Agglutinated Worm Tube from the Late Cretaceous of Colombia

Home , Alvinellidae, Conejo Formation, Pectinariidae, Petta (polychaete), Terebellida

Cretaceous Research 41 (2013) 107e110

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Cretaceous Research

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First record of a pectinariid-like (Polychaeta, Annelida) agglutinated worm tube from the Late Cretaceous of Colombia

Olev Vinn a,*, Javier Luque b a Department of Geology, University of Tartu, Ravila 14A, Tartu 50411, Estonia b Smithsonian Tropical Research Institute, Balboa-Ancón 0843-03092, Panamá, Panama article info abstract

Article history: The earliest agglutinated pectinariid-like tube with a modern appearance is described from the Late Received 11 August 2012 Cretaceous (Santonian, w84 My) of Colombia. The rare agglutinated fossil tube is composed of sorted Accepted in revised form 20 November 2012 skeletal material, quartz sand, and unidentified carbonaceous fragments. It’s solitary and noneencrusting Available online 13 December 2012 life mode, the straight conical shape, and the agglutinated tube wall composed of sand-sized grains, supports affinity with the tube-building Family Pectinariidae. This finding suggests that Pectinariidae Keywords: might have first appeared in the Neotropics at least by the late Mesozoic. Polychaeta Ó 2012 Elsevier Ltd. All rights reserved. Pectinariidae Tubeworms Santonian South America

1. Introduction that produce agglutinated tubes are important constituents of modern oceans, their role in past ecosystems is poorly understood Polychaete worms without mineral tubes are rarely preserved (Fournier et al., 2010). as fossils. However, some polychaetes produce organic, aggluti- The aim of the present paper is to: 1) describe the first specimen nated or calcareous tubes, enhancing the likelihood of preserva- of a pectinariid-like agglutinated worm tube from the Mesozoic of tion. Polychaetes with calcareous tubes have a good fossil record South America; and 2) discuss the evolutionary implications of the and are usually well preserved, but only species within the Ser- described pectinariid-like tube. pulidae, Sabellidae, and Cirratulidae produce these tubes (Vinn and Mutvei, 2009). Conversely, agglutinated tubes have a poor 2. Material fossil record, despite the fact that several modern polychaete families produce them. Therefore, fossil records of agglutinated One pectinariid-like tube was found in light-beige to greyish tubes help us better understand the evolution of tube-building terrigenous mudstones from the Upper Cretaceous Conejo Forma- strategies in polychaete annelids. The oldest agglutinated fossil tion (lower to middle Santonian, w84 My), outcropping near the tubes are known from as far back as the Cambrian (Signor and town of Toca in the Department of Boyacá, Cordillera Oriental McMenamin, 1988). However, it is possible that these fossils do (Latitude 53702800; Longitude 731205300). Other fauna found in this not belong to polychaetes, but to various problematic Palaeozoic deposit include abundant bivalves, principally corbulids and limids, tube-producing worms (Zaton et al., 2012) with affinities that scattered echinoderm remains, linguloid brachiopods, and sporadic suggest that they probably should not be assigned to the annelids decapod crustaceans (Luque et al., in preparation). The agglutinate (Vinn and Mutvei, 2009). The earliest records of agglutinated pectinarid-like tube, as well as most associated fossil invertebrates, tubes with strong polychaete affinities are known from the Late are compressed parallel to lamination. The presence of the Palaeozoic and Mesozoic (Ettensohn, 1981; Zaton et al., 2012). The ammonite Pseudoschloenbachia inconstans (Grossouvre, 1893) diversity and evolution of polychaetes with agglutinated tubes, suggests a lower to middle Santonian age for the beds (Kennedy however, remain poorly known. In addition, although polychaetes et al., 1995; Etayo, personal communication). The specimen is deposited in the Museo Geológico Nacional José Royo y Gómez, Servicio Geológico Colombiano (formerly INGEO- * Corresponding author. MINAS), Bogotá DC, Colombia, under the acronym and catalogue E-mail addresses: [email protected] (O. Vinn), [email protected] (J. Luque). number IMG p880662.

3. Results terrigenous fraction consists of fine to medium (w0.34 mm to w0.55 mm), sub-rounded quartz sand grains. The fossil pectinariid tube is composed of a mixture of biogenic Measurements.dtube length: 23.5 mm; anterior diameter: and terrigenous sand grains, generally elongated in shape, and 7.7 mm. agglutinated nearly perpendicular to the tube’s main axis (Fig. 1). The grain size varies from 0.34 to 1.13 mm (n ¼ 21, mean 0.61 mm, 4. Discussion s ¼ 0.19). The principal biogenic constituents of the tube are fine to very coarse sand-sized particles dominated by angular to sub- The grains of the agglutinated tube we describe here are sorted, angular shell fragments, probably of molluscan and/or crustacean with only certain grain sizes present, in contrast to the finer grain origin, followed in minor proportion by fish scales debris and size of the surrounding sediment (average particle sizes ranging unidentified carbonaceous fragments (w0.45 mm to w0.80 mm). from coarse to medium-size silt (w0.060 mm) to clay-size No microfossil tests or debris are present. The far-less-abundant (w0.002 mm)). However, the surrounding sediment contains

Fig. 1. Pectinariidae. A, An extant pectinariid polychaete dwelling its agglutinated tube (aperture right) (Photo by Hans Hillewaert). BeE, Agglutinated tube of pectinariid-like specimen IMG p880662 , IMG p880662, from the Upper Cretaceous, Conejo Formation (lower to middle Santonian, w84 My), Department of Boyacá, Cordillera Oriental, Colombia. B, overview (aperture right). C, Detailed view of the grains at the aperture. D, Detailed view of the grains in the middle part of the tube. E, Detailed view of the grains in the proximal part of the tube. O. Vinn, J. Luque / Cretaceous Research 41 (2013) 107e110 109 a small number of larger grains comparable to grain sizes used in an agglutinated polychaete tube (possibly an Amphicteridae) from the wall of possible pectinariid tube. Thus, the dominance of the Early Jurassic of England (Barnard, 1956) that, based on the biogenic detritus among the grains of the agglutinated tube can be shape and orientation of the grains within the tube, has a tube explained by the rarity of terrigenous sand grains and the lack of similar to our specimen. However, that Early Jurassic specimen has microfossils of suitable size in the habitat to make use of them, as a less conical shape than typical for pectinariids. Agglutinated tubes we did not find microfossils of sand-fraction size in the rock matrix. of possible polychaetes are also known from the Jurassic of Poland The pattern of tube walls composed of large grains despite the (Zaton et al., 2012), and the Cretaceous of Oman (Wilson and Taylor, small grain size of the surrounding sediment observed here can be 2001), but in contrast to our Late Cretaceous agglutinated tube, the explained by the ecology of these animals. Pectinariids are deposit former are encrusting on hard substrates and have a much less feeders that ingest volumes of sediment (Fauchald and Jumars, conical shape. Their tubes are also curved unlike the specimen 1979) that can exceed 400 cm3/year (Gordon, 1966). These worms described here. On the other hand, similar pectinariid tubes to the build their tubes of all sand-fraction size grains available in the one described here are known from the Miocene of Japan, but they environment. Thus, during the processing of large volumes of occur on bedding planes as broken and deformed tubes of fine sand sediment with their tentacles, worms find sufficient larger detritus (Katto, 1976). The middle to late Santonian age of our specimen and sand-sized particles from which to build their tubes, despite represents the earliest record of a pectinariid-like tube constructed living in generally very fine-grained sediment. For example, the from sand and macrofossil debris instead of microfossils. extant Pectinaria regalis (Verril, 1900) from Puerto Rico, has a tube that consists of coral and shell fragments, along with foraminiferan 5. Conclusion and molluscan shell fragments (Long, 1973). Petta assimilis McIntosh, 1885 (Pectinariidae), from the South Atlantic, has a tube The evolutionary history of Pectinariidae is poorly understood, composed of agglutinated foraminifera (Hartman, 1967), because due to their sparse fossil record. Despite this, based on this spec- the sediment in the environment was very fine-grained and fora- imen and the literature, we can conclude that agglutinated fossil minifera tests were probably the most available particles of the tubes of similar appearance to those made by Recent Pectinariidae desired sand size (Finger et al., 2008). occurred already in the Late Cretaceous, and may have their origin There are a considerable number of polychaete families con- in pre-Santonian times. This places the origin of Pectinariidae in taining species that have the ability to incorporate sediment in a similar time period to several other groups of tube-producing their tubes. These include: the Sabellariidae, Sabellidae, Oweniidae, polychaetes, including the serpulids and sabellids, which first Alvinellidae, Pectinariidae, Terebellidae, Trichobranchidae, Apisto- appeared in the Mesozoic (Vinn and Mutvei, 2009). branchidae, Longostomatidae, Chaetopteridae, Arenicolidae, Mal- danidae and Capitellidae (Wilson and Taylor, 2001). The described Acknowledgements fossil tube can be assigned to the order Terebellida based on its straight, agglutinated morphology. Members of Terebellida, such as J.L. thanks Carlos Jaramillo (STRI) for providing funds and facil- Alvinellidae, Pectinariidae, Terebellidae, Trichobranchidae are ities to allow the collection of the specimen and development of the known to produce a variety of agglutinated tubes. We tentatively present research. Harry A. ten Hove, Sancia van der Meij, Micha1 assign the Late Cretaceous fossil tube described here to the family Zaton, Igor A. Jirkov, Mario H. Londoño Mesa, Sergio Salazar, Alex- Pectinariidae (Terebellida) based on its size, solitary, none ander Tzelin and Stephen Gardiner are thanked for their valuable encrusting life mode, the characteristic “ice-cream-cone” shape, comments and advice. Cesar Silva (STRI) and Jakeline Vanegas and agglutinated tube wall composed of sand-sized grains. In assisted in the field. Fernando EtayoeSerna (Colombian Geological addition, the elongated grains are agglutinated relatively perpen- Survey) supplied intellectual support. Kecia Kerr (McGill) assisted dicular to the tube’s main axis similar to the orientation of agglu- with editing of the manuscript. Hans Hillewaert photographed the tinated grains in some possible pectinariids from the Miocene of extant pectinariid polychaete. Jose Arenas (Colombian Geological California (Finger et al., 2008). Somewhat similar agglutinated Survey) provided export permits. O.V. is grateful to The Paleonto- tubes to the Pectinariidae are built by the phylogenetically closely logical Society for receipt of a J.J. Sepkoski Grant, to the Estonian related family Terebellidae. However, with the exception of a few Science Foundation for grant ETF9064 and to a target-financed species, Terebellidae tubes are usually curved. Those Terebellidae project from the Estonian Ministry of Education and Science species possessing straight tubes are less conical, and they have less (SF0180051s08) for financial support. We are grateful to Mark A. orderly agglutinated particles in their tube wall than Late Creta- Wilson (The College of Wooster) and anonymous reviewer for the ceous tube described here. Terebellidae tubes do not differ constructive reviews. We also are grateful to journal editor Peter remarkably from those produced by other tubes builders, such as in Harries for several improvements in the manuscript. the Maldanidae, Ampharetidae, Sabellidae, Onuphidae, and others, so they can easily be misidentified. In contrast, the Pectinariidae References produce unique tubes (Igor A. Jirkov, personal communication, 2012), which are straight, more conical and have particles agglu- Barnard, T., 1956. An unusual worm tube from the Lower Lias. Journal of Paleon- tology 30, 1273e1274. tinated in the wall in more orderly manner, as in the Late Creta- Ettensohn, F.R., 1981. Crininicaminus haneyensis, a new agglutinated worm from the ceous tube described here. These characteristics indicate Chesterian of east-central Kentucky. Journal of Paleontology 55, 479e482. a Pectinariidae affinity for this specimen. Fauchald, K., Jumars, P., 1979. The diet of worms: a study of polychaete feeding e Despite the fact that pectinariids use biomineralization to guilds. Oceanography and Marine Biology 17, 193 284. Finger, K.L., Flenniken, M.M., Lipps, J.H., 2008. Foraminifera used in the construction secrete reinforcing cement between agglutinated grains in the tube of Miocene polychaete worm tubes, Monterey Formation, California, USA. wall (Fournier et al., 2010), their tubes are rarely preserved as fossils Journal of Foraminiferal Research 38, 277e291. fi because they are thin and fragile (Ettensohn, 1981), and usually Fournier, J., Etienne, S., Le Cam, J.-B., 2010. Inter- and intraspeci c variability in the chemical composition of the mineral phase of cements from several tube- disintegrate quickly after the death of the animal, obscuring our building polychaetes. Geobios 43, 191e200. understanding on the origin and evolution of agglutinate tubicu- Gordon Jr., D.C., 1966. The effects of the deposit feeding polychaete Pectinaria gouldii lous worms throughout geologic time. In the literature, the earliest on the intertidal sediments of Barnstable Harbor. Limnology and Oceanography 11, 327e332. record of possible pectinariids is from the Permian (Howell, 1962). Grossouvre, A.D., 1893. Recherches sur la craie supérieure. Partie 2. Paléontologie. However, their detail description is missing. There is also a record of Les ammonites de la craie supérieure. Imprimerie Nationale, Paris. 110 O. Vinn, J. Luque / Cretaceous Research 41 (2013) 107e110

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