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Review of Palaeobotany and Palynology 142 (2006) 153 160 www.elsevier.

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Chaetocladus gracilis n. sp., a non-calcified Dasycladales from the Upper Silurian of Skne, Sweden
Paul Kenrick a,, Jakob Vinther b
a

Department of Palaeontology, The Natural History Museum, London SW7 5BD, England, United Kingdom b Geological Museum, University of Copenhagen, ster Voldgade 5-7, 1350 Copenhagen, Denmark Received 16 September 2005; accepted 21 March 2006 Available online 17 August 2006

Abstract A rare non-calcified dasycladalean alga Chaetocladus gracilis n. sp. is described from the Upper Silurian (Ludlow) Bjrsjlagrd Limestone (Klinta Formation; ved-Ramssa Group), Skne, Sweden. C. gracilis comprises a slender rod-like thallus with euspondyl, acrophorous laterals arrayed in more or less equally spaced verticils. In its morphology Chaetocladus converges on several other groups of contemporaneous flora (e.g., charalean algae, land plants) and fauna (e.g., graptolites), and the potential of misattributing dasyclads of this type to other groups such as green algae and in particular to land plants is noted and discussed. The preservation of such a delicate thallus suggests that special taphonomic conditions prevailed over parts of the Bjrsjlagrd Limestone, involving rapid burial in anoxic mud and low levels of bioturbation. The presence of Chaetocladus is also indicative of an unusual floral/faunal assemblage that has been termed a thallophytic-alga-dominated biota. Assemblages of this type are characterized by thallophytic algae, annelid worms, lightly sclerotized arthropods and a low diversity shelly fauna. 2006 Elsevier B.V. All rights reserved.
Keywords: Dasycladales; Chaetocladus; Silurian; Sweden; Graptolite; Charales

1. Introduction The affinities of fossils are often puzzling, and this can lead to doubtful taxonomic assignment, fostering controversy over issues such as the time or place of origin of major groups of organisms. This is particularly true in ancient rocks and during formative periods when body plans were developing and modern lineages were diverging from a plethora of long extinct types. One problem is convergence, and one common form on which several groups of organisms have converged indepen Corresponding author. E-mail addresses: P.Kenrick@nhm.ac.uk (P. Kenrick), vinther.jakob@gmail.com (J. Vinther). 0034-6667/$ - see front matter 2006 Elsevier B.V. All rights reserved. doi:10.1016/j.revpalbo.2006.03.023

dently is the simple appendage-bearing axis. One sees this type of organisation in the stipes of graptolites, in land plants where it has been acquired independently in the leafy branches of the true mosses, the leafy liverworts, and in the distantly related clubmosses, and it is encountered again in the green algae particularly among the Dasycladales and the Charales. In these groups this simple appendicular structure belies three fundamentally different forms of construction. Graptolites are colonial organisms, land plants are fully integrated multicellular individuals, and the Dasycladales have a siphonous organisation. Differences such as these would seem to form a good basis for formulating a series of criteria for assigning fossils to groups. However, when the taphonomic history is such that soft tissues are not preserved

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and all that remains is little more than a thin film of carbon, distinguishing among these and other alternatives can be tricky. Graptolites and conodonts have been named as plants (Gabbott et al., 1995; Kenrick et al., 1999), green algae have been assigned to the graptolites (LoDuca, 1990; LoDuca, 1997), and within the green algae Dasycladales can be confused with Charales (Kenrick and Li, 1998; Feist et al., 2003). The affinities of many other putative early plant fossils are doubtful (Chaloner, 1960; Lundblad, 1972; Lemoigne, 1988; Zhang, 1988; Cai et al., 1996; Yang et al., 2004). One consequence of misattribution is that it has led to many questionable early records of land based plant life in the Lower Palaeozoic. Here we document an enigmatic appendage-bearing axis from the Silurian of Skne, Sweden. Its affinities are difficult to determine, and the specimen shows attributes of several distantly related groups of organisms. We interpret this fossil as a new species of Dasycladales in the genus Chaetocladus (LoDuca, 1997). Dasycladales has a long and highly diverse geological history that is dominated by calcareous forms (Berger and Kaever, 1992). Chaetocladus is a non-calcified genus. The absence of a robust carbonate skeleton means that this delicate siphonous alga is very much rarer in the fossil record than its carbonate encrusted relatives, and it is only preserved under exceptional taphonomic conditions (LoDuca, 1997; LoDuca et al., 2003).

2. Material and methods The description is based on a single coalified adpression preserved in a fine-grained calcareous sediment. The specimen was collected from an old quarry near the village of Bjrsjlagrd (Fig. 1) (Jeppson and Laufeld, 1986). The sediments exposed here comprise a unit of silty mudstone with intercalated thin limestone beds formed between two major biostromal carbonate successions (Fig. 2). In addition to Chaetocladus, other collectable fossils include articulated crinoids, trilobites, molluscs, scolecodonts, eurypterids, and brachiopods. These sediments form part of the Bjrsjlagrd Limestone, which is the uppermost member of the Klinta Formation. The Bjrsjlagrd Limestone Member has been correlated to the Burgsvik and Sundre Formations on Gotland (Larsson, 1979; Jeppson and Laufeld, 1986; Grahn, 1996) and to the uppermost Eke and Burgsvik Formations also on Gotland (Eriksson, 2002). These are all late Silurian in age (late Whitcliffian: upper part of the Ludfordian stage of the Ludlow epoch). The Klinta Formation and the ved Formation together form the Upper Silurian ved-Ramssa Group (Jeppson and Laufeld, 1986), which is exposed at various places throughout Skne (Scania) in the downfaulted Colonus Shale Trough (Erlstrm et al., 1997). These sediments were deposited during the closing of the marginal basin

Fig. 1. Map of Scania (Skne) outlining Lower Palaeozoic bedrock of the Colonus Trough (large black area). The white dot marks the Bjrsjlagrd locality. Modified from Bergman et al. (2004).

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Fig. 2. Picture (left) of the exposure at the Bjrsjlagrd locality, marked as Bjrsjlagrd 2 in Jeppson and Laufeld (1986). The outline drawing (right) shows the two limestone units and the mudstone unit from which Chaetocladus was collected.

between the Baltic and the Avalonian continental plates. Shallow-water deposits of mudstone and biostromal carbonates mark the transition from the underlying thick succession of Colonus Shale into the Klinta Formation. The overlying ved Formation is a shallow marine sandy deposit. Above the ved-Ramssa Group there is a major depositional hiatus until the Triassic Period. The specimen was imaged using two methods. Light micrographs were made using cross-polarized illumination to enhance contrast between the adpression of Chaetocladus and the surrounding sediment (Rowe, 1999). Scanning electron micrographs were obtained using backscatter electrons on an uncoated surface (Collinson, 1999). The specimen was scanned in a low-pressure vacuum machine (LEO 1455 VP) at about 1520 Pa and 15 kV. 3. Results 3.1. Systematics Order Dasycladales (Pascher, 1931) Family Triploporellaceae (Berger and Kaever, 1992), emend (LoDuca, 1997) Tribe Salpingoporelleae (Bassoullet et al., 1979), emend (LoDuca, 1997) Subtribe Chaetocladinae (LoDuca, 1997) Genus Chaetocladus (Whitfield, 1894), emend (LoDuca, 1997) Species Chaetocladus gracilis new species Specific diagnosis: Thallus non-calcified, slender (thallus b 3 mm wide; axis 1.5 mm1.7 mm wide, N 80 mm long), rod-like, with euspondyl, acrophorous laterals; verticils more or less equally spaced at 0.3 mm

intervals, composed of whorls of 3040 laterals, each at least 1.5 mm long and about 50 m in width. Probably endosporate. Locality: Bjrsjlagrd Quarry, Skne, Sweden. For details see Jeppson and Laufeld (1986). Age: Late Silurian (Ludlow). Repository: The Geological Institute and the Geological Museum, Copenhagen, Denmark. Holotype: MGUH 27641; Plates I and II. Etymology: The specific epithet gracilis refers to the slender, thin or slim nature of the thallus. 3.2. Description The new species is based on a single axis measuring over 8 cm in length and 1.5 mm to 1.7 mm in width (Plate I, 1). The specimen is unbranched, but it is incomplete at the apex and at the base. It does not appear to taper appreciably over its length, and obvious reproductive structures were not observed. The most striking aspect of this fossil is its segmented appearance, and this is caused by whorls of hair-like filaments (laterals) that are distributed at more or less regular intervals of about 0.3 mm (Plate I, 12; Plate II, 1). Viewed in profile, each lateral has a broad base, estimated to be about 100 m wide, which tapers to a narrow, parallel-sided filament about 50 m thick (Plate II, 4). The lengths of laterals are difficult to gauge. Some appear to be very short, whereas others are much longer. It is likely however that all were more or less the same length but that most are very incomplete. Laterals are at least 1.5 mm long, and all are borne at an acute angle. Overall, therefore, the thallus is slender, and its total width (b 3 mm) does not greatly exceed that of the axis itself (1.5 mm1.7 mm).

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We estimate that the number of lateral appendages per whorl is of the order of 3040 filaments. This is double the number that is visible on the axis surface. The reason for this is that visible filaments actually underlie the axis or emerge from the sides. They therefore represent roughly half the filaments that one would expect in the complete whorl. The other half would have been borne on the counterpart, which is missing. We are able to show that the laterals underlie the axis and are therefore attached to one side only by comparing scanning electron micrographs with light micrographs of comparable areas (Plate II, 12). The circular attachment points and the filaments are clearly visible with light microscopy (Plate II, 1), whereas only the circular attachment points are discernable using electron microscopy (Plate II, 23). This implies that the laterals but not their attachment points are covered by a thin film of sediment that is partially translucent to light but opaque to backscatter electrons. In other words, the visible lateral filaments all underlie the axis. 4. Discussion Fossils of the sort recorded here pose problems of interpretation because there is so little structure preserved and there are no remnants of internal soft tissues. The form of preservationa thin layer of carbonis itself suggestive of a relationship with algae or plants rather than animals. Could the specimen be a land plant or perhaps a member of the closely related charophycean algae? Such fossils are exceptionally rare in Silurian sediments, and any find could be of great potential significance. One needs therefore to be especially cautious in attributing affinity and in critically analysing the available evidence. The scientific literature contains examples of organisms attributed to the land plants that on further investigation turned out to be algae, hemichordates, or animal parts (e.g., Gabbott et al., 1995; Kenrick et al., 1999). Others are simply too poorly documented to fully justify a taxonomic assignment (e.g., Kenrick et al., 1999; Lundblad, 1972; Lemoigne, 1988). We therefore feel that it is important to state in a very explicit way why we attribute this fossil to Dasycladales rather than to land plants, charophycean algae, or indeed the graptolites. 4.1. Comparisons and affinities Large macroscopic green algae in the Charales (stoneworts) and land plants in the Equisetales (horsetails) have a
Plate I. Chaetocladus gracilis sp. nov. Holotype specimen MGUH 27641. 1. 2.

highly distinctive jointed appearance with parts borne in whorls. In this respect they resemble the fossil documented here, but the similarity is superficial. The leaves and leaflets of both Charales and Equisetales are larger and much fewer in number. The leaf-like laterals of Charales are more properly termed lateral branches, and these dichotomise in some genera. In living Chara and Lamprothamnium an additional whorl of stipulesspine-like cellsoccurs just below each whorl of leaves. Living Nitella, Nitellopsis, and Tolypella do not possess stipules, yet all Charales have conspicuous external male and female gametangia borne in the axils of leaves and leaflets. Stipules and external gametangia have not been observed in Chaetocladus gracilis. With regard to land plants in the Equisetales, C. gracilis would be most comparable to a lateral branch. These bear minute microphyllous leaves that are united for at least part of their lengths forming a sheath around the stem. Furthermore, the internodal regions have very clear and distinctive longitudinal ribs that reflect the underlying vasculature. Neither sheathed leaves nor longitudinal ribs were observed in C. gracilis. Modern Equisetales are of course multicellular plants with a complex internal anatomy including vascular tissues. They possess a robust cuticularised epidermis and a particularly rough textured surface caused by the deposition of massive quantities of silica (opal). Stomata are arrayed in longitudinal rows in furrows between ridges. Careful inspection of the surface of C. gracilis using scanning electron microscopy would be expected to yield some evidence for these anatomical features, however none was observed. Our analysis does not provide any evidence to support a relationship with Charales, Equisetales, or land plants in general. A further possibility is a relationship with graptolites, especially the most comparable forms of dendroid graptolites in the family Inocaulidae. On close inspection, however, the lateral filaments do not show any of the features of graptolite thecae. Furthermore, they are borne in distinct whorled arrangements, which is not a characteristic of graptolites. Also, even though incomplete our specimen is unbranched, which is inconsistent with the morphology of most if not all dendroid graptolites. Similarities to the unbranched Medusaegraptus (Ruedemann, 1925) and to Diplospirograptus (Ruedemann, 1925) are not indicative of graptolite affinity because these genera have recently been reinterpreted as dasycladalean algae (LoDuca, 1990, 1997).

Axis bearing lateral filaments. Note segmented appearance. Scale bar = 3 mm. Detail of axis showing underlying filaments. Scale bar = 1 mm.

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The new fossil most closely resembles rare noncalcified dasycladalean algae in the genus Chaetocladus (LoDuca, 1997). Key similarities include an unbranched thallus bearing more or less close-set whorls of unbranched laterals (1040 per whorl). Of the five species recognised by LoDuca (1997), Chaetocladus gracilis most closely resembles three of the more slender ones. C. ruedemanni (LoDuca, 1997) possess an axis twice as stout and a wider thallus overall with laterals some 4 times thicker. C. dubius (Spencer, 1884) emend (LoDuca, 1997) has a marginally narrower axis but a wider thallus. The laterals are of broadly similar width. In both species the verticils are more extended, being repeated at 1 mm intervals as opposed to the 0.3 mm intervals in C. gracilis. C. capillatus (Heg and Kiaer, 1926) emend (LoDuca, 1997) is the closest species. The axis, thallus, and laterals are all about twice the width of those of C. gracilis, yet the thallus is somewhat shorter. In summary, C. gracilis is considered to represent a distinct species because it is the most slender yet documented bearing the shortest or least distinct laterals. The differences noted here are comparable to those that have been used in the segregation of other species within the genus (LoDuca, 1997). 4.2. Distribution and palaeoenvironment In 1997, LoDuca published a critical review of the genus Chaetocladus recognising a total of six species. These are known from only a handful of sites in North America and northern Europe spanning an age range of Mid Ordovician to Early Devonian. The rarity of Chaetocladus is attributable partly to the delicate nature of the thallus and partly to the narrowly constrained environmental conditions in which it thrived. This distinctive dasyclad seems to be associated with rather unusual floral/ faunal assemblages, and these have been termed thallophytic-alga-dominated biotas (LoDuca, 1997). Typically, they comprise several species of thallophytic algae, annelid worms, lightly sclerotized arthropods (e.g., eurypterids, phyllocarids) and a low diversity shelly fauna. Chaetocladus gracilis is found in association with eurypterids, scolecodonts and lightly sclerotized bivalved arthropod carapaces indicating that similar associations prevailed over parts of the Bjrsjlagrd Limestone. Yet, a

diverse brachiopod fauna and the occurrence of various molluscs would seem to be atypical. Further collecting is needed to clarify more precisely the associated faunal diversity. Modern dasyclads are abundant in clear, calm, warm, shallow marine environments, as typically developed in tropical lagoons, interreef areas, and embayments. This habitat interpretation is consistent with the sediments in which Chaetocladus is found. Typically, these are finegrained laminated limestone, dolostone, and shale that cap sharp-based normally graded beds that are essentially devoid of bioturbation. At the Bjrsjlagrd Quarry locality, the mudstone containing the fossil is intercalated between two larger carbonate units. The Bjrsjlagrd Limestone member is characterized by great lateral variation, and this can be interpreted as biostromes isolated from each other by stagnant waters comprising muddy sediments. The fossilisation of such a delicate non-calcified siphonous alga is also indicative of a distinctive taphonomic environment that in addition to a lack of bioturbation is characterized by poorly oxygenated conditions at the sedimentwater interface and episodic sedimentation involving rapid burial in anoxic mud (LoDuca, 1997). Such conditions might also have occurred in stagnant but periodically storm-agitated lagoons or embayments. The discovery of rare non-calcified fossil algae such as Chaetocladus provides additional insights into the geological history of groups that are known principally through biomineralized forms. The fossil record of Dasycladales is dominated by species that develop an encrusting layer of calcite or aragonite. These carbonate minerals are far more resilient than the delicate siphonous thallus, which they envelop preserving the original algal cell as a cast. Despite the fact that the fossil record of mineralized Dasycladales is lengthy and diverse (Berger and Kaever, 1992), there is some evidence to suggest that the time of origin of certain key features and groups has been significantly underestimated. Based on non-calcified specimens from China, Kenrick and Li (1998) documented unequivocal evidence of choristospore gametangia in the Early Devonian, implying a very large range extension (ca. 240 million years) to the family Dasycladaceae. These same data also extended the known range of a mode of reproduction involving operculate cysts from the Early

Plate II. Chaetocladus gracilis sp. nov. Details of holotype specimen MGUH 27641. 1 and 2. Details of axis in same field of view imaged using two different methods: 1. Cross-polarized light. 2. Scanning electron microscopy (SEM) (backscatter electrons). Scale bars = 1 mm. Comparison of the two images illustrates that all the filaments visible in image 1 actually underlie the axis. 3. SEM (backscatter electrons) of filament attachment points. Scale bar = 200 m. 4. SEM (backscatter electrons) of filament (incomplete at apex). Scale bar = 200 m.

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P. Kenrick, J. Vinther / Review of Palaeobotany and Palynology 142 (2006) 153160 Gabbott, S.E., Aldridge, R.J., Theron, J.N., 1995. A giant conodont with preserved muscle tissue from the Upper Ordovician of South Africa. Nature 374, 800803. Grahn, Y., 1996 Upper Silurian (Upper WenlockLower Pridoli) Chitinozoa and biostratigraphy of Skne, southern Sweden. Geologiska Freningens I Stockholms Frhandlinger1181996, 237250. Heg, O.A., Kiaer, J., 1926. A new plant bearing horizon in the marine Ludlow of Ringerike. Avhandlingar utgitt av det Norske Videnskapsakademi i Oslo. I, Matematisk-Naturvidenskapelig Klasse, vol. 1, pp. 112. Jeppson, L., Laufeld, S., 1986. The late Silurian ved-Ramssa Group in Skne, South Sweden. Sveriges Geologiska Underskning. Ser. Ca, Avhandlingar Och Uppsatser 58, 145. Kenrick, P., Li, C.-S., 1998. An early non-calcified dasycladalean alga from the Lower Devonian of Yunnan Province, China. Review of Palaeobotany and Palynology 100, 7388. Kenrick, P., Kvacek, Z., Bengtson, S., 1999. Semblant land plants from the middle Ordovician of the Prague Basin reinterpreted as animals. Palaeontology 42, 9911002. Larsson, K., 1979. Silurian Tentaculitids from Gotland and Scania. Fossils and Strata 1180. Lemoigne, Y., 1988. La flore au cours des temps gologiques, I. Gobios. Mmoire Spcial 10, 1384. LoDuca, S.T., 1990. Medusaegraptus mirabilis Ruedemann as a noncalcified dasyclad alga. Journal of Paleontology 64 (3), 469474. LoDuca, S.T., 1997. The green alga Chaetocladus (Dasycladales). Journal of Paleontology 71 (5), 940949. LoDuca, S.T., Kluessendorf, J., Mikulic, D.G., 2003. A new noncalcified dasycladalean alga from the Silurian of Wisconsin. Journal of Paleontology 77 (6), 11521158. Lundblad, B., 1972. A reconsideration of Psilophyton (?) hedei, Silurian of Gotland (Sweden). Review of Palaeobotany and Palynology 14, 135139. Pascher, A., 1931. Systematische bersicht ber die mit Flagellaten in Zusammenhang stehenden Algenreihen und Versuch einer Einreihung dieser Algenstmme in die Stmme des Pflanzenreichs. Beihefte zum Botanischen Centralblatt 48, 317332. Rowe, N.P., 1999 Macrophotography. In: Jones, T.P., Rowe, N.P. (Eds.), Fossil Plants and Spores: Modern Techniques. Geological Society, London, pp. 4146. Ruedemann, R., 1925. Some Silurian (Ontarian) fossils of New York. New York State Museum Bulletin 265, 184. Spencer, J., 1884. Niagaran fossils. Transactions of the Academy of Science of Saint Louis 4, 555610. Whitfield, R.P., 1894. On new forms of marine algae from the Trenton Limestone, with observations on Buthograptus laxus Hall. Bulletin of the American Museum of Natural History 6, 351358. Yang, R.D., Mao, J.R., Zhang, W.H., Jiang, L.J., Gao, H., 2004. Bryophyte-like fossil (Parafunaria sinensis) from EarlyMiddle Cambrian Kaili Formation in Guizhou Province, China. Acta Botanica Sinica 46 (2), 180185. Zhang, Z.-Y., 1988. Longfengshania Du emend.: an earliest record of bryophyte-like fossils. Acta Palaeontology Sinica 27, 416426.

Mesozoic to the mid Palaeozoic. Similarly, the presence of euspondyl branching in Chaetocladusa key feature in dasycladalean systematicsextends the stratigraphic range of euspondyl taxa from the Early Devonian to the mid Ordovician (LoDuca, 1997). Evidence such as this indicates that there exists a significant representational bias in the fossil record of Dasycladales, favouring heavily mineralized forms. The distribution, diversity, and probably also the ecological significance of non-mineralized or weakly mineralized species of Dasycladales in the warm, shallow marine environments of the Palaeozoic are very probably underestimated. Acknowledgements We thank Dr. Ben Williamson and Dr. Alex Ball (NHM) for assistance with scanning electron microscopy and Mr. Phil Crabb (NHM) for the light micrographs. Thanks also to Dr. Per Ahlberg (Lund) and Dr. Eckart Hkansson who provided valuable information on locality and geology, and Dr. David A. T. Harper who introduced the authors and provided very helpful criticism. Paul Kenrick gratefully acknowledges Muriel Fairon-Demaret's help and support during his tenure as a Royal Society Research Fellow in Lige in 1989. References
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