First report on the cirratulid (Annelida, Polychaeta) reefs from the Miocene Chilcatay and Pisco Formations (East Pisco Basin, Peru)
Introduction
The extant worms assigned to the family Cirratulidae are known as deposit-feeding, marine sedentary polychaetes that typically live below the water-sediment interface, exposing only their long branchiae and tentacles at the seafloor surface. They mostly burrow or crawl through the bottom sediments, although some species of at least one genus (i.e., Dodecaceria) are able to bore into shells and calcareous rocks or to construct calcareous build-ups in shallow settings (Blake and Magalhães, 2019). These reefs, consisting of adjacent tubes, develop in estuaries and coastal environments rich in organic debris. Nowadays, conspicuous cirratulid bioconstructions are known from the Gulf of Mexico near Puerto Chicxulub, Yucatan (Mexico) (ten Hove and van den Hurk, 1993) and from the Pacific coasts of Washington state (USA), in the San Juan Islands (Fischer et al., 2000).
Living cirratulid species use their tentacles for non-selectively collecting sediment particles from the surface of rocks (Gibson, 1992). As the sediment particles are trapped into mucus in each ciliated groove, the worm transfers the mucus laden with particles to the mouth (Blake and Magalhães, 2019). Exactly how these worms build their tubes is unknown, but it seems that some kind of cementing material is used in the making (Blake and Magalhães, 2019).
The construction of patch reefs by cirratulids and other tube-dwelling invertebrates (such as sabellariids, serpulids and vermetid gastropods) is the result of gregarious larval settlement as well as of asexual reproduction and/or combination of both processes, except for gastropods that reproduce only sexually (Nischi and Nishihira, 1994; Hoeksema and ten Hove, 2011; Sanfilippo et al., 2013, 2019, 2020; Rosso et al., 2016; Blake and Magalhães, 2019).
Among the extant polychaetes of the subclass Sedentaria, calcareous tube walls are known from: i) all the members of Serpulidae; ii) a single species of Sabellidae (i.e., Calcisabella (= Glomerula) piloseta (Perkins, 1991)); and iii) three cirratulid species (i.e., Dodecaceria coralii (Leidy, 1855), Dodecaceria fistulicola Ehlers, 1901, and Dodecaceria caulleryi Dehorne, 1933) (Fischer et al., 1989, 2000; Fischer, 1990; Vinn, 2005, 2009; Taylor et al., 2010). The fossil cirratulid Diplochaetetes mexicanus Wilson, 1986, which is the subject of the present study, has a mostly calcareous exoskeleton (Wilson, 1986; Fischer et al., 1989, 2000; Fischer, 1990; Vinn, 2005, 2009; Taylor et al., 2010).
The modern family Cirratulidae was investigated by Ehlers (1901) and Hartman (1959), and it was recently redefined and summarized by Blake and Magalhães (2019), who distinguished some 278 species in 11 genera, among which only Dodecaceria goes into the geological past. Fossil records of cirratulids are to date scarce: the geologically oldest cirratulid patch reefs date back to the late Oligocene of Baja California, Mexico (Fischer, 1990; Fischer et al., 1989, 2000), where they are represented by the extinct genus Diplochaetetes (Table 1). Other fossil cirratulids, belonging to the genus Dodecaceria, were reported by Fischer et al. (2000) from younger sediments of the same area (early Miocene of Baja Formation of California), and from Plio-Pleistocene deposits of Oregon and California (ten Hove and van den Hurk, 1993) (Table 1). The microstructure of tube-building polychaetes was mainly investigated by several authors, with respect to both serpulids (ten Hove and van den Hurk, 1993; Weedon, 1994; Aliani et al., 1995; Sanfilippo, 1996, 2009; Vinn, 2005, 2007, 2008; Vinn and Furrer, 2008; Vinn and Mutvei, 2009, Vinn et al., 2008c, 2008d; Taylor et al., 2010; Elorza and Astibia, 2018; Khodaverdi Hassan-Vand et al., 2018) and sabellids (Vinn et al., 2008a). As regards cirratulids, a few detailed studies by Vinn (2009) and Taylor et al. (2010) revealed that the tube wall of tube-dwelling Diplochaetetes and Dodecaceria is composed of micritic peloidal lamellae that form a stromatolitic fabric with intercalated lenses of fibrous calcite/aragonite. In particular, Vinn (2009) documented for the Oligocene species Diplochaetetes mexicanus a homogenous structure composed of irregularly oriented calcareous rods, that does not have an exact analogy in other calcareous polychaetes. Following the hypothesis by Fischer et al. (2000), according to which some structures of the cirratulid tube wall could be formed by calcification of bacterial rods (not controlled by the worm itself), Vinn (2009) supposed that the homogeneous structure of the species D. mexicanus could be likely formed by bacterial activity, but much still needs to be explored. The occurrence of modern polychaete colonies along the southern coast of Peru was mentioned by Olsson (1961: 135), who reported on bivalve Lithophaga borings into massive tube aggregates of a polychaete worm identified as the sabellariidae Gunnarea. Indeed, these masses of calcareous tubes would probably have been cirratulids of the genus Dodecaceria. Similar tube masses were recorded from Plio-Pleistocene deposits of northern Peru (DeVries, 1986: 73, 158, 255, 281, 301, and pl. 25, Fig. 1; 1988: 129) and in similarly aged deposits of southern Peru (DeVries, 2003: 345).
The present paper aims at: i) describing the first finds of fossil cirratulid reefs from the Miocene deposits of the Pisco and Chilcatay Formations of Peru; and ii) briefly discussing their palaeoenvironmental and palaeobiogeographic context.
Section snippets
Geological and paleontological framework
The southern coast of Peru hosts the Ica Desert, a district of the much larger Peru-Chile Desert, which is thought to have formed during a late Pliocene phase of increased aridity (Hartley and Chong, 2002). This Peruvian coastal desert hosts a diverse and exquisitely preserved marine fossil assemblage that includes vertebrates (e.g., Lambert et al., 2010, 2017, 2020; Collareta et al., 2015, 2017, 2020, in press; Bianucci et al., 2016a, b, c, 2018a, b; Landini et al., 2017, 2019; Marx et al.,
Material and methods
During fieldwork along the western side of the Ica River Valley (East Pisco Basin, southern Peru) in the years 2015–2017, reef boulders made by cirratulid tube aggregates were discovered at three localities (Fig. 1B):
- 1
– Corre Viento (14°26′36″ S; 75°45′34″ W), Pisco Formation, P2 allomember, upper Miocene (Tortonian) (Fig. 1, Fig. 2).
- 2
– Surroundings of Cerro Geoglifo, south of Cerro Colorado (approximated coordinates: 14°23′50″ S; 75°53′27″ W), Chilcatay Formation, unknown allomember, lower
Systematic palaeontology
Following the systematic schemes proposed by Ehlers (1901), Weissermel (1913), Reish (1952), Wilson (1986), Fischer et al. (1989, 2000), Fischer (1990) and Blake and Magalhães (2019), the examined material, including reef portions as well as polished and thin sections, was assigned to Diplochaetetes mexicanus Wilson, 1986.
Class Pleistoannelida Struck, 2011
Subclass Sedentaria Lamarck, 1818
Order Cirratulida Dales, 1963.
Suborder Cirratuliformia Fauchald, 1977
Family Cirratulidae Ryckholt, 1851
Genus
Palaeoecological inferences
During the Miocene, the East Pisco Basin was a shallow-marine, semi-enclosed embayment, protected by a chain of islands (i.e, the Gran Tablazo Archipelago of DeVries and Jud, 2018), which dampened wave energy within the bay, as reconstructed in Fig. 1B on the basis of the distribution of the pre-Cenozoic igneous rocks (Thornburg and Kulm, 1981; Marocco and Muizon, 1988; DeVries and Schrader, 1997; Bianucci et al., 2018b; Di Celma et al., 2017, 2018a, b). More in general, for most of the
Palaeobiogeographic remarks
The discovery of cirratulid specimens referable to Diplochaetetes mexicanus in the Miocene (both Burdigalian and Tortonian strata) of southern Peru stimulates some considerations on the palaeobiogeographic distribution of this extinct polychaete species. The geologically oldest record of D. mexicanus is from the southern tip of the Baja California peninsula (Pacific Mexico). The type horizon of the D. mexicanus, which occurs in the El Cien Formation, was regarded by Wilson (1986) as upper
Neogene and Quaternary cirratulid reefs from Peru
Polychaete aggregates occur in Peruvian Neogene deposits of all ages. DeVries (2020) found in situ polychaete colonies that could actually belong to cirratulids (DeVries, pers. comm., 2020) in the Plio-Pleistocene of the Sacaco area (southern Peru). The same author (DeVries, 2003) also found a small polychaete colony, preserved in situ and Lithophaga borings, in the early Pliocene south of Chala; this record could also be reassigned to Cirratulidae (DeVries, pers. comm., 2020). Polychaete reefs
Conclusions and perspectives
Exceptionally preserved cirratulid reefs have been described for the first time from Peru, South America.
Cirratulid tube-aggregates (“patch reefs”) belonging to the species Diplochaetetes mexicanus Wilson, 1986 were recorded from the lower Miocene (Burdigalian) shallow-marine sediments of the Chilcatay Formation as well as in the late Miocene (Tortonian) basal sandstones of the P2 allomember of the Pisco Formation. Such finds reveal the broad time range of this species in the East Pisco Basin.
Credit author statement
Tomáš Kočí: contributing in conceptualization and design of the study, original draft preparation, writing-reviewing and editing, writing of the first draft of the manuscript, performing tube observations and the majority of imaging data, contributing to the palaeobiogeographic section, treating and observing thin tube sections under SEM, preparing figures, taking photos under light microscope, final editing of the manuscript. Giulia Bosio: contributing in conceptualization and design of the
Declaration of competing interest
The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.
Acknowledgements
We are indebted to Jan Sklenář (NM Prague) for providing T. K. free access to the camera and optical microscope as well as for his help with photography. We are indebted to Lenka Váchová (NM Prague) for help with detailed SEM images. We are grateful to Giovanni Bianucci (Università di Pisa) for his thorough logistic support, and to Claudio Di Celma (Università di Camerino) for his invaluable stratigraphic work on the Chilcatay and Pisco strata. We would like to thank Thomas J. DeVries (Burke
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