[Palaeontology, Vol. 50, Part 4, 2007, pp. 951–959]

SOFT-TISSUE PRESERVATION OF THE HIND GUT IN A NEW GENUS OF CLADID FROM THE MISSISSIPPIAN (VISEAN, ASBIAN) AT ST ANDREWS, SCOTLAND by THOMAS W. KAMMER* and WILLIAM I. AUSICH *Department of Geology and Geography, West Virginia University, Morgantown, WV 26506-6300, USA; e-mail: [email protected] Department of Geological Sciences, 155 South Oval Mall, The Ohio State University, Columbus, OH 43210, USA; e-mail: [email protected]

Typescript received 20 February 2006; accepted in revised form 11 August 2006

Abstract: Soft-tissue preservation of the hind gut, or anal by pointing down current between the arms to avoid fouling sac, in the tegmen of Tubulusocrinus (gen. nov.) doliolus of the ambulacra. Its smaller size may also have required less (Wright) from the Mississippian (Visean, Asbian) Pitten- energy to produce than a heavy-plated sac. The hind gut is weem Formation at St Andrews, Scotland, is the first of its apparently preserved by haematite replacement, which was kind known in ; it sheds important new light on the probably altered from original authigenic pyrite that formed nature of the cladid tegmen. Many cladid crinoids had a cal- soon after catastrophic burial. cite-plated anal sac that may have functioned like a chimney to prevent fouling of the ambulacra. The tubular, uncalcified Key words: Anal sac, crinoids, Mississippian, Scotland, teg- anal sac of Tubulusocrinus may have functioned like a hose men, soft tissues.

The crinoid theca contains the visceral mass of the thought to be the result of rapid disintegration of the and consists of the aboral cup and tegmen, tegmen plates shortly after death, presumably because which are composed of calcite plates. The free arms they were loosely embedded in a leathery integument typically extend from the theca at the junction between rather than being closely sutured together. Alternatively, the aboral cup and tegmen. In the Palaeozoic subclass the tegmen of many cladids may have lacked calcitic Cladida, the aboral cup consists of closely sutured plates and consisted simply of a leathery integument plates that are nearly always better preserved than the like the epidermis of other classes or the tegmen, which was commonly more loosely sutured. naked tegmen of comatulids (Breimer 1978). However, The cladid tegmen includes the mouth, madreporite this latter possibility has been difficult to evaluate and anal sac (Wachsmuth and Springer 1897, pl. 3; because soft-tissue preservation in crinoids is essentially Kammer and Ausich 1996, figs 7.21, 7.22). The pre- unknown. The discovery of soft-tissue preservation of sumed functions of the cladid anal sac include housing the hind gut in the tegmen of Tubulusocrinus gen. nov. the hind gut and gonads (Lane 1984), and physically from the Mississippian, or Lower (Heckel separating the anus from the mouth to avoid faecal and Clayton 2005), of Scotland is remarkable and sheds contamination of ingested food (Ausich et al. 1999), important new light on the nature of the cladid thereby serving a function similar to the anal tube in tegmen. camerate crinoids (Baumiller 1990). In the cladids the anal structure is variously shaped and sized, and may range from a small opening in the aboral cup, such as SYSTEMATIC PALAEONTOLOGY in Gasterocoma (Moore et al. 1978, figs 376.6, 377.5), to a large variety of inflated and heavily plated sacs Terminology follows Ubaghs (1978) and higher-level (Ubaghs 1978, fig. 159). However, in the great majority taxonomy follows Moore and Teichert (1978), Simms and of taxa, the tegmen and anal sac are poorly known or Sevastopulo (1993), Ausich (1998) and McIntosh (2001). not known at all, even when the aboral cup and arms All specimens are in the Wright Collection at the are well preserved (Strimple 1973). This is generally National Museums of Scotland, Edinburgh (NMS).

ª The Palaeontological Association 951 952 PALAEONTOLOGY, VOLUME 50

Class CRINOIDEA Miller, 1821 1952b Ureocrinus doliolus (Wright); Wright, p. 325, pl. Subclass CLADIDA Moore and Laudon, 1943 13, figs 4–5. Order DENDROCRINIDA Bather, 1899 Types. Holotype, NMS G. 1958.1.1279, a complete crown preser- Remarks. We follow McIntosh (2001) in placing pinnulate ving an uncalcified anal sac. Two paratypes, NMS G. cladid crinoids in the Order Dendrocrinida, which contains 1958.1.1280 and 1281, isolated cups. both pinnulate and non-pinnulate taxa. The Poteriocrinida as defined by Moore et al. (1978) is probably a polyphyletic Description. Crown well preserved, very tall, narrow, vase-shaped where proximal arms are constricted in a girdle shape. Aboral grouping composed of several pinnulate taxa derived from cup cone-shaped, becoming slightly constricted at the radials, the Dendrocrinida sensu Moore et al. (1978). height to width ratio c.1Æ1; plates smooth, lacking ornamenta- tion; infrabasals pentagonal, prominent, clearly visible in side view and one-third to one-half cup height, with height to width Family SCYTALOCRINIDAE Moore and Laudon, 1943 ratio c.1Æ6; basals hexagonal, elongate, height to width ratio 1Æ1– 1Æ6, increasing with ontogeny; radials pentagonal, height to Genus TUBULUSOCRINUS gen. nov. width ratio c.0Æ8, with plenary facets and distinct fulcral ridge extending the entire width of facets, fossae shallow. Three anal Derivation of name. Latin tubulus, little tube. plates in cup; radianal largest; anals arranged in typical advanced cladid arrangement with anal X above and to the left of radianal, Type species. Ulocrinus doliolus Wright, 1936. and right tube plate above radianal. Anal sac expressed as an adoral tube, non-calcified and Diagnosis. Crown tall, slender, constricted at top of non-plated, elongate, with a terminal circular opening now preserved as an ellipse. Fine annular ridges along the uncalci- aboral cup and proximal primibrachials in a girdle shape, fied anal sac indicate an original leathery integument. Preser- five atomous arms with long pinnules; aboral cup cone- vation apparently iron oxide replacement of soft tissues of shaped with thin plates, higher than wide; infrabasals and sac, as indicated by red-brown colour. Uncalcified anal sac is basals higher than wide; radials wider than high with slightly three-dimensional, indicating filling with either excre- extremely shallow plenary facets; uncalcified anal sac long ment or post-mortem mud; grey mud is visible at terminal and slender, presumably an originally leathery tegmen. opening. Anal sac exits tegmen area at CD interray between arms and above primibrachial 4 of the C ray. Lower part of the anal structure consists of plates above the anals of the Tubulusocrinus doliolus (Wright, 1936) cup and adjacent to primibrachials 1–3 of the C ray. At least Plate 1, figures 1–7; Table 1 11 small tegmen plates have been deformed post-mortem and squeezed between the arms of the C and E rays; pores are present along the sutures between tegmen plates. 1934 Hydriocrinus sp. Wright, p. 244, pl. 13, fig. 1; Arms five, atomous in all rays; brachials longer than wide, text-fig. 3. subcuneate with a distinct pinnule attachment scar along the 1936 Ulocrinus doliolus Wright, p. 404, pl. 9, fig. 5; ambulacrum at the distal end of each brachial on the longest text-figs 28–32. side of the brachial; pinnules extremely long, extending to as 1938 Ulocrinus doliolus Wright; Wright, p. 337, pl. 14, much as ten brachials on the well-preserved D ray arm. Column fig. 2. subpentagonal proximally, becoming circular by fourth colum- 1939 Ulocrinus doliolus Wright; Wright, p. 30, pl. 2, nal; four preserved columnals are homeomorphic with crenulate fig. 2; pl. 8, fig. 12; text-figs 31–35. sutures indicating symplexy. 1945 Ureocrinus doliolus (Wright); Wright and Strimple, p. 224, pl. 9, fig. 6. 1952a Ureocrinus doliolus (Wright); Wright, p. 116, Remarks. Wright (1952a) apparently did not recognize pl. 19, figs 10, 13; text-figs 61–65. the soft anal sac and interpreted it as an arm (his text-

EXPLANATION OF PLATE 1 Figs 1–7. Tubulusocrinus doliolus (Wright, 1936). 1–3, NMS G. 1958.1.1279, holotype; Encrinite Bed, Pittenweem Formation, Strathclyde Group, St. Andrews, Fife, Scotland. 1, complete crown with long arms (from left to right) D, E and A rays preserved with just the proximal portions of B and C ray arms preserved, uncalcified anal sac compressed between D and E rays on slab; · 1Æ15. 2)3, close-up of uncalcified anal sac coated with ammonium chloride (2) and uncoated, in colour (3); note also oral surface of E ray arm with pinnule attachment scars, and a small cluster of tegmen plates near the base of the arm and above the primibrachials of the C ray; · 5Æ0. 4)5, NMS G. 1958.1.1281, paratype; Macgregor Marine Bands, Aberlady Formation, Strathclyde Group, Thornton Burn, East Lothian, Scotland; A–C ray and D–E ray views; · 2Æ0. 6–7, NMS G. 1958.1.1280, paratype; Encrinite Bed, Pittenweem Formation, Strathclyde Group, St. Andrews, Fife, Scotland; B–C ray and D–A ray views; · 3Æ0. PLATE 1

2

1

4 5

67

3 KAMMER and AUSICH, Tubulusocrinus 954 PALAEONTOLOGY, VOLUME 50

TABLE 1. Measurements (in mm) of Tubulusocrinus doliolus (Wright, 1936).

Crown Cup Infrabasal Basal Radial Arm Pinnule Anal sac H H,W H,W H,W H,W L L L Ray CBCBD

NMS G. 1958.1.1279 107 13Æ1, 11Æ94Æ6, 3Æ06Æ2, 4Æ03Æ9, 4Æ8911316 NMS G. 1958.1.1280 – 8Æ4, 7Æ94Æ4, 2Æ63Æ1, 2Æ72Æ7, 3Æ0– – – NMS G. 1958.1.1281 – 10Æ7, 9Æ74Æ2, 2Æ75Æ6, 4Æ03Æ1, 4Æ1– – –

fig. 63) drawing in subcuneate brachials, although the heavily plated, plicate anal sac unlike the uncalcified anal sac is barely visible in his photograph (pl. 19, fig. 10). sac of Tubulusocrinus. The anal sac does appear to have sutures suggestive of brachials, but this is the result of the soft sac being Occurrence. The holotype, NMS G. 1958.1.1279, and one para- draped over loose ossicles on the substrate. The radi- type, NMS G. 1958.1.1280, are from the Encrinite Bed along anal, anal X and right tube plates are termed the prim- the shore at St. Andrews (Fife, Scotland: British National Grid anal, secundanal and tertanal, respectively, by Webster Reference NO 513 169; Wright 1952a). The Encrinite Bed of and Maples (2006). Geikie (1902) is within the Witch Lake Marine Band of the predominantly terrestrial Pittenweem Formation, Strathclyde Group (Forsyth and Chisholm 1977, p. 37; Browne et al. Comparisons. Wright (1934, 1936) and Wright and 1999, p. 10). According to Browne et al. (1999), the Pittenw- Strimple (1945) assigned their specimens of the present eem Marine Band is within the Asbian Macgregor Marine species to Hydriocrinus, then Ulocrinus and finally Ureoc- Bands of Wilson (1974). The crinoidal band of the Encrinite rinus. Hydriocrinus shows an essentially identical cup, but Bed is 15 cm thick and is near the centre of the 3Æ6mof it has more than five arms with bifurcations at primibra- marine mudstone of the Witch Lake Marine Band at chial 1 or higher and is only known from the Upper Car- St Andrews (Forsyth and Chisholm 1977, p. 38). boniferous (Middle Pennsylvanian) (Moore et al. 1978, The other paratype, NMS G. 1958.1.1281, is from the Macgre- p. T643). Ulocrinus has a thick-plated conical cup and ten gor Marine Bands, Thorton Burn, East Lothian, Scotland (NT arms, and is also known only from the Middle and Upper 739 740; Wright 1952b). Wilson (1989) correlated these beds Pennsylvanian (Upper Carboniferous) (Moore et al. 1978, with the Asbian Substage. Formerly, they were placed in the Upper Lothian Group of the Calciferous Sandstone Measures p. T701). Wright and Strimple (1945) created Ureocrinus (Wilson 1974), but they are now assigned to the Aberlady For- with Poteriocrinus bockschii Geinitz, 1846 as the type spe- mation of the Strathclyde Group (Browne et al. 1999). cies and included Ulocrinus doliolus Wright in that genus. Tubulusocrinus doliolus is similar to Ureocrinus bockschii in having five arms. However, it is distinct in having a DISCUSSION conical cup composed of thin plates, whereas Ureocrinus has a high-bowl, globose cup with thick plates and is con- Preservation of echinoderm soft tissue stricted at the top of the cup rather than at the proximal primibrachials (Moore et al. 1978, p. T701). Tubulusocrinus The echinoderm mesodermal skeleton is highly suscept- doliolus and U. bockschii co-occur in the Encrinite Bed at ible to disarticulation; thus, the preservation of complete St Andrews (Wright 1952a). echinoderm skeletons requires rapid burial (see Ausich Tubulusocrinus is placed in the Scytalocrinidae because 2001 and references therein). In fact, it is probable that of the following shared characters (Moore et al. 1978): most complete fossil were buried alive. Per- tall, slender crown; conical cup with infrabasals visible in haps unexpectedly, the special conditions leading to side view; wide radial facets; three anal plates in cup; uni- exceptional echinoderm specimens very rarely led to pre- serial arms; and some genera with five arms, although servation of soft tissues. Organic films (Springer 1901; some others may have up to four bifurcations in line per Meyer and Milsom 2001) can be associated with fossil cri- ray. Other scytalocrinids with five arms include Anemetoc- noids, but certain identifiable soft parts are not known rinus, Ophiurocrinus and Gilmocrinus. Anemetocrinus has among them [see Ausich and Babcock (1998, 2000) and robust, biserial arms, while Ophiurocrinus has arms that Sprinkle and Collins (1999) for a discussion of Echmato- are large and round, with short brachials that give the crinus, now regarded to be a cnidarian]. appearance of columnals. Tubulusocrinus is most similar Within Mississippian camerate crinoids, lightly miner- to Gilmocrinus, but differs in having the proximal arms alized internal organs with a spicular endoskeleton have constricted inwards as in Ureocrinus, rather than project- been preserved by secondary calcification and silicifica- ing outwards as in Gilmocrinus. Gilmocrinus iowensis tion (Haugh 1975; Haugh and Bell 1980). This type of Laudon, 1933, type species of the genus, has a large, KAMMER AND AUSICH: SOFT-TISSUE PRESERVATION IN A CARBONIFEROUS CRINOID 955 preservation is intermediate between the heavy-plated atomous arms that are all clearly visible on NMS G. mesodermal skeleton of crinoids, and truly soft tissues 1958.1.1279 (Pl. 1, fig. 1). Furthermore, the uncalcified anal such as tube feet and the uncalcified anal sac of Tubu- sac lacks any evidence of plating, neither impressions in the lusocrinus gen. nov. bedding surface nor sutural subdivisions. Thus, this tubular The most remarkable soft-tissue preservation among structure cannot be an aberrant or regenerated arm. Also, echinoderms is the calcium-phosphate replacement in a the fact that the structure is filled with matrix and com- ophiuroid (Pinna 1985). Perhaps the most com- pressed indicates that it could not have been a typically mon soft-bodied preservation among fossil echinoderms preserved, plated structure of a cladid crinoid. Another is seen in the holothurian Achistrum from the Pennsylva- potential interpretation is that this tubular structure repre- nian Mazon Creek fauna of Illinois (USA) (Sroka and sents a typical anal sac that is preserved in the process of Blake 1997). In this instance the holothurian body is regenerating. This would follow the hypothesis of Lane faintly preserved as a mould or composite impression (1984) that anal sacs functioned as an ‘offering’ to preda- with some mineralization and is encased within a siderite tors to promote non-lethal predation, a strategy apparently concretion. Other holothurian soft-body preservation was used by the modern comatulid crinoid Antedon where the reported by Pawson (1980) and Smith and Gallemı´ gonad-bearing pinnules are taken by predators in prefer- (1991). asteroid tube feet have been reported ence to the more vulnerable calyx (Nichols 1996). However, as moulds (Gale 1987) and in pyritized form (Glass and in contrast to the completely uncalcified tubular structure Blake 2002, 2004; Glass 2006), and plated tube feet have of Tubulusocrinus, regenerating anal sacs do have plating as been reported in ophiocistioids (Jell 1983). Skin-like they develop (Lane 1984). structures and rare pyrite preservation of other soft tis- Preserved tube feet are unknown in crinoids despite sues have also been reported for asteroids, ophiuroids, occurrences in Lagersta¨tten where one might reasonably holothurians and crinoids from the Hunsru¨ck Slate expect such preservation, such as the Solnhofen Lime- (Lower , Germany) (Schmidt 1934; Lehmann stone of Germany (Barthel et al. 1990) or the Mazon 1957, 1958; Bergstro¨m 1990; Bartels and Blind 1995; Brig- Creek fauna of Illinois (Sroka and Blake 1997). Addition- gs et al. 1996; Su¨dkamp 1997). All accounts of soft-tissue ally, complete crinoids were subject to rapid burial, com- preservation in the Hunsru¨ck Slate have been contested monly in fine-grained sediments, and some faunas (Otto 1994, 2000). However, the evidence put forward by underwent pyritization (Hunsru¨ck Slate, Germany; Silica Glass and Blake (2002, 2004) is convincing. They docu- Shale, USA; Kesling and Chilman 1975), yet definitive mented tube foot preservation through pyritization in the tube feet have not been reported in crinoids. Thus, the ophiuroids Bundenbachia beneckei Stu¨rtz and Palaeo- anal sac in Tubulusocrinus must have had some special phiomyxa grandis (Stu¨rtz). Furthermore, Glass (2006) attributes that allowed it to be preserved. In addition to made a convincing argument for preserved tube feet in a rapid burial in fine-grained sediments, perhaps the Late Ordovician asteroid, Protasterina flexuosa (Miller and integument of Tubulusocrinus consisted of more leathery Dyer) from Kentucky (USA). tissue than was typically present on other crinoid uncalci- fied tegmens or tube feet. This is speculation, but it sug- gests that careful cleaning of other closely related crinoids Unique preservation of Tubulusocrinus may reveal similar structures.

The discussion above emphasizes the rarity of echinoderm soft-tissue preservation. Where it does occur, it is typic- Functional morphology of the anal sac of Tubulusocrinus ally of isolated morphological features, such as a few contiguous tube feet or a ‘ghost’ of tissue, as among holo- Before discussing the tegmen of Tubulusocrinus,itis thurians. Thus, the soft-tissue preservation of a complete necessary to review the terminology associated with cri- anal sac in Tubulusocrinus is truly remarkable. We do not noid tegmen structures. In the subclasses Cladida, Flexi- know the precise stratigraphic context of this Tubulusocri- bilia and Disparida, the anal structure is referred to as an nus specimen, but neither the crinoid (with the exception anal or ventral sac, because of its often non-rigid and of the anal sac) nor the matrix appears any different from inflated form and its location on the ventral or adoral other crinoid material from the Pittenweem Formation. side of the body. In cladids, the anal sac is commonly Thus, it follows that some aspect of the anal sac of Tubu- robustly plated (e.g. Lane 1975). Homologous to the anal lusocrinus may have been unique, allowing a greater sac of cladids, the structure in the Articulata is commonly potential for preservation. referred to as the anal cone or tube. In the subclass Cam- What are other possible explanations and why are other erata, the anal structure is referred to as an anal tube soft structures not preserved? Tubulusocrinus doliolus has a because of its often rigid and more tubular structure. very simple filtration fan consisting exclusively of five These different names are also thought to indicate inde- 956 PALAEONTOLOGY, VOLUME 50 pendent evolutionary origins (Ausich et al. 1999; Ausich Lane’s (1984) hypothesis is correct, where were the and Kammer, work in progress). gonads? The tegminal plates preserved between the B and The structure preserved on Tubulusocrinus was clearly a E ray arms (Pl. 1, figs 2–3) may have been part of a small soft tube that arose from the tegmen, and we regard it as tegmen that was squeezed between the arms by compac- homologous with the anal sac of a cladid. Thus, although tion after death and burial. These small plates have very distinct from a typical heavy-plated anal sac, it is con- numerous pores along their sutures that may have sidered to be an uncalcified anal sac that housed the hind allowed for the passage of gametes or brooded larvae, as gut. It is non-calcified, non-plated, and smooth with no suggested by Lane. Thus, the gonads may have been in a evidence of any sutures or articulations. It is elongate, small tegmen separate from the uncalcified anal sac that essentially 2 mm wide along its entire length of 16 mm. At contained only the hind gut. the terminal end there is an elliptical opening formed by compression of an originally circular opening. A rim is vis- ible within the tube, defined by grey mud. There is no evi- Taphonomy of Tubulusocrinus dence of tearing, so the tubular structure appears to be complete and not truncated. The sac is slightly three- The holotype specimen of Tubulusocrinus doliolus con- dimensional, indicating that it was filled with excrement sists of calcite plates preserved on a small slab of crinoi- or mud post-mortem. The structure is also slightly undu- dal packstone (Pl. 1, fig. 1). Preservation of the lating from being laid out over an uneven surface of loose uncalcified anal sac, or hind gut, is apparently by haem- ossicles on the sea-floor. These undulations are not atite replacement of the soft tissues, as indicated by the brachial plate sutures as drawn by Wright (1952a, text- red-brown colour and smooth, shiny surface in plain fig. 63). The original leathery integument of the uncalcified light (Pl. 1, fig. 3). Much of the surface of the holotype sac is evident from fine annular ridges preserved near the slab is also stained red-brown. Original replacement may base, visible at high magnification (·25). The uncalcified have been by authigenic pyrite associated with sulphate anal sac exits the tegmen area at the CD interray between reduction under anoxic conditions in the sediment after the arms above primibrachial 4 of the C ray and is laid out rapid burial (Briggs et al. 1991, 1996; Briggs 2003), on the bedding surface between the D and E ray arms. although the bottom waters may have been well oxygen- This position at the CD interray is the precise location for ated (Raiswell et al. 1988). The thin coating of haematite the base of the cladid anal structure and, moreover, sup- over much of the slab suggests that a microbial mat ports the interpretation that this is the anal sac. once covered the specimen and surrounding sea-floor The non-rigid character of the sac may have allowed (Allison and Briggs 1991). Allison and Briggs (1991) it to point down-current, like a hose, through the long emphasized the importance of rapid mineralization after arms of Tubulusocrinus to avoid fouling the ambulacra burial because anaerobic decay can be fairly rapid. Allen with waste. This would have been an advantage over a (2002) demonstrated in laboratory experiments that rigid, calcified anal sac. Also, calcified sacs were often pyritic sulphide can begin to form around a buried car- as long as the arms to avoid fouling of the ambulacra cass after only three days. This is certainly fast enough (Ubaghs 1978, fig. 159). They were much larger and to preserve soft tissues before they decay. Liddell (1975) required more energy to produce than the short, uncal- reported that modern comatulid crinoids begin to disar- cified sac of Tubulusocrinus, although their function ticulate on the sea-floor one day after death, but when may have included more than simply serving as a waste buried are still articulated after six days. Therefore, chimney. although the correct sequence and timing of burial and Lane (1984) argued that the anal sac of the advanced diagenesis must occur, the timing constraints would cladids was enlarged to hold not only the hind gut but allow for soft-tissue preservation. also the gonads. If so, the pores present at the plate The pyrite apparently oxidized to haematite before dis- boundaries on most preserved anal sacs were probably for covery of the holotype specimen by James Wright on out- the release of gametes or brooded larvae. Eggs in living crop along the shore at St Andrews. Wilby et al. (1996) crinoids range from 135 to 150 lm in diameter, and reported reddened lithologies from iron oxidation associ- would easily pass through these pores (Mortensen 1920; ated with pyrite preservation of soft tissues. They also Nakano et al. 2003; Sevastopulo 2005). Modern crinoids recorded pyrite infilling within the gut of Jurassic deca- have a small tegmen with an anal cone and lack a com- pod crustaceans. Presumably, the hind gut of Tubulusocri- parable anal sac like those of the Cladida, and the gonads nus was pyritized in a similar manner. In view of the fact are located on the arms in genital pinnules (Breimer that specimen NMS G. 1958.1.1279 is the holotype of 1978). The uncalcified anal sac of Tubulusocrinus appears Ulocrinus doliolus Wright, 1936, and because it is the only to have functioned strictly as a hind gut because of its known example of soft tissue preservation of a crinoid narrow, tubular structure and single terminal opening. If tegmen, it is inappropriate to damage the specimen by KAMMER AND AUSICH: SOFT-TISSUE PRESERVATION IN A CARBONIFEROUS CRINOID 957 taking a sample for mineralogical analysis. Thus, the pres- preservation of this feature. Many cladid crinoid genera ence of haematite is inferred. do not preserve any evidence of a plated anal sac, so the discovery of Tubulusocrinus raises the possibility that many of these crinoids may have had an uncalcified Palaeoecology of Tubulusocrinus rather than a plated anal sac. The non-rigid character of such a sac may have allowed it to be directed down cur- Asbian sedimentary rocks in central Scotland are predom- rent through the arms to avoid fouling the ambulacra inantly non-marine siliciclastics ranging from mudstones with waste, which would have been an advantage over to sandstones, and include lacustrine oil shales, coals, rigid, calcified anal sacs, which were often very long, like seatearths and ironstones (Wilson 1989), representing flu- a chimney, to avoid fouling of the ambulacra. They were vial, lacustrine and deltaic depositional environments also much larger and required more energy to produce (Read et al. 2002). The marine rocks include limestones, than the short, uncalcified anal sac of Tubulusocrinus, mudstones and silty mudstones. These strata comprise although the longer plated sacs may have had the addi- Yoredale-type cyclothems in which marine rocks are suc- tional function of housing the gonads. ceeded by upward-coarsening, fluvial–deltaic deposits capped by palaeosols and coals (Browne et al. 1999; Read Acknowledgements. L. Anderson (National Museums of Scotland, et al. 2002; Kassi et al. 2004). At the St Andrews shore Edinburgh) provided access to the Wright Collection and section, c. 300 m of the Asbian Pittenweem Formation arranged the loan of the type specimens. This research was sup- are exposed, including three marine bands that total c.9 ported by the US National Science Foundation grants EAR- m in thickness, namely the West Sands Marine Band (or 0206307 to TWK and EAR-02059068 to WIA. S. K. Donovan, T. K. Baumiller and an anonymous reviewer improved an earlier Cuniger Rock Marine Band), Witch Lake Marine Band draft of this manuscript. and St Andrews Castle Marine Band (Forsyth and Chis- holm 1977; Browne et al. 1999). The Witch Lake Marine Band consists of c.3Æ6 m of mudstone and includes the REFERENCES 15-cm crinoidal limestone Encrinite bed near the centre (Forsyth and Chisholm 1977), which may record maxi- ALLEN, R. E. 2002. Role of diffusion-precipitation reactions in mum water depth. authigenic pyritization. Chemical Geology, 182, 461–472. The Witch Lake Marine Band contains crinoids, brachi- ALLISON, P. A. and BRIGGS, D. E. G. 1991. Taphonomy of opods, molluscs and bryozoans (Forsyth and Chisholm nonmineralized tissues. 25–70. In ALLISON, P. A. and 1977). It probably represents shallow-marine deposition BRIGGS, D. E. G. (eds). Taphonomy. Releasing the data locked associated with a rapid glacio-eustatic rise in sea level that in the fossil record. 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