fossil sponges of argentina: a review - ib.usp.brmcust/7thiss/09-book/pdf/beresi - fossil sponges...

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11 PORIFERA RESEARCH: BIODIVERSITY, INNOVATION AND SUSTAINABILITY - 2007 Introduction The actual knowledge of the record fossil in Argentina is based on relatively rigid whole skeletons, fragments of skeletal nets and spicules seen in thin sections or recovered from acid residues for obtaining conodonts. The fossil record of sponges comes from several geological provinces of Argentina with different lithologic, paleontologic and environmental characteristics. Sponge faunas were collected from marine sediments of the Puna, Cordillera Oriental, Sierras Subandinas basins of northern Argentina; Famatina Range, Precordillera Terrane, San Rafael Block, Neuquén Basin (western Argentina), from lacustrine deposits of the North Patagonian Massif, and from the Chaco-Paranaense Basin. Occurrences of sponge faunas have been reported from the Lower Paleozoic (upper-Lower Cambrian) to the Cenozoic (Tertiary). Only a few publications have dealt with sponge and sponge spicules in Argentina. Most of them concern the fauna from the Cambrian and Ordovician rocks of the Precordillera terrane. Associated with protospongiid spicules, well-preserved chancelloriid sclerites occur in the Cambrian carbonate platform of the Precordillera. Chancelloriids are Cambrian enigmatic organisms constituting the monophyletic taxon Coeloscleritophora (Bengtson and Missarzhevsky 1981). Sclerites of chancelloriids (Family Chancelloridae) were first described from the Burgess Shale by Walcott (1920), who interpreted them as heteractinid sponges. This traditional view of the fossil group as sponges was accepted for more than 50 years. In the Precordillera, chancelloriid sclerites associated with spicules are a common and distinguishing features of the Lower to Middle Cambrian fossil fauna. Protospongia and Chancelloria crucensis (Rusconi, 1955) were the first Cambrian species of the Mendoza Precordillera. Cambrian Protospongia Salter, 1864 and hexactinellid spicules were mentioned by Pernas (1964), Devizia (1973), Bordonaro and Martos (1985), Heredia et al. (1987) and Beresi and Heredia (1995) from the San Isidro area in the Precordillera of Mendoza. Afterwards, assemblages tentatively identified as Kiwetinokia utahensis Walcott, 1920, protospongiid skeletal nets and Chancelloria eros Walcott, 1920 were reported from early-Middle Cambrian carbonates blocks of San Juan and Mendoza, Precordillera, by Beresi and Rigby (1994). Two small associated specimens assigned to Protospongia sp. occur in the western Precordillera of San Fossil sponges of Argentina: a review Matilde Sylvia Beresi CONICET-CRICYT: Ianigla, Dto. de Geología y Paleontología, Avda Ruiz Leal s/n, 5500 Mendoza, Argentina. [email protected] Abstract: This is a review on fossil sponges and sponge spicules reported from several regions in Argentina and in strata ranging in age from Early Cambrian to Tertiary. Sponges have been collected from marine sediments of the Puna, Cordillera Oriental and Sierras Subandinas basins, northern Argentina; Famatina Range; Precordillera terrane, San Rafael block, Neuquén basin and from lacustrine deposits of the North Patagonian Massif. Knowledge of the sponge fossil record is based on whole relatively rigid skeletons, fragments of skeletal nets and spicules seen in thin sections or recovered from acetic acid residues. Early to Middle Cambrian Porifera and Chancelloriids are known from the carbonate platform and slope facies of the Precordillera terrane. Specimens with body preservation of Protospongia, Diagoniella, Kiwetinokia, fragments of hexactinellid, and anthaspidellid sponges and sclerites of Chancelloria had been reported from Cambrian of the Precordillera. Remains of hexactinellid sponges, Pelicaspongiidae and Protospongiidae, have been found in Ordovician rocks of the Puna and of the Famatina System, western margin of Gondwana. Protospongia sp. and hexactinellid mesh were reported from Upper Cambrian-Lower Ordovician siliciclastic sediments in the Cordillera Oriental and Sierras Subandinas. The most significant fossil record of Lower-Middle Ordovician sponge faunas is from the carbonate platform of the San Juan Precordillera. Sponge faunas are dominated by orchoclad lithistid demosponge genera, although hexactinellids are known from loose spicules and root tufts, and calcareous heteractinid sponges are known from isolated octactine spicules and only one genus. Hexactinellid, calcarean and demosponge spicules were reported from diverse localities of the Precordillera. A Late Jurassic (Oxfordian) carbonate complex of the Neuquén Basin, west-central Argentina, contains siliceous sponges dominated by hexactinellids (Hexactinosa and Lyssakinosa). Palaeospongilla chubutensis, a fresh water sponge, was described from lacustrine Cretaceous deposits of the Chubut River valley. Oxeas and strongyles, belonging to the Family Spongillidae, have been mentioned from Tertiary sediments of the Paraná basin, northeastern Argentina. Keywords: biostratigraphy, fossil sponges, Argentina.

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11Porifera research: Biodiversity, innovation and sustainaBility - 2007

Introduction

The actual knowledge of the record fossil in Argentina is based on relatively rigid whole skeletons, fragments of skeletal nets and spicules seen in thin sections or recovered from acid residues for obtaining conodonts. The fossil record of sponges comes from several geological provinces of Argentina with different lithologic, paleontologic and environmental characteristics.

Sponge faunas were collected from marine sediments of the Puna, Cordillera Oriental, Sierras Subandinas basins of northern Argentina; Famatina Range, Precordillera Terrane, San Rafael Block, Neuquén Basin (western Argentina), from lacustrine deposits of the North Patagonian Massif, and from the Chaco-Paranaense Basin. Occurrences of sponge faunas have been reported from the Lower Paleozoic (upper-Lower Cambrian) to the Cenozoic (Tertiary). Only a few publications have dealt with sponge and sponge spicules in Argentina. Most of them concern the fauna from the Cambrian and Ordovician rocks of the Precordillera terrane.

Associated with protospongiid spicules, well-preserved chancelloriid sclerites occur in the Cambrian carbonate platform of the Precordillera. Chancelloriids are Cambrian

enigmatic organisms constituting the monophyletic taxon Coeloscleritophora (Bengtson and Missarzhevsky 1981). Sclerites of chancelloriids (Family Chancelloridae) were first described from the Burgess Shale by Walcott (1920), who interpreted them as heteractinid sponges. This traditional view of the fossil group as sponges was accepted for more than 50 years.

In the Precordillera, chancelloriid sclerites associated with spicules are a common and distinguishing features of the Lower to Middle Cambrian fossil fauna. Protospongia and Chancelloria crucensis (Rusconi, 1955) were the first Cambrian species of the Mendoza Precordillera.

Cambrian Protospongia Salter, 1864 and hexactinellid spicules were mentioned by Pernas (1964), Devizia (1973), Bordonaro and Martos (1985), Heredia et al. (1987) and Beresi and Heredia (1995) from the San Isidro area in the Precordillera of Mendoza. Afterwards, assemblages tentatively identified as Kiwetinokia utahensis Walcott, 1920, protospongiid skeletal nets and Chancelloria eros Walcott, 1920 were reported from early-Middle Cambrian carbonates blocks of San Juan and Mendoza, Precordillera, by Beresi and Rigby (1994). Two small associated specimens assigned to Protospongia sp. occur in the western Precordillera of San

Fossil sponges of Argentina: a reviewMatilde Sylvia Beresi

CONICET-CRICYT: Ianigla, Dto. de Geología y Paleontología, Avda Ruiz Leal s/n, 5500 Mendoza, Argentina. [email protected]

Abstract: This is a review on fossil sponges and sponge spicules reported from several regions in Argentina and in strata ranging in age from Early Cambrian to Tertiary. Sponges have been collected from marine sediments of the Puna, Cordillera Oriental and Sierras Subandinas basins, northern Argentina; Famatina Range; Precordillera terrane, San Rafael block, Neuquén basin and from lacustrine deposits of the North Patagonian Massif. Knowledge of the sponge fossil record is based on whole relatively rigid skeletons, fragments of skeletal nets and spicules seen in thin sections or recovered from acetic acid residues. Early to Middle Cambrian Porifera and Chancelloriids are known from the carbonate platform and slope facies of the Precordillera terrane. Specimens with body preservation of Protospongia, Diagoniella, Kiwetinokia, fragments of hexactinellid, and anthaspidellid sponges and sclerites of Chancelloria had been reported from Cambrian of the Precordillera. Remains of hexactinellid sponges, Pelicaspongiidae and Protospongiidae, have been found in Ordovician rocks of the Puna and of the Famatina System, western margin of Gondwana. Protospongia sp. and hexactinellid mesh were reported from Upper Cambrian-Lower Ordovician siliciclastic sediments in the Cordillera Oriental and Sierras Subandinas. The most significant fossil record of Lower-Middle Ordovician sponge faunas is from the carbonate platform of the San Juan Precordillera. Sponge faunas are dominated by orchoclad lithistid demosponge genera, although hexactinellids are known from loose spicules and root tufts, and calcareous heteractinid sponges are known from isolated octactine spicules and only one genus. Hexactinellid, calcarean and demosponge spicules were reported from diverse localities of the Precordillera. A Late Jurassic (Oxfordian) carbonate complex of the Neuquén Basin, west-central Argentina, contains siliceous sponges dominated by hexactinellids (Hexactinosa and Lyssakinosa). Palaeospongilla chubutensis, a fresh water sponge, was described from lacustrine Cretaceous deposits of the Chubut River valley. Oxeas and strongyles, belonging to the Family Spongillidae, have been mentioned from Tertiary sediments of the Paraná basin, northeastern Argentina.

Keywords: biostratigraphy, fossil sponges, Argentina.

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Juan (Beresi and Banchig 1997). A synthesis of the Cambrian sponge occurrence in the Argentine Precordillera was given by Beresi (2003a).

Silicified sponge spicules from residues of conodonts from diverse Cambrian and Ordovician sections in the Precordillera, were described by Mehl and Lehnert (1997). Anthaspidellid skeletal fragments from the Middle Cambrian rocks of the Precordillera, in the province of San Juan, document the only known occurrence from South America (Beresi and Rigby 1994). Protachileum kayseri Zittel, 1877 from the San Juan province, was the first report of a Precordilleran sponge. Taxonomic studies have been concentrated in the rich sponge fauna of the warm carbonate platform (San Juan Formation) from the Lower-Middle Ordovician of the Precordillera basin (Beresi and Rigby 1993, Carrera 1996a, 1996b).

The purpose of this paper is to review the occurrence of fossil sponges and their biostratigraphic distribution in the diverse geological provinces of Argentina.

Cambrian and Ordovician fauna from the Northwest Argentinian region

Siliciclastic sediments with minor carbonates dominate the Cambrian and Ordovician of Northern Argentina and the Central Andean Basin of South America. Cambrian and Ordovician sponges and spicules from Northwest Argentina provide additional paleontological data from the siliciclastic platforms of western Gondwana.

Puna

In the Puna region (Fig. 1A-B), a single specimen (Fig. 2M-N) of a complete Ordovician hexactinellid sponge was discovered. It has been collected from volcaniclastic rocks of the Las Vicuñas Formation (Tremadocian) in Lari Creek, southwest of the Salar del Rincon area, Salta province, Argentina. The material was assigned to the new genus and species Larispongia magdalenae (Carrera, 1998), that belong to the family Pelicaspongiidae, and it is the first record of the family in western Gondwana.

Subandean Ranges

The first report on the occurrence of one completely preserved hexactinellid sponge is a part and counterpart of a round small sponge described as Diagoniella sp. (Beresi et al. 2006). This sponge (Fig. 2H) was reported for the first time from the Upper Cambrian-Lower Ordovician siliciclastic rocks of the Orcomato Formation, in the Candelaria Range, Sub-Andean Ranges, in Salta Province (Fig. 1B).

The fossiliferous record of the unit suggests a Cambrian-Ordovician age. The sponge is preserved in greenish-yellowish shales, as dark stained flattened rounded bodies (somewhat deformed and fragmentary).

Eastern CordilleraHexactinellid meshes of Protospongia (Beresi et al. 2006)

were recently reported from Lower Tremadocian siliciclastic sediments in the Eastern Cordillera, Salta Province (Fig. 1). The hexactinellid spicules were recovered from the lower

Fig. 1: A. South America showing the position of Argentina. B. Location of the Basins of Argentina with sponge fauna: (Pu) Puna; (CO) Cordillera Oriental, (SS) Sierras Subandinas, (F) Famatina, (Pa) Paraná basin, (P) Precordillera, (SR) San Rafael Block, (N) Neuquén basin, (MP) North Patagonian Massif. C. Map of Argentine showing the San Juan and Mendoza provinces. D. Sponge localities within the Precordillera Precordillera of San Juan and northern Mendoza provinces.

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levels of the Saladillo Formation, at the Angosto de la Quesera section, Eastern Cordillera.

Spicules of Protospongia (Fig. 2K) are preserved on upper surfaces of yellow-brownish shales and sandstones belonging to the lower section of the unit, sharing the stratigraphic position with Tremadocian graptolites. The sedimentation of the greenish and yellowish shales and sandstones of the Saladillo Formation indicates a transition to an upper offshore-lower offshore environmental setting.

Famatina system

Catamarca ProvinceThe only discovery of Ordovician sponges in this

province corresponds to isolated hexactinellid spicules,

which have been described by Aceñolaza and Toselli (1977) for the Chaschuil region. The material appears dispersed in carbonate concretions of the Suri Formation (Arenig).

La Rioja ProvinceThe sponge material came from black siliceous graptolitic

shales of the upper part of the Volcancito Formation of Lower Tremadocian age, outcropping in the Peña Negra location in the Famatina range (Fig. 1B). Fragments of a reticulated skeletal net of Protospongia species (A and B) were described by Esteban and Rigby (1998), in the siliclastic Famatina basin, western margin of Gondwana (Fig. 2L).

The sponges are associated with planktonic graptolites and this level can be assigned to the Lower Tremadoc (Esteban and Gutierrez-Marco 1997).

Fig. 2: A. Cambrian anthaspidellid sponge fragment, genus and species indeterminant from San Isidro, Mendoza. B-G. Coeloscleritophora: Chancelloria eros Walcott, 1920, sclerites from the upper Lower–Middle Cambrian of Zonda Range and San Isidro area. H. Diagoniella sp. from Salta province, Sub-Andean ranges. I. Cambrian fragment of bioclastic limestone showing sclerites of Chancelloria Walcott, 1920, San Isidro. J. Root tuft, large monaxons or monactine-like spicules, in bundle with small stauracts or hexacts (Tontal Range). K. Protospongia sp. showing stauractine-based skeleton and long rayed hexactines forming marginal spines along the left margin (Tontal Range). L. Protospongia sp. Esteban and Rigby, 1998, specimen PIL 14.192 from Peña Negra section, Famatina region. M-N. Larispongia magdalenae Carrera, 1998, holotype CEGH-UNC 17365 from Lari Creek, Puna region. N. Detail of the same specimen showing dermal hexactines surrounding major gaps.

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Precordillera (Cuyania) Terrane, Western Argentina

The Argentine Precordillera is situated along the forefront of the high Andes at approximately 28º to 37º S, and it is a major geologic province in northwestern Argentina. It contains a complete thick sequence of Early Paleozoic rocks.

The Precordillera, as part of the Cuyania Terrane, was formed during the Andean (Tertiary) crustal shortening. This distinctive terrane can be recognized mainly on the basis of its key stratigraphic composition, involving biostratigraphic, sedimentary and magmatic events; its boundaries with adjacent geologic regions are abrupt (Ramos et al. 1986). In accordance with the Terrane concept, the present Precordillera, plus the San Rafael Block and San Jorge Limestones, integrate a unique geologic entity, the so-called Precordillera Terrane or Cuyania Terrane.

Two hypotheses exist regarding the origin of the Precordillera: 1) the Precordillera represents a terrane of Laurentian origin that became attached to Gondwana (western Argentina) already during Ordovician times (Thomas and Astini 2003). It includes either the classical Precordillera as well as the San Rafael Block, to the south in the province of Mendoza, and the San Jorge Limestones cropping out in the Province of La Pampa, within the Sierras Pampeanas structural setting as an allochthonous terrane “Cuyania”, accreted to Gondwana during the lower Paleozoic. 2) the Precordillera is considered as an autochthonous Gondwanan fragment (Baldis et al. 1989, Aceñolaza et al. 1999, 2002) displaced by simple transcurrence mechanics, from a hypothetical intermediate sector between South America, Africa, and Antarctica. Recently, U-Pb geochronology of detrital zircons indicated a Gondwanan provenance for Lower Cambrian and Upper Ordovician sandstones of the Precordillera of western Argentina, supporting the autochthonous Gondwanan nature of the Precordillera Terrane (Finney et al. 2003a, 2003b).

Cambrian PoriferaThe Cambrian system of the Argentinian Precordillera

is represented by a carbonate platform, in the east, and of a continental slope, in the west. Cambrian platform and slope facies, containing spicules and chancelloriids sclerites are located in the Precordillera of San Juan and northern Mendoza provinces, western Argentina (Fig. 1C-D).

Cambrian sponges are known mainly from fragments of skeletal nets and dissociated spicules from the shallow carbonate platform sequences of the upper-Lower to Middle Cambrian in the eastern and central belts, and from the slope olistostromic sequences with allochthonous blocks in the western part of the Precordillera of San Juan Province. The spicules from the Upper Cambrian were collected in Tontal Range, San Juan Precordillera (Beresi and Banchig 1997) and in the La Cruz Olistolith, San Isidro area (Beresi and Heredia 1995), Precordillera of Mendoza Province.

Sclerites of Chancelloria (Coeloscleritophora) occur in shallow carbonate platforms and allochthonous blocks in

the Precordillera of San Juan and Mendoza. Well preserved sclerites are associated with protospongiid spicules (Fig. 2B-G).

Cambrian assemblagesTwo spicule assemblages occur in the Cambrian facies

(Beresi and Rigby 1994, Beresi 2003a). The autochthonous assemblage corresponding to material collected from the upper Lower to Middle Cambrian platform sequence of the eastern Precordillera of San Juan. This assemblage consists of a variety of stauractines and sclerites of Chancelloria eros (Walcott, 1920). The Protospongiidae are represented by triradiate prodianes, pentactines and hexactines, all belonging to Kiwetinokia utahensis Walcott, 1920, Protospongia and anthaspidellid fragments. This fossil fauna represents the oldest assemblages known of Argentina.

The allochthonous assemblage proceeds from the diverse Cambrian carbonate olistoliths of slope sequences of the western San Juan Precordillera and from the classical area of San Isidro, Mendoza Province. The assemblages consist of the first precordilleran Protospongia with body preservation, Diagoniella Rauff, 1894, Kiwetinokia Walcott, 1920 and Chancelloria and skeletal net with hexactines and monaxons (Beresi and Banchig 1997).

Demosponges have a limited record in the Cambrian of the Precordillera. Typically anthaspidellid fragments with dendroclones (Fig. 2A) have been reported from the carbonate platform and slope sequences of the San Juan and Mendoza Precordillera (Beresi and Rigby 1994).

Ordovician PoriferaDeposits of Ordovician carbonate basins occur in the

central and eastern Precordillera. The Lower-Middle Ordovician sediments of the Precordillera represent a drowning carbonate platform with a diverse and relatively complete fossil record.

Well-preserved and diverse faunas of sponges have been collected from limestones of the San Juan Formation (Upper Tremadoc-Early Llanvirn) in the Precordillera basin of San Juan province (Fig. 1C-D). This fauna represents the most significant Ordovician sponge fauna known from South America and provides the first extensive record of sponges derived from a stable carbonate platform, constituting one of the most important Early Ordovician sponge associations of the world.

Precordilleran sponge faunas are dominated by orchoclad lithistid demosponge genera, although hexactinellids are known from loose spicules and root tufts, and calcareous heteractinid sponges are known from isolated octactine spicules and only one genus. Spicules assemblages were reported from diverse localities of the Precordillera (Beresi and Esteban 2003, Carrera 2003).

The San Juan Formation was deposited on an open carbonate shelf, bounded to the west by continental slope and oceanic basin deposits. The diverse marine fauna and the lack of specific structures indicative for shallow water

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conditions point to low energy, subtidal conditions within an open-platform environment during the entire sedimentation interval. The age of the San Juan Formation is well constrained by conodonts (Sarmiento 1985, Albanesi and Ortega 2002) spanning from the late Tremadocian (Paltodus deltifer Zone) to the early Darriwilian (Lenodus variabilis – Eoplacognathus suecicus zones).

The first report of a Precordilleran sponge was Protachileum kayseri Zittel, 1877 from the Talacasto Gulch, Precordillera of San Juan Province (Fig. 3J).

Anthaspidellid genera first appear in the basal beds of the San Juan Formation (Upper Tremadoc) at the Niquivil Range, Eastern Precordillera of San Juan, associated with

reef-mound (Cañas and Carrera 1993). The cosmopolitan Archaeoscyphia Hinde, 1889 and Rhopalocoelia Raymond and Okulitch, 1940 are the predominant genera in this sponge-algal association (Fig. 3H, 3E).

Diverse and well-preserved sponge faunas are from the middle and upper part of the San Juan Formation (Arenig-Lower Llanvirn). In this carbonate platform the fauna is dominated by orchocladine lithistid demosponges. Their first taxonomic study was made by Beresi and Rigby (1993) and afterwards by Carrera (1996a, 1996b, 1998). Apart from orchocladine demosponges there are hexactine spicules, hexactinellid root tufts, and isolated octactine spicules that document the presence of the Heteractinida.

Fig. 3: Early and Middle Ordovician orchoclad lithistid sponges from the Precordillera of San Juan. (A-B-C-D-G-K; Beresi and Rigby 1993). A. Talacastonia chela Ianigla PI T-2. B. Tangential thin section. C. Anthaspidella annulata Ianigla PI T-49. D. Calycocoelia perforata Ianigla PI VI-2. E. Rhopalocoelia clarkii Raymond and Okulitch, 1940, Ianigla PI T-22. F. Hudsonospongia cyclostoma Raymond and Okulitch, 1940, Ianigla PI VI-2. G. Hudsonospongia talacastensis. Ianigla PI T-32. H. Archaeoscyphia minganensis Billings, 1859 Ianigla PI T-47. I. Incrassospongia ramis Carrera, 1996b, CEGH-UNC 9308. J. Protachileum kayseri Zittel, 1877, Ianigla PI H-43. K. Aulocopium sanjuanensis Ianigla PI VI-13.

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The greatest generic and specific diversity of lithistid sponges occurs during the Lower Llanvirnian (Darriwilian) in the upper most part of the San Juan Formation. A total of 15 demospongiid genera and 20 species are described. This fauna shows a variety of external morphologies and body plan.

Many new Ordovician species were described for the San Juan Formation by Beresi and Rigby (1993): Anthaspidella inornata, A. annulata (Fig. 3C), and A. alveola, Archaeoscyphia nana, Aulocopium sanjuanensis (Fig. 3K), Calycocoelia perforata (Fig. 3D), Hudsonospongia talacastensis (Fig. 3G), H. cyclostoma (Fig. 3F), Patellispongia robusta, Psarodictyum magna, Rhopalocoelia rama, among other species. The new species Talacastonia chela (Fig. 3A-B) was described from the classical Ordovician Talacasto section, Central Precordillera (Fig. 1B). Root tufts of hexactinellids also occur (see Table 1).

New megamorinid genera as Rugospongia viejoensis (family Saccospongidae) (Carrera, 1996a) and the tricranocladine sponge Eoscheiella concave (Family Hindiidae Rauff, 1893) have been recovered from the top of the San Juan Formation in the Cerro Viejo, Huaco (Carrera 2007). Nexospongia sillaensis (family Nexospongiidae Carrera, 1996a) was described from the Cerro La Silla, Niquivil Range, Eastern Precordillera.

In the upper levels of the San Juan Formation from the Early Llanvirnian, at the Cerro La Chilca section, the calcareous heteractinid Chilcaia bimuralis (Carrera, 1994) and a lithistid species Incrassospongia ramis (Carrera, 1996a) were described (Fig. 3I).

Endemic genera such as Protachileum and Talacastonia Beresi and Rigby, 1993, from the Talacasto Gulch and Rugospongia Carrera, 1996 and Chilcaia from different localities of the San Juan Formation occur in the Precordillera.

Ordovician sponges from the Precordillera show changes from algal-sponge (reef ecosystems) in the Early Arenig to stromatoporid associations in the Middle Arenig to anthaspidellid demosponge dominated associations in the Upper Arenig to Lower Llanvirn. From the Llanvirn up to the Upper Ordovician, the effects of diverse abiotic factors such as volcanic activity, sea level fluctuations and finally the global climatic cooling, could have contributed to the decrease of the sponges diversity.

The diversification of the orchoclad demosponges in the Lower Ordovician carbonate platform of the Precordillera was similar to worldwide radiation pattern (Carrera and Rigby 1999).

Ordovician sponge spicules

Precordillera of San Juan ProvinceThe oldest spicule assemblage comes from the Lower

Ordovician limestones (Oepikodus intermedius Zone) of the San Juan Formation. The Arenigian silicified spicule

assemblages were documented by Gnoli and Serpagli (1980) in the Pachaco section, western Precordillera.

Calcarean and demosponge spicules assemblages recovered from residues of conodont samples of several Lower to Middle Ordovician sections of the San Juan Precordillera were described by Mehl and Lehnert (1997). The well preserved silicified spicules include: Polyactinellidae, Heteractinellidae (Calcarea) and hexactinellid and demosponge spicules. The species Dodecaactinella oncera Mehl and Lehnert, 1997, Sardospongia cynodonta Mehl and Lehnert, 1997, Praephobetractinia sp. and Eiffelia sp. were reported from Lower Ordovician (Arenig) strata of the San Juan Formation (Fig. 4A-E).

These spicule assemblages were collected from reef-mound horizons and biostromes with sponges, stromatoporoids and receptaculitids of the San Juan Formation (Lower Arenig-Lower Llanvirn) and from the Gualcamayo and Las Aguaditas formations (Lower Llanvirn to Caradocian).

San Rafael BlockSponge spicules are derived from residues of conodont

samples from Middle Ordovician strata, in the geological province of the San Rafael Block, southern Mendoza Province, Argentina (Fig. 1C-D). Spicules (Fig. 4F-Q) have been recovered from the Ponón Trehué Formation, a clastic-carbonate sequence. Poriferan taxa (Beresi and Heredia 2000) include two spicule assemblages: 1) associations of exclusively heteractinellid spicules (sexiradiates) restricted to Arenigian allochthonous blocks of the Oepikodus evae Zone (Heredia 2001) from the shallow platform of the San Juan Formation; and 2) associations of hexactinelliid spicules, calcarean triaene and monaxons, from Upper Llanvirnian autochthonous limestones and carbonate sandstones of the Pigodus serra Zone and the P. anserinus Zone (Heredia 2001) from the outer platform and slope.

The spicule associations of the Ponón Trehué Formation represent the most austral Ordovician assemblage described in the context of the Precordillera (Cuyania) Terrane.

Jurassic sponges from the Neuquén Basin

A late Jurasic (Oxfordian) carbonate complex was developed on the foreland side of the Neuquén Basin (Fig. 1B), west- central Argentina and form part of the Cordillera Principal. Shelf carbonates facies are exposed throughout Mendoza and Neuquén provinces.

One of these facies consists of small siliceous sponge buildups of the La Manga Formation (Plicatilis Zone), well developed at the Río Poti Malal section, in southern Mendoza Province. The siliceous sponges with moderate diversity are fossilized in their original shape and exhibit calcareous preservation.

Sponge fauna is dominated by hexactinellids (Hexactinosida and Lyssacinosida, 95%) and lithistid demosponges (5%). Up to now, approximately 20% of the material has been preliminary determinated (Beresi 2003b).

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Sponge taxa

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DEMOSPONGIAEFamily Anthaspidellidae Carbonate platform

Allosacus sp. Carrera, 1994 XAnthaspidella alveola Beresi and Rigby, 1993 XAnthaspidella annulata Beresi and Rigby, 1993 XAnthaspidella inornata Beresi and Rigby, 1993 XArchaeoscyphia minganensis Beresi and Rigby, 1993 XArchaeoscyphia nan Beresi and Rigby, 1993 XArchaeoscyphia pulchra Bassler, 1941 XAulocopium sanjuanensis Beresi and Rigby, 1993 XCalycocoelia perforata Beresi and Rigby, 1993 XHudsonospongia cyclostoma Raymond and Okulitch, 1940 XHudsonospongia talacastensis Beresi and Rigby, 1993 XIncrassospongia ramis Carrera, 1996 XPatellispongia argentina Carrera, 1994 XPatellispongia occulata Bassler, 1941 XPatellispongia robusta Beresi and Rigby, 1993 XPatellispongia magna sp. XProtachilleum kayseri Zittel, 1877 XPsarodictium magna Beresi and Rigby, 1993 XRhopalocoelia clarikii Raymond and Okulitch, 1940 XRhopalocoelia rama Beresi and Rigby, 1993 XRhopalocoelia regularis Raymond and Okulitch, 1940 XRhopalocoelia tenuis Carrera, 1994 XTalacastonia chela Beresi and Rigby, 1993 X

Family HindiidaeEoscheiella concave Carrera, 2007 X Carbonate platform

Family NexospongiidaeNexospongia sillaensis Carrera, 1996 X Carbonate platform

Family SaccospongiidaeRugospongia viejoensis Carrera, 1996 X Carbonate platform

HEXACTINELLIDAFamily Pelicaspongiidae

Larispongia magdalenae Carrera, 1998 X Marine volcaniclasticFamily Protospongiidae

Protospongia sp. A X Silicoclastic platformProtospongia sp. B X Silicoclastic platformDiagoniella sp. X X X Carbonate platformProtospongia sp. X X X Carbonate platformKiwetinokia utahensis Walcott, 1920 X X Carbonate platform

Family uncertainRoot tuft X Carbonate platform

HETERACTINIDA Carbonate platformChilcaia bimuralis Carrera, 1994 XDodecaactinella oncera Mehl and Lehnert, 1997 XSardospongia cynodonta Mehl and Lehnert, 1997 XPraephobetractinia sp. XEiffelia sp. X XOctactine spicules XTriactine spicules X

Table 1: Biogeographic distribution of sponge taxa from the Cambrian and Ordovician Argentine Basins.

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The following species have been identified: Laocoetis sp., Laocoetis procumbens and Laocoetis parallela (Hexactinosida Schrammen, 1903, Family Craticulariidae Rauff, 1893) (Fig. 5C-J); Cribrospongia sp., Cribrospongia clathrata and Cribrospongia cucullata (family Cribrospongiidae Roemer, 1864) (Fig. 5A-B) and Polygonatium sp. (Lyssacinosida Zittel, 1877).

Sponges belonging to the Family Cribrospongiidae are cup-shaped (Cribrospongia reticulata), tubular and conical. Only a few specimens are triangular in shape and compressed (Cribrospongia cucculata). Fragments of cylindrical to

tubular sponges belong to the genus Laocoetis (=Craticularia Zittel, 1877; emend. Schrammen, 1937).

Cretaceous freshwater sponge from the North Patagonian Massif

The only freshwater sponge was described from Lower Cretaceous lacustrine sediments of the Chubut River Valley, in the Chubut Province, North Patagonian Massif (Fig. 1B). The sponge was determined as Palaeospongilla chubutensis by Ott and Volkheimer (1972). The encrusting sponge belongs to the monogeneric family Palaeospongillidae Volkmer-

Fig. 4: Ordovician sponge spicules. A-E. Calcarean and demospongid spicules from the Lower-Middle Ordovician of the San Juan Precordillera (Mehl and Lehnert 1997). CEGH-UNC 15951. A. Sardospongia cynodonta. B. Calcarean triactine. C. Demospongid oxea. D-E. Dodecaactinella oncera. F-Q. Spicules from the Ponón Trehué Formation, San Rafael Block (Beresi and Heredia 2000). F. Octactine spicule shows the distal and two of the tangential rays broken. G. Octactine spicule. H. The proximal-distal vertical ray shows a prominent node. I. Monaxon spicule appears to have been sheared diagonally by diagenetic processes. J. Calcarean triactine. K-Q. Scanning electron microscope (SEM) photomicrographs. K, M, N, P. Octactine spicules. L. monaxon spicule shows the central circular canal. O, Q. Hexactine spicules.

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Ribeiro and Reitner, 1991. It is characterized by acanthoxeas to acanthostrongyles gemmoscleres (Fig. 5K-O).

Tertiary spicules of the Paraná basin

Isolated oxeas and strongyles, possibly belonging to the species Trochospongilla repens (family Spongillidae) were collected from Tertiary (Miocene) pelitic sediments of the Paraná basin, northeast of Argentina (Fig. 1).

Borings of Cliona entrerriana and C. ameghinoi on calcareous shells of Ostrea patagonica have also been found in Tertiary sediments.

Remarks

Fossil sponges are known from several geologic basins with different lithologic, sedimentologic and environmental characteristics (Table 2). There are sponges and loose spicules representative of the Classes: Hexactinellida, Demospongida and Calcarea.

The oldest sponge fauna known in Argentina is from the upper Lower Cambrian of the Precordillera, the youngest one occurs in the Middle Tertiary of the Paraná Basin.

Protospongiids characterized the western old Gondwana continent and are known from the Cambrian and Lower

Fig. 5: Oxfordian hexactinellid sponges at the Río Potimalal section, Neuquén Basin (Beresi 1997) A. Cribrospongia cuccullata Quenstedt, 1878, lateral view. B. Dermal surface with craticulariid diplorhysis. C. Laocoetis procumbens Goldfuss, 1826, lateral view. D. Upper view of the osculum sponge showing in H. E. Upper view of the narrow osculum and folded wall in a cribrospongiid sponge. F. Lateral view of a cylindrical sponge. G. Laocoetis clathrata Goldfuss, 1833, lateral view. H. Longitudinal section of a tubular cribrospongiid sponge. I. Longitudinal section of a tubular cribrospongiid sponge. J. Upper view of the same sponge, showed in I. K-M. Palaeospongilla chubutensis Ott and Volkheimer, 1972. K. Megasclere with central canal. L. Gemmules. M. Gemmule and spicular texture of the sponge skelton. N-O. Tertiary megascleres, Paraná basin.

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Ordovician of the Puna, Eastern Cordillera and Sub-Andean Ranges of northern Argentina and the Famatina System, occurring also in Cambrian rocks of the Precordillera.

Middle Ordovician sponge faunas are mostly dominated by demosponges, whereas calcareans and hexactinellids do not occur frequently in the warm carbonate platform of the Precordillera.

Within the range of the Oxfordian Plicatilis Zone Hexactinellid sponges are common in the carbonate facies of the Neuquén basin, similarly to the Oxfordian European sponge facies.

There is still a long way to go concerning the fossil sponges of Argentina, especially in the Cambrian of the Precordillera and northern regions, the Jurassic of Neuquén Basin and freshwater sponges from Mesozoic and Cenozoic lacustrine sediments.

Acknowledgements

The author thanks the Scientific Committee of the 7th International Sponge Symposium for the invitation to participate as Invited Lecturer in the session “Palaeontology”, and Dr. W. Volkheimer (Conicet-Ianigla, Mendoza) for critic reading of the manuscript. This paper is a contribution to the CONICET-Project: PIP 5222, Argentina.

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Taxon sponges and spicules types Formation Basin Age Environment

DEMOSPONGIAEFamily Spongilliidae Paraná Chaco-Paranense Middle

Tertiary Fluviomarine ? Trochospongilla repens Bonetto and Ezcurra de Drago, 1969

Family PalaeospongilliidaeAgrio North Patagonian

Massif Cretaceous: Hauterivian Lacustrine Paleoespongilla chubutensis Ott and Volkheimer, 1972

HEXACTINELLIDAOrder HexactinosidaFamily Craticulariidae

La Manga Neuquén Upper Jurassic Oxfordian

Marine, carbonate bioherms

Laocoetis paradoxa Goldfuss, 1833Laocoetis procumbens Goldfuss, 1826Laocoetis parallela Goldfuss, 1826Laocoetis clathrata Goldfuss, 1826

Family CribrospongiidaeCribrospongia sp.Cribrospongia cucullata Quenstedt,1878

Order LyssacinosidaPolygonatium sp.; Schrammen, 1937

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