ofthelocality,where crinoids occur,intheGrupera Formation.Mapmodified from Jiménez-Hernández Subclass andOrder UncertainMoore and Class CrinoideaMiller, 1821 http://dx.doi.org/10.18268/BSGM2020v72n2a280819 Boletín de la Sociedad Geológica Mexicana Mexicana Geológica Sociedad la de Boletín et al ., - - nady et uv, h inr ns abutting ends inner the curve, bent inwardly straight crenulae, their outer ends aligned in articular pores;medium-length facets having a star-shaped belt of radial of development no with pitswider than high sutures indented forming size; in columnals,alternating 14 with internodals tastellate stem; nodals with 5 elliptical scars; cirrus Diagnosis. and Jeffords, 1968. Type species. Family Pentacauliscidae Moore and Jeffords, 1968 moderately large andcircular. concave lumen slightly floor; a with areolastellate Genus [Group Pentameri] Moore andJeffords, 1968 Pentagonopternix Heteromorphic andtransversely Heteromorphic pen Pentagonopternix Figure 3Aand3B

Pentagonopternix Jeffords, 1968 Moore andJeffords, 1968 / 72(2)2020 ? sp.

insculptus Moore 3 3 et -

GEOLOGICAL SETTING / Permian crinoids from the Grupera Formation, Chiapas, Mexico SYSTEMATIC PALEONTOLOGY Permian crinoids from the Grupera Formation, Chiapas, Mexico SYSTEMATIC PALEONTOLOGY Figure 2 4 4 Permian (Roadian‒Wordian) of the Transbaikal the (Roadian‒Wordian)of Permian Jeffords, and HorizonfromAntiinskiy the (Moorein middle 1968), the Texas of age sylvanian) Penn (Upper Gzhelian from Limestone Chaffin does notallow makingareliable assignment. noditaxis, a of lack the as well as facet, articular ossicle. the several the features of poorconservationThe of of shape stellate the to due mainly to themorphogenus assigned Remarks. ian), Chiapas, Mexico. (Asselian‒Sakmar Formation Grupera stones, Occurrence. Material. pentagonal lumen. sub small, preserved; poorly crenularium shape; Description. / Paleozoic stratigraphy oftheChicomuseloregion Paleozoic Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín hs opoeu hs en on i the in found been has morphogenus This h seies r questionably are specimens The FCMP 1349,1350. Fine-grained calcareous sand Fine-grained Columnals with a pentastellate http://dx.doi.org/10.18268/BSGM2020v72n2a280819 Pentagonopternix . - - - - - /72(2)2020

ls wls atclr ae crua; crenularium circular; facet articular walls; ulose gran profile; longitudinal in rounded eter,gently Diagnosis. unit where thespecimens were collected. articular facet, aswell as to the lithostratigraphic Etymology. Type species. Sonora (Buitrón-Sánchez Verde,Agua Sierra of Pennsylvanian Middle the in recorded been only had it Mexico, In 2016). Verkhoyansk, Russia(Biakov of (Lopingian) Imtachan Formation from the upper Permian Russia(Kotlyar region of Family Cyclomischidae Moore andJeffords, 1968 [Group Cyclici] Moore andJeffords, 1968 Hmoopi tmo small diam stem of Homeomorphic Genus Rfr otecrua hp fthe Refersto the circular shapeof

Cyclogrupera minor Cyclogrupera et al et al ., 2015). n.gen. ., 2006), and in the in and 2006), ., n.sp. Article A280819 et al ., - - Article A280819 lumen surrounding the circular lumen. peri a of presence and crenularium broad stem, duetothe slender to thefamilyLeptocarphiidae Remarks. Chiapas, Mexico. Formationstones,Grupera (Asselian‒Sakmarian), Occurrence. Material. and crenularium 0.7mmwidth. diameter crenulae,mm 13 displays0.5 this lumen Description. Diagnosis. specimens. Etymology. small, lumen outline; circular. lumen the marked, delineating gently perilumen areola; without size; the same coarse, withcrenellae and culminaof crenulaeperilumen; of large,outline reachingthe outline. in quinquelobate or subpentagonal circular, and size, in medium to small lumen perilumen; a ing becom and or lacking small, areola crenulae; straight long with wide crenularium taller; slightly and scars, notwider than internodals ing cirrus bear nodals convexprofile; gently longitudinal to Diagnosis. fords, 1968. Type species. thearticularfacet. from thesurfaceof and ring-shaped perilumen surroundingthe lumen, Nothrosterigma ulus Genus Cyclocaudex typicus differs from differs Heterostelechus Cyclocaudex FCMP1351. The Chiapasmaterial The assigned was tm eeoopi; straight-sided heteromorphic; Stem Asthegenus. differs by its coarser, convex and convex coarser, its by differs Rfr otetn ieo the Refers to the tiny size of Cyclogrupera minor Fine-grained calcareous sand Fine-grained ih ris f te eu; besides genus; the of traits With

Figure 3Cand3D Cyclocaudex typicus Cyclogrupera Figure 3Eto 3N displaysan elevated perilumen Moore andJeffords, 1968 Moore andJeffords, 1968 by the lack of areola; of lack the by n.sp. http://dx.doi.org/10.18268/BSGM2020v72n2a280819 Moore and Jef Boletín de la Sociedad Geológica Mexicana Mexicana Geológica Sociedad la de Boletín Cyclocaudic ------mation resemble mation Remarks. Chiapas, Mexico. Formationstones,Grupera (Asselian‒Sakmarian), Occurrence. Material. is observed. lumen; circular the lumen, acircular and minute jugula a large, inside of surrounds which flat, not conspicuous, reduced in size, and granulose is areola bifurcated; many crenulae, broad,slender with crenularium subcircular; crenulate; facet articular gently sutures stem; heteromorphic rounded longitudinal profile, withgranulose walls; Description. Material. to circular inoutline. irregular size, reduced of lumen flat; and narrow areola margins; near bifurcating crenellae, than wider culmina crenulae; narrow with wide very crenularium columnals; four of noditaxis imum least three different heights, which a min suggests at crenulate;suturesslightly nodals; the of ossicles cirrus,conspicuous which endow a greater size to Description. NorthAmerica. lower Permian of the in taxon the of record first the is This 1997). Mexico Oaxaca in (Vachard dle Permian of mid and 1996) (Esquivel-Macías, Puebla of nian (Villanueva-Olea Oaxaca of Carboniferous the in as well as 1968), Jeffords, and (Moore States of United the Pennsylvanian in Texas Upper the in reported been surface. depressed a with Cyclocaudex jucundus Lomalegnum culmina typical of nor the peripheral ends of C. typicus h mtra cn lo e itnuse from distinguished be also can material The Cyclocaudex plenus hasneither the sutures strongly crenulate FCMP1352-1357. FCMP1358-1360. . Teseieso the Grupera For of specimens The Fine-grained calcareous sand Fine-grained Columnwithstraight to slightly Ossicles circular of shape in outline; Figure 3Oto3S Lomalegnum hormidium. Lomalegnum t al. et by the lack of a broad areola broad a of lack the by Moore andJeffords, 1968 21) Upr Pennsylva Upper 2011), , / 72(2)2020 Cyclocaudex typicus However, et al. et has 5 5 - - - - - ,

Permian crinoids from the Grupera Formation, Chiapas, Mexico SYSTEMATIC PALEONTOLOGY Permian crinoids from the Grupera Formation, Chiapas, Mexico SYSTEMATIC PALEONTOLOGY 6 6 1968) by its symmetric articular facet and suture and facet articular symmetric its by 1968) TexasGraham Formation (Mooreof and Jeffords, guished from Remarks. Chiapas, Mexico. Formationstones,Grupera (Asselian‒Sakmarian), Occurrence. idePrino Sonora(Buitrón middle Permian of et al. Upper PennsylvanianSonora (Buitrón-Sánchez of Pemuxco, Hidalgo(Arellano Salvador Patlanoaya, Puebla, the lower Permian of Buitrón, (Esquivel Permian and 1992) and León de (Velasco Pennsylvanian (Buitrón Hidalgo Calnali, of Pennsylvanian the to addition in 1968), fords, TexasUpper Pennsylvanian (Moore and Jef of Remarks. Chiapas, Mexico. Formationstones,Grupera (Asselian‒Sakmarian), Occurrence. Material. thelumen. towards theinteriorof deepens which depressed, broad, areola circular; and large lumen crenellae; than wider with culmina narrow crenulae half; outer the bifur at some cations with size, medium of crenularium circular; facet articular crenulate; sutures height; w columnals convex; and Description. (Arellano (Moreno and Patiño 1981) and Pemuxco, Hidalgo Calnali of Permian lower the in and 1968) fords, TexasUpper Pennsylvanian (Moore and Jef of areola. the of edge the near bifurcated crenulae the and boundary crenularium, the and areola reduced defined the between its poorly by 1968) Jeffords, Texasthe Upper Pennsylvanian (Moore and of Cyclocaudex typicus strongly crenulate. / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Cyclocaudex jucundus , 2012; Villanueva-Olea Cyclocaudex plenus et al Temtra fChiapas is distin material The of

FCMP 1361. Cyclocaudex jucundus ., 1998; Buitrón-Sánchez Fine-grained calcareous sand Fine-grained Ltso columnalsslightlyrounded Latus of Mooreanteris waylandensis Figure 3Tand3U Fine-grained calcareous sand Fine-grained fo h hit omto f from Formation the Thrifty of Cyclocaudex plenus Moore andJeffords, 1968 has been reported in the in reported been has ith slight differences in differences slight ith et al et al et al ., 1987), the Upper the 1987), ., et al., et ., 2016) and in the http://dx.doi.org/10.18268/BSGM2020v72n2a280819 ., 1998), Middle– 1998), ., occurs inthe occurs is dissimilar to dissimilar is 2000) of San of 2000) et al. et al. from the , 2007a). , 2017). ------/72(2)2020 ecig h rm f te lumen. the of rim the reaching Cyclocaudex plenus Remarks. Chiapas, Mexico. Formationstones,Grupera (Asselian‒Sakmarian), Occurrence. Material. mediumsize andcircular. lumen of part; medium the in bifurcating some crenellae, lumen; the crenulae very narrow, withculminawider than of rim the reaching wide, very ium Description. Etymology. sized stellate lumen. medium and areola flat crenulae, length median stem columnals with straight and heteromorphic Diagnosis. Etymology. Type species. from thePermian. al., (Villanueva-Olea Oaxaca Ixtaltepec, Santiago of Pennsylvanian Lower–Middle and sissippian (Buitrón-Sánchez Calnali, Hidalgo as well as inthe Pennsylvanian of ( Kansas sylvanian of Cyclocaudex insaturatus edges. rounded inner without crenularium, flat its by 1968) Jeffords, and (Moore Kentucky of pian from differs species The lumen. larger a and crenulae less abundant and coarser its by and 1968) (Moore Jeffords, Kentucky of Mississippian Lower uratus Cyclocaudex insaturatus Family Moore Floricyclidae andJeffords, 1968 21) hsi h is eodo of record first the is This 2011). is dissimilar to isdissimilar Cyclocaudex congregalis Floricyclocion heteromorpha FCMP1362. The specimen of Chiapas differs from Chiapasdiffers specimenof The Genus lrccia caatrzd y a by characterized Floricyclidae Refers stem. totheheteromorph Refers lumen. tothefloriform riua fct iclr crenular circular; facet Articular Floricyclocion heteromorpha Figure 4Aand4B by its larger lumen and crenulae and lumen larger its by Fine-grained calcareous sand Fine-grained et al., Floricyclocion Figure 3V occurs inthe Middle Pennoccurs Moore and Jeffords, 1968), Jeffords, and Moore Moore andJeffords, 1968 1987) and the Middle Mis Cyclocaudex aptus o the Lower Mississip of n.gen. Cyclocaudexinsat n.sp. C. insaturatus C. Article A280819 n.sp. from the et ------Article A280819 tagtlniuia rfl fthestem straight longitudinalprofile of a of presence the and lumen floriform its by dae Floricyclocion crenulae. some of division with perilumen, ulose gran and marked very its by morphogenus new the Likewise, crenulae. areola, in addition to very slender and numerous the genus hand, other the On keeled. strongly nodals and crenulate strongly sutures lumen; larger and guished from Remarks. Chiapas, Mexico. Formationstones,Grupera (Asselian‒Sakmarian), Occurrence. Material. lumenmediumsizedand granulose; andstellate. flatareola crenellae; than coarser slightly culmina crenulae; coarse and straight with of different size, medium crenularium circular; of facet articular columnals length; with heteromorphic, Description. Diagnosis. lae; areola fromgranulose, crenulariumlae; off the set crenelthan wider culmina periphery; the to close bifurcating some crenulae, slender with wide ium Description. perilumen. narrow a by it from separated or lumen, broad pentalobate the of edge the smooth to extending surface, with bowl-shaped areola edge; sharp a by areola the from off set straight, crenulae ium; crenular broad moderately with facet articular indented; not noncrenulate, sutures convex; alit tle or straight profile longitudinal columnals; Diagnosis. Jeffords, 1968. Type species. Lamprosterigma erathense Genus Lamprosterigma Lamprosterigma FCMP 1363. isassignedto the family Floricycli oemrhc tm low-height stem; Homeomorphic Asthegenus. The morphogenus morphogenus The Floricyclocion Fine-grained calcareous sand Fine-grained riua fct iclr crenular circular; facet Articular Columnstraight in lateral view,

Lamprosterigma mirificum Figure 4C displaysthe in a depression Moore andJeffords, 1968 Moore andJeffords, 1968 Floricyclus by its very short crenulae http://dx.doi.org/10.18268/BSGM2020v72n2a280819 Cyclocion differs from the from differs Boletín de la Sociedad Geológica Mexicana Mexicana Geológica Sociedad la de Boletín Moore and . is distin is ------crenulae, and strongly floriform perilumen, with perilumen, floriform strongly and crenulae, phogenus (Moore and Jeffords, 1968) is dissimilar to the mor Plummeranteris mirificum howeverJeffords, 1968) and (Moore Kansas of mirificum Lamprosterigma Remarks. Chiapas, Mexico. Formationstones,Grupera (Asselian‒Sakmarian), Occurrence. Material. oped petals. devel poorly with floriform, wide, lumen lumen; peri discrete a into grades and depression a by wider than crenellae; areola wide and granulose granulose and wide areola crenellae; than wider culmina crenulae; coarse and crenularium slender gitudinal profile; articular facet circular with a very Description. claustra. and jugularramparts inthe center of columnalplates, jugulum claustra at mid-height of with lumen areola; the of edge accentuated an or perilumen narrow a by bordered lumen, circular vermiculate or granulose irregularities; large of and marks with convex, or concave slightly areola, flat medium to coarse crenulae set off from a broad and narrow; crenularium profile; longitudinal rounded width; column largest the than narrower slightly Diagnosis. Jeffords, 1968 Type species. Family Moore Leptocarphiidae and Jeffords, 1968 erathense al. et (Buitrón-Sánchez Sonora of (Atokan) sylvanian Middle Penn the Jeffords, and and 1968) (Moore Texasreported fromthe MiddlePennsylvanian of petals. well-developed Preptopremnum Genus 20b. hs s h frt curne of occurrence first the is This 2007b). , inPermian rocks. does not display a developed perilumen. developed a display not does Preptopremnum Lamprosterigma The specimen wascompared The with FCMP 1364. Medium-sized column; articular facets fo h enyvna fTexas from the Pennsylvanian of Fine-grained calcareous sand Fine-grained Column with aslightly rounded lon cf.

Preptopremnum rugosum Figure 4Dto4H laeve Moore andJeffords, 1968 Moore andJeffords, 1968 Mdl Pennsylvanian) (Middle by its coarse and straight and coarse its by apotrga erathense Lamprosterigma / 72(2)2020 Moore and 7 7 L. L. is ------

Permian crinoids from the Grupera Formation, Chiapas, Mexico SYSTEMATIC PALEONTOLOGY Permian crinoids from the Grupera Formation, Chiapas, Mexico SYSTEMATIC PALEONTOLOGY mm; C,D=1mm. (V) 1361. 1353. (O-S) 1351. (E-N) Figure 3 8 8 / (A,B) Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Cyclocaudex insaturatus Cyclocaudex typicus Cyclocaudex plenus Pentagonopternix , samples O, P) FCMP 1359; Q) FCMP 1360; R, S) FCMP1358.(T,U) , samplesO,P)FCMP1359;Q) 1360;R,S) , samplesE)FCMP1354;F,G) 1357;H,I) FCMP1352;J,K) FCMP1355;L)1356;M,N) FCMP ? sp.,samplesA)FCMP1350;B) FCMP1349.(C,D) , sample FCMP 1362. All specimens illustrated are rubber casts. Scale bars = 5 mm, except A, B, L = 2 = L B, A, except mm, 5 = bars Scale rubbercasts. are illustrated specimens All 1362. FCMP sample , http://dx.doi.org/10.18268/BSGM2020v72n2a280819 /72(2)2020 Cyclogrupera minor Cyclocaudex jucundus n.genand n.sp.,sampleFCMP , sampleFCMP Article A280819 Article A280819 California, Mexico (Navas-ParejoCalifornia, Baja Pintas, Las Sierra of Permian vanian–lower Pennsyl Middle the in and 2015) 2012; 2007b, Sonora (Buitrón-Sánchez sylvanian of (Moore and Jeffords, 1968), as well as in the Penn States United the in Texas of Pennsylvanian dle columnals. the of latus the on granules occurrence the of quantity great of of because (1968) Jeffords and Moore by described those to different slightly are Nevertheless,specimens lumen. largeChiapas the and rounded medium-sized areola, and acircular from Remarks. Chiapas, Mexico. Formationstones,Grupera (Asselian‒Sakmarian), Occurrence. Material. resembling abroad perilumen; lumen circular. in the area surrounding lumen, slightly elevated, Occurrence. Material. onal lumen. areola size; same medium-sizedsubpentag elevated granulose; and the of crenella and culmina crenulae; coarse with crenularium medium-sized circular; facet articular crenulate; sutures 2IN); the height in 1INand columnals(nodalsandinternodals of sizes different three of noditaxis Description. jeffordsi Heterostelechus cular lumen,alsoclassifiable asaperilumen. cir around tract elevatedconcave, areola length; medium of crenulae straight coarse wider; much not but internodals, than taller are which nodals, and indented c profile; longitudinal in rounded Diagnosis. euae uue; n o mr cirru more or one sutures; renulate Genus P. laeve Heterostelechus FCMP1369-1373.

FCMP 1365-1368. Preptopremnumlaeve tm eeoopi; columnals heteromorphic; Stem Preptopremnumrugosum by the longer crenulae, a granulose granulose a crenulae, longer the by Fine-grained calcareous sand Fine-grained Fine-grained calcareous sandstones, Fine-grained oun ih srih profile; straight a with Column Figure 4Ito4N Jeffords, 1968) ilr 16 (n or and Moore (in 1968 Miller, Moore andJeffords, 1968 occurs inthe Mid occurs http://dx.doi.org/10.18268/BSGM2020v72n2a280819 is distinguished is et al., Boletín de la Sociedad Geológica Mexicana Mexicana Geológica Sociedad la de Boletín et al 2018). s scars in s scars ., 2005, ------Heterostelechus in theearly Permian. 2004). This is the the first morphospecies report of and Buitrón-Sánchez, 1982; Esquivel-Macías fords, 1968) and Guerrero, Mexico (Flores de Dios Texasthe Late Pennsylvanian(Moore andJef of recordedin been taxonhas rounded. This slightly the mainly morphospecies by its column profile of jeffordsi Heterostelechus to the morphospecies by the articular facet features. Remarks. apas, Mexico. Chi (Asselian‒Sakmarian), Formation Grupera file; nodals bearing one single scar; crenularium scar; single one bearing nodals file; Diagnosis. Jeffords, 1968. Type species. NorthAmerica. the Permian of al. (Buitrón-Sánchez Mexico in Sonora and 1992) Buitrón-Sánchez, and León de (Velasco Puebla addition to Hidalgo (Buitrón-Sánchez in TexasJeffords, 1968), and (Moore of sylvanian Heterostelechuskeithi lumen does not allow making a reliable assignment. keithi Remarks. Chiapas, Mexico. Formationstones,Grupera (Asselian‒Sakmarian), Occurrence. Material. and circular. small lumen crenularium; the of edge inner the slightly concave, apentagonal outline in forming areola width; same the of culmina and crenellae with of crenulae, straight and crenularium coarse size; medium circular; facet articular latus; Description. Genus , 2007b, 2008, 2015). This is the first report in report first the is This 2015). 2008, 2007b, , but its straight longitudinal profile and smaller Cyclocaudiculus Temtra fChiapaswasassigned material The of The material resembles material The FCMP 1374,1375. Stem with a straight longitudinal pro cf. Fine-grained calcareous sand Fine-grained Plate with asmoothandstraight

Figure 4Oand4P Cyclocaudiculus regularis keithi Jeffords, 1968) has been recorded in the Pennthe recordedin been has is distinguishedfrom the rest Miller, 1968 (in Moore and Moore (in Miller,1968 Moore andJeffords, 1968 / 72(2)2020 Heterostelechus et al., Moore and 1987), et al. 9 9 et - - - - - ,

Permian crinoids from the Grupera Formation, Chiapas, Mexico SYSTEMATIC PALEONTOLOGY SYSTEMATIC PALEONTOLOGY / Permian crinoids from the Grupera Formation, Chiapas, Mexico DISCUSSION 4.1. DEPOSITIONAL PALEOENVIRONMENT 4. Discussion 10 10 Description. small ormedium. circular,lumen crenulae; straight and coarse with the author focused on a group of sedimentary sedimentary the author focused on agroup of because shallow extremely is interpretation This waves. the from protected oxygenated, and nated low energy, Grupera unit well-illumi asshallow waters of the of paleoenvironment the described tion isthat done by (1973),who Hernández-García the Grupera Forma anenvironment of tation of differences.lithological previous only The interpre underlays the Paso HondoFormation, having clear et al (Torres-Martínezdescribed were ramp homoclinal carbonated a whichfeaturesin the of included was (Artinskian‒Roadian) Formation Hondo Paso from the succession carbonate one of study the only Previously, Formation. Grupera the scarce, for especially very general, in been, has region muselo the Chico Paleozoic environments of studyThe of Permian is the This Mesteñas, (Villanueva-Olea Las Mexico Sierra Sonora, of Pennsylvanian the in and Jeffords, 1968) and (Moore States United the in Texas of Pennsylvanian) (Upper Gzhelian the Remarks. Chiapas, Mexico. Formationstones,Grupera (Asselian‒Sakmarian), Occurrence. Material. and subcircular. small crenularium perilumen; with flat areola lae; crenel than wider culmina crenulae; coarse with crenularium columnal; the of width the occupies which circular facet articular profile; longitudinal / Cyclocaudiculus regularis Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín , 07. oehls, h Guea Formation Grupera the Nonetheless, 2017). ., . FCMP 1376-1379. first record of the taxon in the lower the in taxon the of record first Cyclocaudiculus regularis Fine-grained calcareous sand Fine-grained tm eeoopi, tagt in straight heteromorphic, Stem Figure 4Qto4V Moore andJeffords, 1968 http://dx.doi.org/10.18268/BSGM2020v72n2a280819 is reportedis in et al , 2016). ., ------/72(2)2020 AGE 4.2. PALEOBIOGEOGRAPHY ANDIMPLICATIONS IN the contrary, the occurrence of microbial growth, microbial of occurrence contrary,the the on waters; open with nearness suggests algae loid matrix. fine-grained the ironin wellas oxides aredispersed as pyrite and crystals dolomite Euhedral observed. loides the genera foraminifera of crinoids lated whereas of phylloid algae and benthic articu pleuricolumnals and ossicles abundant by represented mainly are grains skeletal The cement. calcareous and matrix clay-rich by characterized is which grain, fine of sandstone calcareous in ded changes through the unit. features without considering the lithological al. (Yancey,ments Shen 2004; Shi, and Shen 1975; environ subtropical and tropical of brachiopods and cephalopods fusulinids, of taxa numerous by occurred in the Grandian Province, characterized ico (Torres-Martínez south-central United States the Mex and the south of linked which sea epicontinental an through that, established has during this period, there was a marine connection faunas of similarity The New Mexico. and Texas from localities different in described those with resemblance great a have early‒middle Permian southern Chiapas faunas of al. Torres-Martínez 1944; Miller, and Thompson In previous studies (Müllerried possible by produced was association the that chance a is there Nonetheless, 2010). (Flügel, transport longer without settings quiet-water in burial indicates nals associated with well-preserved crinoidpleuricolum crinoidfragments disarticulated, the poorsorting of mostly are studied ossicles crinoid Although ramp. the inner the environment to restricted waters of suggests crinoids high productivity inthe euphotic zone, relating of abundance The zone. tidal clay and sands indicates a proximity with the inter , 2013). the that out pointed been has it 2019), 2016, , The presence of benthic foraminifera and phyland foraminifera benthic of presence The context,this parataxaIn theare studied embed are rare. Scattered microbial components are components microbial Scatteredrare. are scavenging crinoidremains. dispersion of et al. et , 2016). This connection This 2016). , Geinitzina and et al Article A280819 Nodosinel , 1941; ., et et ------Article A280819 FCMP 1369;J)1370;K) 1373;L, M)FCMP1370;N) FCMP1372.(O,P) (D-H) Figure 4 except C =5mm;G,H1mm. except (Q-V) 1375. FCMP Preptopremnum (A, B) (A, Floricyclocion heteromorpha Cyclocaudiculus cf. laeve , samplesD,E)FCMP1365;F) 1368;G,H)FCMP1366.(I-N)

regularis http://dx.doi.org/10.18268/BSGM2020v72n2a280819 , samples Q) FCMP 1377; R, S) FCMP 1376; T) FCMP 1378; U, V) FCMP 1379. Scale bars = 2 mm, 2 = bars Scale 1379. FCMP V) U, 1378; FCMP T) 1376; FCMP S) R, 1377; FCMP Q) samples , n. gen and n. sp., sample FCMP 1363. (C) 1363. FCMP sample sp., gen andn. n. Boletín de la Sociedad Geológica Mexicana Mexicana Geológica Sociedad la de Boletín Heterostelechus Lamprosterigma erathense, cf. Heterostelechus jeffordsi keithi / 72(2)2020 , samplesO)FCMP1371;P) sample FCMP 1364. FCMP sample , samplesI) 11 11

Permian crinoids from the Grupera Formation, Chiapas, Mexico DISCUSSION Permian crinoids from the Grupera Formation, Chiapas, Mexico DISCUSSION / CONCLUSIONS 12 12 lo ulse dsrpin o ema material Permian of descriptions published also some taxa. Nonetheless, Russian work groups have withpreviousthe only country Permian records of is Mexico States, United the of localities vanian) (Middle‒Late Gzhelian or Moscovian in highlighting that all known taxa had been reported Regarding columnar ossicles herein described and a kon rm h Lwr issipa o of Mississippian Lower Oaxaca and Oaxaca to the Pennsylvanian of the from known was Oaxaca. of Permian the range fromgraphic the Lowerto Mississippian a very broad temporal distribution. In thismanner,observed itis that, sometaxa have detailed. been have reference bibliographic their and occurrences the 1, Table In Texas. of nian the Pennsylva them have recordslocalities of in of all and Mexico of Permian or Carboniferous the of localities diverse to referred been havealready work this in described are that crinoids of from Venezuela. Palmarito and States United the Texasfrom and Mexico New Mexico, from Chiapas and Sonora to the Grandian Paleoprovince, located related in Coahuila, localities Permian in found be might crinoids of forms common that hypothesized be can it fusulinids, or brachiopods as such fauna, the associated considering the composition of Texas.However,except Mexico, outside analysis there are no coeval associations allowing comparative that noted be can it Formation, Grupera the from described ossicles columnar the cerning Con correlations. or comparisons allow not does America North of Permian the in reports of lack Sour-Tovar,the Nonetheless,and 2015). va-Olea nent region (Villanueva-Olea the Mid-Conti with those recorded in localities of and Pennsylvanian ages, have high similarities Mexican localities, from Mississippian of ossicles the current data, it can be stated that the columnar With States. United south-central the or Mexico not recorded any taxa from the late Paleozoic of (Kotlyar / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín For example, ossicles columnar the above, mentioned As Except for the genus et al ., 2006;Biakov Cyclocaudextypicus Pentagonopternix et al Cyclocaudex insaturatus et al. ., 2016). http://dx.doi.org/10.18268/BSGM2020v72n2a280819 , 2011; Villanue has a strati has , theyhave Pennsyl ------/72(2)2020

5. Conclusions Rosa Formation (Carboniferous) of Chicomuselo of (Carboniferous) FormationRosa and crinoidsinthe region were of of Cisuralian) (early Chiapas, Mexico, sincethe only previous reports Permian the for first the represents parataxa all of recordrero. Hence,the age, Hidalgo and Guer includingBaja California, Permian reported in other Mexican localities of leave localities. vanian whose previous records were limited to Pennsyl keithi Heterostelechus for established is similar Something Permian. lower the to distribution ral tempo workpresentits the in Hidalgo,extending fMexico, Venezof America which groups different localities Permian North of Province Grandian biotic the with ated regioncles were associ inageographic deposited ossi crinoid The Cisuralian. lower the until least allowsextending range at their stratigraphical FormationGrupera their the from rocks in but occurrence America North of Pennsylvanian the had only been recorded in different localities from erathense Lamprosterigma the taxa Previously, illumination. good and productivity characterizedby well-oxygenated conditions, high theinnerramp, restricted marine environment of Cyclogrupera two newmorphogenera: rence of made in Chiapas, Mexico, highlighting the occur first the is taxa these of record The Sakmarian). muselo Chico region during the early Permian (Asselian– the inhabited that invertebrates marine the allowedhas ossicles recognizing part of crinoid elevenof morphospecies report of The useful toestablish environmental features. yet,still underdiscussion, utility is theyare very biostratigraphic their ranges, temporal long have parataxa these Because 1977). (Buitrón-Sánchez, and Lamprosterigma mirificum Lamprosterigma Cyclocaudiculusregularis, Heterostelechus keithi, Heterostelechus jeffordsi Tee rnis ee eoie i a in deposited were crinoids These . uela andtheUnited States. Cyclocaudex plenus, Preptopremnum and and apotrga erathense, Lamprosterigma Cyclocaudiculus regularis , both from the Santa the from both , have been previously Cyclocaudex insaturatus Cylindrocauliscus fiski Floricyclocion Article A280819 and ? ------

Article A280819 Table 1.Occurrencesand stratigraphicrange of crinoid ossiclesherein studied. Cyclocaudex insaturatus Cyclocaudex erathense Cyclocaudex Moore Jeffords, and Moore Jeffords, and Moore Jeffords, and Moore Jeffords, and Pentagonopternix Lamprosterigma Lamprosterigma Morphotype Jeffords, 1968 Jeffords, Jeffords, 1968 Jeffords, Cyclocaudex Cyclocaudex 1968 1968 1968 1968

Moore and and Moore Moore and and Moore

jucundus jucundus

typicus typicus plenus plenus (Villanueva-Olea al et (Villanueva-Olea al et - Santiago Fm., Visean, - Santiago Fm., Visean, Mississippian Oaxaca, Mexico Oaxaca, Mexico 2011). 2011). http://dx.doi.org/10.18268/BSGM2020v72n2a280819 Boletín de la Sociedad Geológica Mexicana Mexicana Geológica Sociedad la de Boletín ., ., USA (Moore 1968). Jeffords, and - Mingus Moscovian, Shale, Texas, 2007b). al et Mexico (Buitrón-Sánchez Atokan, Agua Sonora, Verde, - Sierra USA (Moore 1968). Jeffords, and - Fm., Canabiss Moscovian, Kansas, Sánchez al et Moscovian, Mexico (Buitrón- Oaxaca, - Fm., Ixtaltepec Bashkirian- Sánchez al et (Buitrón- Mexico Hidalgo, Calnali, - Monte Del Fm., Pennsylvanian, (Moore 1968). Jeffords, and USA - Thrifty Fm, Texas, Gzhelian, 1992). Sánchez, Buitrón- León and de Mexico (Velasco - Patlanoaya Fm, Gzhelian, Puebla, Sánchez al et Kasimovian, Sonora, Mexico (Buitrón- Moscovian- Tule, El Cerros - al Sonora, Mexico (Villanueva-Olea et - Sierra Las Mesteñas, Pennsylvanian, Sánchez al et (Buitrón- Mexico Hidalgo, Calnali, - Monte Del Fm., Pennsylvanian, 1968). USA(Moore Jeffords, and Texas, Thrifty- and fm., Graham Gzhelian, Patiño-Ruiz, 1981). - Calnali, Hidalgo (Moreno-Cano and (Moore 1968). Jeffords, and USA - Thrifty Fm., Texas, Gzhelian, 1987; Esquivel-Macías, 1996). Mexico (Villaseñor-Martínez al et - Patlanoaya Fm., Gzhelian, Puebla, (Villanueva-Olea al et Moscovian, Oaxaca, Mexico - Fm., Ixtaltepec Bashkirian- USA (Moore 1968). Jeffords, and - Limestone, Chaffin Texas, Gzhelian, al Sonora, et Mexico (Buitrón-Sánchez - Sierra Agua Verde, Moscovian, ., 2016). ., 2015). Pennsylvanian ., 2000). ., 1987). ., 2012). ., 1987). ., 2011). ., ., 2016). al et (Biakov Rusia Verkhoyansk, - Imtachan Fm. (Lopingian), (Kotlyar al et Wordian), Russia Transbaikal, - Antiinskiy (Roadian- Horizon 2007a). al et Mexico (Buitrón-Sánchez - Cerros Los Monos, Wordian, Sonora, Sánchez al et Artinskian, Hidalgo, Mexico (Buitrón- - Tuzancoa Fm., Sakmarian- al et (Arellano-Gil Artinskian, Pemuxco, Hidalgo, Mexico - Guacamaya Fm., Sakmarian- al Puebla, Mexico (Esquivel-Macías et - Patlanoaya Fm, lower Permian, Oaxaca (Vachard al et - Fm., La Cruz Wordian, Ihualtepec, Sánchez al et al et (Arellano-Gil Artinskian, Pemuxco, Hidalgo - Tuzancoa Fm., Sakmarian- ., 2000). ., 2006). ., 1987). ., 2017). Permian / 72(2)2020 ., 1998). ., 1998; Buitrón- ., 1997). ., ., 13 13

Permian crinoids from the Grupera Formation, Chiapas, Mexico CONCLUSIONS CONCLUSIONS / Permian crinoids from the Grupera Formation, Chiapas, Mexico ACKNOWLEDGEMENTS / REFERENCES Acknowledgements Table 1.(Continuation) Occurrencesandrange stratigraphicofcrinoid ossiclesherein studied. 14 14 Heterostelechus jeffordsi Moore andJeffords,1968 México, through the project PAPIIT de IA102618. Autónoma Nacional Universidad (DGAPA), General de Asuntos del Personal Académico Dirección the by supported was work This labor. technical his for Navarro Daniel and casts rubber for her photographic work, Leonora Martin for the rey Nyborg for his language revisions, Erika Porras in improving the manuscript. helpful Likewise, we thank Tor very been have anon which referee, an ymous and México) de Autónoma Nacional Carolina Martín-Cao-Romero (Universidad University), State (Washington Webster D. Gary by suggestions and comments all appreciate We Miller, 1968(inMoore Miller, 1968(inMoore Preptopremnum laeve Heterostelechus keithi / regularis and Jeffords,1968). and Jeffords,1968). Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Cyclocaudiculus Morphotype Jeffords, 1968 Mooreand Mississippian http://dx.doi.org/10.18268/BSGM2020v72n2a280819 Calnali, Hidalgo,Mexico(Buitrón- - DelMonteFm.,Pennsylvanian, (Moore andJeffords,1968). - GrahamFm.,Gzhelian,Texas,USA USA (MooreandJeffords,1968). - MingusShale,Moscovian,Texas, Parejo etal.,2018). – lowerPermianBajaCalifornia(Navas- - SierraLasPintas,upperPennsylvanian Mexico (Buitrón-Sánchezetal.,2012). - CerrosElTule,Bashkirian,Sonora, 2005, 2007b,2015). Sonora, Mexico(Buitrón-Sánchezetal., - SierraAguaVerde,Pennsylvanian, (Moore andJeffords,1968). - ThriftyFm.,Gzhelian,Texas,USA 2016). 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Permian crinoids from the Grupera Formation, Chiapas, Mexico REFERENCES