Acta Geologica Polonica, Vol. 55 (2005), No.3, pp. 269-282

Lower Kimmeridgian comatulid crinoids of the Holy Cross Mountains, Central Poland

URSZULA RADWANSKA

Institute of Geology, University of Wm:mw, AI. Zwirki i WigUl)" 93; PL-02-089 Warszawa, Poland. E-mail: u.radwallska@ Ulv.edupl

ABSTRACT:

RADWANSKA, U. 2005. Lower Kimmeridgian comatulid crinoids of the Holy Cross Mountains, Central Poland. Acta Geologica Polonica, 55 (3), 269-282. Warszawa.

An assemblage of feather stars or comatulids (free-living crinoids of the order Comatulida AH. CLI\RK, 1908) is reported for the first time from Upper Jurassic sequences of Poland, precisely from Lower Kimmeridgian strata of the Holy Cross Mountains. The major part of this assemblage comes from oolitic deposits exposed at Malogoszcz Quarry, others from oyster (Actinostreon, and Nanogyra) shellbeds higher up section at Malogoszcz, as well as from the coeval strata of the Karsy section. Taxonomically recognizable skeleton elements such as calyces, isolated een­ trodorsals and radials are here assigned to seven taxa, three of which are new to science: Comatulina malogoslialla sp.nov., Palaeocomaster learsensis sp.nov., and Solanocrinites sanctacruccnsis sp.nov. The majority of the material available was contained in burrows made by some ancestral stock of alpheid shrimp, closely comparable to those of present-day snapping shrimp (genus Alpheus WEBER, 1795), and its allies. These burrows, situated at the tops of oolitic shoals/banks at Malogoszcz, casually served both as habitats of cryptic faunas (mostly comatulids, dwarf-sized gastropods) and as preservational/taphonomic traps for others, primarily echinoderms (ten taxa of echinoids, three stalked crinoids, two ophiuroids, one asteroid) swept into by highly agitated waters, most likely during storms, to pro­ duce an Echinodermenlagerstdlte. Comatulid remains from the oyster shellbeds undelwent longer periods of trans­ port, to be entombed far from their habitats.

Keywords: Crinoidea, Comatulida, Taxonomy, New Species, Eco-taphonomy, Upper Jurassic, Poland.

INTRODUCTION in recent years: the free-living crinoids of the comatulid order, or feather stars, as commonly referred to in the lit­ The Upper Jurassic shallow-marine carbonate erature (see HENDLER & al. 1995, MESSING 1997). These sequence of the Holy Cross Mountains, Central Poland, tiny animals, mostly hidden from view in present-day seas has long been known to yield ubiquitous fossils (see KUTEK due to their nocturnal mode of life, have long been un­ 1968,1969; MATYJA 1991; KUTEK &al. 1992a, b). Amongst recognized also in Jurassic deposits of Poland. The situa­ echinoderms, echinoids have first been taken into investi­ tion changed when an intense search was done in the gation by the present author (RADWANSKA 1999, 2003a, Malogoszcz sequence of Kimmeridgian age (see KUTEK 2004a, b), on the basis of material for a large part coming & al. 1992b; RADWANSKI 1995; RADWANSKA 1999; from huge exposures of cement works at Malogoszcz, RADWANSKA & RADWANSKI 2003, 2004a, b, 2005). established in the mid-1970s. Comatulid crinoids from the Lower Kimmeridgian The present report deals with one of the more fasci­ portion of the sequence exposed at Malogoszcz, first nating discoveries of Late Jurassic echinoderms in Poland recorded by RADwANSKJ (1995, p. 12), have recently been 270 URSZULA RADWANSKA the subject of to a preliminaIY account by the present sequences of Poland. Previous records of comatulid author (RADWANSKA 2004b). That account concerned crinoids in Poland only involved those from Middle remains of comatulid crinoids from the Upper Oolite Miocene (Badenian) sequences along the southern Member, where they are confined to tiered burrows of slopes of the Holy Cross Mountains: the Piilczow alpheid shrimp (see RAOWANSKA & RADWANSKI 2004b). Limestone (RAOWANSKlI977, p. 747 and fig. 172/8) and Apart from this location, eomatulids also occur in the the Korytnica Clays (RAOWANSKA 1987, 2003b). marly intervals of oyster coquinas (Actinostreon/Nanogyra shellbeds) higher up section at Malogoszcz (see Text-fig. SYSTEMATIC ACCOUNT 1), and in coeval beds sequence exposed at Karsy near Korytnica, some 30 km away (see RADWANSKA 1999, pp. The taxonomy and terminology of comatulids used 297-298; MACHALSKI 1983; KIN 2005). here follows that proposed by Ti'eatise on Invertebrate These two localities, Malogoszez and Karsy, have as Paleontology (WIENBERG-RAsMUSSEN 1978; see com­ yet remained the only sites of the eomatulid occurrence ments by MESSING 1997, p. 4). throughout all Upper Jurassic and other Mesozoic Repositories

Malogoszcz Quarry All comatulid material described here is housed in the Lithostratigraphic unit Department of Palaeontology, Faculty of Geology, Sandstone University of Warsaw. It is kept under the collection num­ bers preceded by the character C (comatulids), followed by a lettered symbol Km indicative of its Kimmeridgian age. Acanthicum Top Shales

Order Comatulida AH. CLARK, 1908 Superfamily Solanocrinitacea JAEKEL, 1918 Family Solanocrinitidae JAEKEL, 1918

Upper Platy Limestones Divisum Genus Comatulina O'ORBIGNY, 1852

TYPE SPECIES: Solanocri.ni.tes costatus GOLDFUSS, coquinas 1829; OD

REMARKS: The genus Comatulina O'ORBIGNY, 1852,

Shaly Llmeslones and is diagnosed (see WIENBERG RASMUSSEN 1978, p. T875) Underlying Shales as featured by centro dorsals truncated conical to trun­ Hypselocyclum cated subhemispherical, of dorsal (aboral) side flat or ~g Oolite-Platy member concave, and by closely placed cirrus sockets arranged in more than 10 (generally 11-15, exceptionally up to 20) columns in adult specimens. In the material studied, such features are shared by four taxa, the centro dorsals of which range in shape Banded Limestone member from conically hemispherical (C malogostiana), truncat­ Hypselocyclum :J ed conical (C peroni, C beltremieuxi), through to almost and/or Lower OolilA low cylindrical (classified below as Comatulina sp). Platynota Underlying Pelitic limestones

Fig. 1. Lower to low-Upper Kimmeridgian seqnence exposed at Comatulina peroni. (DE LORIOL, 1889) Malogoszcz Quarry in the Holy Cross Mountains, Central Poland (adapt­ (Text-fig. 2 and PI. 1, Figs 2-4) cd from: KUTEK & ai. 1992b, fig. 5; MATYJi\ & WIERZBOWSKI 1996, fig. 3;

RADWANSK;\ & RADWAKSKI 2005, fig. 2), to show occurrence sites 1889. Alltedoll Peroni P. DE LORIOL, 1888; P. DE LORIOL, pp. (arrowed) of comatulids studied from oolitic shoals (Upper Oolite 490-492, pI. 222, figs 2, 2a-2c. Member) and onkolitie, marly oyster-coquinas (Actinostreon/Nanogyra 2004b. Comatulina peroni (DE LORIOL, 1889); u. RADWANSKA, sheIIbeds of the Skork6w LumaeheIIe Member) p. 126, pI. 2, fig. 11. KIMMERIDGIAN COMATULID CRINOIDS 271

1888". Due to the general shape of the centrodorsal and the number of columns of cirrus sockets, they are assigned to the genus Comatulina D'ORBIGNY, 1852. The studied calyces of Comatulina peroni display a slight variability in the steepness and height of the artic­ ular facet of the radials (see PI. 1, Figs 2, 4b). Moreover, their aboral side is relatively flat widely.

Comatulina cf. beltremieuxi (DE LORIOL, 1889) (PI. 1, Figs 1a-1c)

MATERIAL: Three centrodorsals and a single radial plate; all from tiered burrows of alpheid shrimp at Malogoszcz (Upper Oolite Member). b DESCRIPTION: The centro dorsals are moderately tall, truncated conical, with 15 columns of one to three close­ Fig. 2. Comallilina peroni (DE LORIOl., 1889); a - lateral (interrradial) ly spaced, large eirrus sockets (PI. 1 , Fig. 1b). The abo­ view of calyx, to show basal plate; b - lateral (radial) view of calyx, to ral side of the centro dorsal is slightly concave and cov­ show radial plate; CKm/008; x 7 ered by furrows (PI. 1, Fig. la). Cirrus soekets are oval in outline and deep, with a large lumen. The adoral side MATERIAL: Four calyces and five centrodorsals; all of the centro dorsal has a very narrow eavity, about 17% from tiered burrows of alpheid shrimp at Malogoszcz of eentrodorsal diameter. Basals are not observable. (Upper Oolite Member). The radial plate is trapezoidal, with low free aboral sur­ face, covered by prominent granules, arranged in a sin­ DESCRIPTION: The centrodorsais are moderately tall, gle horizontal row (PI. 1, Fig. lc). The articular facet of truncated conical to almost discoidal, with 15 columns the radials is moderately tall and steep. The aboralliga­ of three large, closely spaced, cirrus sockets (Text-figs ment fossa is low, with a large, oval ligament pit. 2a-2b and PI. 1, Figs 2, 3a, 4a-4b); in one speci­ Interarticular ligament fossae are low, and triangular in men, the median column bifurcates to yield four sockets, shape. Adoral muscular fossae are triangular. what amounts 46 sockets in total (see Text-fig. 2a and PI. 1, Fig. 4a). The aboral side of the centro dorsals is rela­ REMARKS: The studied centrodorsal and the radial tively large (about 75% of centro dorsal diameter), flat plate are very close to those of the species Comatulina or slightly eoncave, covered by granules (PI. 1, Fig. 3b). beltremieuxi (DE LORIOL, 1889), described from the Cirrus soekets are oval in outline and deep, with a large Kimmeridgian of France. Alternatively, the single radi­ lumen. The adoral side of centro dorsal has a shallow al plate might be assigned to Solanocrinites jutieri (DE cavity, about 25% of centrodorsal diameter (PI. 1, LORIOL, 1879), from the Lower Kimmeridgian of Fig. 3a). Basals are distinctly exposed at interradial France (see DE LORIOL 1879, pp. 265-266, pI. 21, figs I­ points (Text-fig. 2a and PI. 1, Fig. 4a). The radial ring S), but no centrodorsal which may be ascribed to that is lower than the slightly overhanging eentrodorsal (PI. 1, species has yet been found in the investigated material. Figs 4a-4b). Radials are trapezoidal, with a very low The poor preservation of the collected specimens (iso­ free aboral surface, and distinctly undulated aboral lated centro dorsal and radial plate, both with corroded margin (Text-figs 2a-2b and PI. 1, Figs 2, 3a, 4b). The surface) does not permit a preeise specific determina­ articular facet of radials is moderately tall and oblique tion, contr31Y to the previous statement (RADWANSKA (PI. 1, Figs 2, 3a, 4a-4b). The aboral ligament fossa is 2004b, pI. 2, figs 3-4). low, with a large, oval ligament pit. Interarticular liga­ ment fossae are triangular in shape. Adoral muscular fossae are arc-shaped (Text-fig. 2b and PI. 1, Fig. 4b). Comatulina malogostiana sp.nov. (Text-fig. 3 and PI. 2, Figs 1-3) REMARKS: The studied ealyces are concordant with those described from the Kimmeridgian of France by 2004b. Comafulina costa fa (GOLDFUSS, 1829); u. RADWANSKA, DE LORIOL (1889) as "Antedon peroni P. DE LORIOL, p. 126, pI. 2, fig. 6. 272 URSZULA RADWANSKA

DESCRIPTION: The centro dorsals are moderately tall, conically hemispherical, with 15 columns of a single to three closely spaced cirrus sockets (Text-figs 3a-3b and PI. 2, Figs 2a-2b, 3b-3c). The aboral side of centrodor­ sals is pointed with a single, centrally placed cirrus sock­ et surrounded by five small ones (PI. 2, Figs 1b, 2c, 3d). Cirrus sockets are oval in outline and deep, with a small lumen; their number is 41 in the largest specimen. Relatively distinct basals, exposed in interradial points, are rhomboidal in cross-section (Text-fig. 3a and PI. 2, Fig. 3b). The radial ring is slightly taller than centrodor­ sal, slightly overhanging (PI. 2, Figs 2a, 2b, 3b). Radials are trapezoidal, with a very high free aboral surface (PI. 2, Fig. 3b) The articular facet of radials is moderately tall and steep at 4SO. The aboral ligament fossa is large with a long, deep ligament pit. Interarticular ligament fossae are tall, and triangular in shape. Adoral muscular fossae are low, and arc-shaped (Text-fig. 3b and PI. 2, b Fig. 3c). The radial cavity is moderately large and deep, reaching about 50% of calyx diameter (PI. 2, Fig. 3a).

Fig. 3. Comatuiil1(1 maiogostial1(1 sp.nov.; holotype: a -lateral (interradial) REMARKS: The calyces studied are assigned to the view of calyx, to show basal plate; b - lateral (radial) view of calyx, genus Comatulina D'ORBIGNY, 1852. The newly estab­ to show radial plate; CKm/OOl; x 7 lished species Comatulina malogostiana sp.nov. differs from the closely related species, C. costata (GOLDFUSS, HOLOTYPE: Calyx (specimen number CKm/OOl), 1829), in having a lower centro dorsal, a smaller number presented in Text-fig. 3 and PI. 2, Figs 3a-3d. of cirrus sockets, and a distinctly taller free aboral sur­ face of radials. From other congeners, e.g. C. peroni (DE PARATYPES: Centrodorsal (specimen number CKm/003) LORIOL, 1889), it differs in its more hemispherical shape pressented in PI. 2, Fig. 1 and calyx (specimen numbcr of centrodorsal, and in the more prominent basals. CKm/002), presented in PI. 2, Fig. 2.

TYPE LOCALITY: Malogoszcz Quarry, south-western Comatulina sp. margin of the Holy Cross Mountains, Central Poland. (PI. 3, Figs 1a-1b)

TYPE HORIZON: Low Kimmeridgian, Katroliceras divi­ MATERIAL: Two centro dorsals, from Karsy. sum Zone; Skork6w Lumachelle Member. DESCRIPTION: The large centrodorsals (the larger DERIVATION OF THE NAME: Adjectival name, from reaching 13 mm in diameter) arc low cylindrical, with neo-Latinized name of the medieval town of Malogoszcz. up to 17 columns of two or three, closely spaced, large cirrus sockets (PI. 3, Fig. 1a). The aboral side of cen­ MATERIAL: The holotype calyx from oyster coquinas at trodorsal is large, flattened and slightly concave, cov­ Malogoszcz; the other three calyces and six centro dorsals ered by radiating furrows. Cirrus sockets arc large, oval from tiered bUlTOWS of alpheid shrimp at Malogoszcz. in outline and deeply incised, with a large lumen; their number totals 49 in the larger specimen. The adoral DIAGNOSIS: Centro dorsal conically hemispherical, side of centro dorsal has a very narrow cavity, about with 15 columns of one to three large, closely spaced 14% of centro dorsal diameter (PI. 3, Fig. 1b). cirrus sockets; aboral side of the centro dorsal pointed with one, centrally placed cirrus socket surrounded by REMARKS: The studied centro dorsals arc assigned to five small ones; basals exposed in interradial points, the genus Coma/ulina D'ORBIGNY, 1852. In view of their rhomboidal in cross-section; radial ring a bit taller than size, number and arrangement of cirrus sockets, they centrodorsal, slightly overhanging; radials trapezoidal, probably belong to the species Comatulina lambel1i DE with a very tall free aboral surface. LoRIOL, 1889, described from the Lower Kimmeridgian of KIMMERIDGIAN COMATULID CRINOIDS 273

France (see DE LoRlOL 1889, pp. 515-519, pI. 225, figs 2-4), MATERIAL: One calyx and a single centrodorsal; both but because other ossicles (basals and radials) are lacking, from Karsy. they are assigned to the genus level only. Amongst the congeners, these centrodorsals are extremely wide. DIAGNOSIS: Centrodorsal large, discoidal with numerous (c. 50) deeply incised, relatively small cirrus sockets; basals invisible; radial ring low, not overhang­ Genus Palaeocomaster GISLEN, 1924 ing; radial plates with very low free aboral surface, articular facet fairly steep at 35°. TYPE SPECIES: Actinometra guirandi DE LORIOL, 1889; OD DESCRIPTION: The large centro dorsals (the larger reaching 14.5 mm in diameter) are relatively low, slight­ ly truncated conical to almost discoidal, with closely Palaeocomaster karsensis sp.nov. spaced, small cirrus sockets, not well arranged in (Text-fig. 4 and PI. 3, Figs 2a-2e) columns (Text-figs 4a-4b and PI. 3, Figs 2a-2b). The abo­ ral side of centro dorsal is large, and concave, covered by HOLOTYPE: Calyx (specimen number CKm/011), radial furrows (PI. 3, Fig. 2e). Cirrus sockets are small, presented in Text-fig. 4 and PI. 3, Figs 2a-2e. deeply incised, with a small lumen; their number is about 50 in the larger specimen. Basals are invisible (Text-fig. PARATYPE: Centrodorsal (specimen number 4a and PI. 3, Fig. 2a). Radial ring is low, not overhanging CKm/014). (PI. 3, Figs 2a-2c). Radials are trapezoidal, with very low aboral surface (Text-fig. 4b and PI. 3, Fig. 2c). The articu­ TYPE LOCALITY: Karsy, south-western margin of lar facet of radials is moderately tall and fairly steep at the Holy Cross Mountains, Central Poland. 35° (PI. 3, Figs 2a-2b). The aboral ligament fossa is large with a long, deep ligament pit. Interarticular ligament TYPE HORIZON: Lowcr Kimmcridgian, Katroliceras fossae are moderately tall, and triangular in shape. divisum Zone; Skork6w Lumachelle Member. Adoral muscular fossac are low, and arc-shaped. The radial cavity is moderately large and deep, reaching DERIVATION OF THE NAME: Adjectival namc, about 50% of calyx diameter (PI. 3, Fig. 2d). from neo-Latinized name of the will age of Karsy. REMARKS: The studied specimens with their almost discoidal shape of centrodorsal and crowded cirrus sockets, not forming distinct columns, are assigned to the genus Palaeocomaster GISLEN, 1924. The newly established species Palaeocomaster karsensis sp.nov. differs from the most similar species Palaeocomaster vagnasensis (DE LORIOL, 1889) by having more numer­ ous but smaller cirrus sockets, and by showing distinct­ ly steep articular facets of radials. From the type species, Palaeocomaster guirandi (DE LORIOL, 1889) described from the Oxfordian of France (see DE LORIOL 1889, pp. 535-537, pI. 227, figs 2, 2a-2c), it differs by the more sloping articular facets of radial plates and in the absence of exposed basals.

Palaeocomaster sp. (PI. 1, Figs 5a-5b)

b 2004b. Palaeocol11aster vagnasensis (DE LORIOL, 1889); U. RADWANSKA, p. 126, pI. 2, fig. 12. Fig. 4. Palaeocomaster karsensis sp.nov.; holotype: a - lateral (interra­ dial) view of calyx, to show a lack of basal plate; b -lateral (radial) MATERIAL: A single centrodorsal, from tiered burrows view of calyx, to show radial plate; CKm/011; x 7 of alpheid shrimp at Malogoszcz (Upper Oolite Member). 274 URSZULA RADWANSKA

DESCRIPTION: The centro dorsal is relatively large PARATYPE: Calyx (specimen number CKm/013), pre­ (diameter reaching 10 mm), pentagonal in outline, low sented in Text-fig. Sc and PI. 4, Figs 2a-2b. discoidal, with irregularly arranged cirrus sockets, which do not form distinct columns (PI. 1, Fig. Sb). The TYPE LOCALITY: Malogoszcz Quarry, south-western aboral side of centrodorsal is large, flattened and slight­ margin of the Holy Cross Mountains, Central Poland. ly concave, covered by radial furrows. Cirrus sockets are large, oval in outline, and relatively deeply incised; their TYPE HORIZON: Lower Kimmeridgian, Ataxioceras number is 34. Six small cirrus sockets are developed on hypselocyclum Zone; Upper Oolite Member. adoral side of centro dorsal (PI. 1, Fig. Sa). DERIVATION OF THE NAME: Adjective sanctacrucen­ REMARKS: The studied specimen with its low dis­ sis - neo-Latinized, in reference to the Holy Cross region. coidal shape and crowded cirrus sockets, not forming distinct columns, is assigned to the genus Palaeo­ MATERIAL: Six calyces, four centrodorsals, plus a sin­ comaster GISLEN, 1924. In view of its size, and general gle radial plate; all from tiered burrows of alpheid shape, as well as number and arrangement of cirrus shrimp at Malogoszcz. sockets, it probably belongs to Palaeocomaster vagnasensis (DE LORIOL, 1889), described from the DIAGNOSIS: Centro dorsal truncated conical, with 10 Jurassic of France (see DE LORIOL 1889, pp. S38-S40, pI. columns of three, closely spaced, large cirrus sockets; 227, figs 3, 3a-3b), but through a lack of other plates aboral side of centro dorsal sharply pointed; radial ring (basals and radials), it is assigned to the genus level slightly lower than centrodorsal, distinctly overhanging; only. From Palaeocomaster karsensis sp.nov. it differs in radials low-trapezoidal, with moderately large, free its lesser height, in being more pentagonal in outline, aboral surface. and in having cirrus sockets conspicuously larger. DESCRIPTION: The centrodorsals are truncated coni­ cal, with 10 columns of three, closely spaced cirrus sock­ Genus Solanocrinites GOLDFUSS, 1829 ets (Text-fig. S and PI. 4, Figs Id, 2b). The aboral side of centro dorsals is sharply pointed (PI. 4, Fig. Id). Cirrus TYPE SPECIES: Solanocrinites costatus GOLDFUSS, sockets are large, oval in outline with a large lumen. 1829; SD DE LORIOL (1889, p. S26). Basals are inconspicuous (Text-figs Sa, Sc and PI. 4, Figs 1b, 2a-2b). The radial ring is slightly lower than centro dorsal, distinctly overhanging (PI. 4, Fig 1d, 2b). Radials are low Solanocrinites sanctacrucensis sp.nov. trapezoidal, with moderately large aboral surface (PI. 4, (Text-fig. S and PI. 4, Figs 1-2) Figs 1a-1b, 1d, 2a). The articular facet ofradials is rela­ tively steep (PI. 4, Fig. 1d). The aboral ligament fossa 2004b. Solanocrinites sp.; U. RADWANSKA, p. 126, pi. 2, figs is large. Interarticular ligament fossae are moderately Sa-Sb. tall and triangular in outline (Text-fig. Sb-Sc and PI. 4, Figs la, 1d). Adoral muscular fossae are low, arc-shaped HOLOTYPE: Calyx (specimen number CKm/012), (PI. 4, Fig. 1a). The radial cavity is deep and large, reach­ presented in Text-figs Sa-Sb and PI. 4, Figs 1a-1d. ing 80% of calyx diameter (PI. 4, Fig. lc).

Fig. 5. Soianocrinites sanctaclUcensis sp.nov.; a-b - holotype, a - lateral (interradial) view of calyx, to show basal plate; b - lateral (radial) view of calyx, to show radial plate; CKm/012; x 10; c - paratype, lateral (interradial) view of calyx; CKm/013; x 10 KIMMERIDGIAN COMATULID CRINOIDS 275

REMARKS: The studied calyces are assigned to the comatulids living in shallow-water habitats (see genus Solanocrinites GOLDFUSS, 1829. The newly estab­ MLADENOV 1983). lished species Solanocrinites sanctacrucensis sp.nov. dif­ fers from the two most similar species, S. lambeltsi Echinoids. A typical feature of echinoid remains col­ SIEVERTS-DoRECK, 1958, and S. costatus (GOLDFUSS, lected is the relatively high frequency of small-sized, 1829), in having a more conical centrodorsal, a distinct­ complete tests (see Text-fig. 8.10-14). In comparison to ly overhanging radial ring, and by showing more the size of conspecific specimens from other parts of depressed articular facet of radials. the Malogoszcz sequence (see RAoWANSKA 1999), these may be interpreted mostly as juveniles (see Text-fig. 8.10 and 8.13). One species, Pseudosalenia malogostiana ECO-TAPHONOMY RADWANSKA, 1999, represented by both smaller and larger specimens, interpreted herein as juveniles and The majority of the comatulids studied come from adults (see Text-fig. 8.11-12), is generally small-sized one location of the alpheid burrows in oolite banks com­ and has to date only been recorded from alpheid bur­ posing the Upper Oolite Member at Malogoszcz. An rows of the Upper Oolite Member. These burrows may abundance of various echinoderms contained in these thus be regarded as habitats for a crevice fauna of some burrows, and their wide-ranging mode of preservation, echinoids but, on the other hand, as preservational justifies eco-taphonomic analysis, to recognize the path­ traps, the small openings of which (4-5 cm in diameter) ways the comatulids and associated echinoderms have functioned as a taphonomic filter for any skeleton undergone from life to final burial. The alpheid burrows swept over. A significant number of juvenile specimens of Malogoszcz were long enough exposed with their is a feature typifying their assemblage as a thana to­ openings at the sediment/water interface to become coenosis, that was formed by catastrophic death of a habitats for more or less cryptic faunas, and deadly traps live community. It is suggested that storm agitation is for others (see Text-fig. 6). The burrowed surface then the most probable agent that killed the animals on the evolved from soft to firm (see Text-fig. 6A), and finally oolitic shoals studied. to the hard-bottom type (see Text-fig. 6B). Following a proposal of SMITH (1984) and results All echinoderm remains contained in the studied from taphonomic analyses of other echinoids, both pre­ alpheid burrows (see Text-figs 7-8) are slightly corroded sent-day (NEBELSICK 1992) and fossil (GORDON & at their surface (? erosionally worn), and some are sili­ DONOVAN 1992), six classes of echinoid preservation are cified to varying extent. Their mode of disarticulation here selected, to be briefly characterized in successive and inferred taphonomic interpretation may briefly be order of their taphonomic significance. characterized, as follows. (i) Tests, lacking spines and Aristotle's lantern, with or without apical disc, always empty (see Text-fig. 8.11- Comatnlid crinoids. An ubiquity of comatulid remains 13), represent live specimens (see also ASLIN 1968) of which only complete calyces, centro dorsals, and radi­ swept into burrows, where their corpses decom­ al plates are diagnostic (see Pis 1-4; and WIENBERG posed. Then, the burrows could still be empty or, RASMUSSEN 1978, MESSING 1997), is thought to repre­ possibly, inhabited by their alpheid producers. sent a case of live specimens, skeletons of which disin­ (ii) Tests such as above, but partially filled with ooids tegrated into component ossicles wthin the alpheid bur­ (see Text-fig. 8.10), represent live specimens mor­ rows. In these, the comatulids either lived cryptically, tally entrapped in burrows when the oolitic sedi­ having been curled-up at daytime (comp. MESSING ment had been swept into emptied tests. 1997, p. 18), or had been swept in with sediment by (iii) Tests partlially filled with sediment, but with their which they were suffocated, buried and disarticulated spine canopy in position (see Text-fig. 8.14), repre­ after their death. Of isolated ossicles, in majority undis­ sent live specimens violently swept into burrows, tinguishable in their taxonomy (radials, brachials, pin­ together with a bulk of sediment which buried nulars, cirrals), only some radials may specifically be them, and infilled tests when corpses decomposed. recognized (see PI. 1, Fig. lc). Hundreds of isolated (iv) Test fragments broken across component plates ossicles other than centro dorsals and/or complete (interambulacral in particular) represent those calyces suggest that dead crinoids within the burrows swept into burrows when their collagenous fibres disarticulated rapidly, in terms even of a few days (see were still intact; such fragments are interpreted as DONOVAN 1991, pp. 245 and 254; MESSING 1997, p. 21). coming from damaged tests of live specimens that A part of disarticulated ossicles may have resulted from were crushed by water agitation rather than by arm autotomy, a process important in present-day predators. 276 URSZULA RADWANSKA

(v) Spines, or their fragments, isolated teeth and other bodies/corpses swept violently into burrows, or an pieces of Aristotle's lantern, disarticulated plates of allochthonous detritus for which the burrows test, or larger fragments of the latter, represent became sedimentary traps. either further steps in the disarticulation process of (vi) Isolated copepod cysts, composed of echinoid cal- A

B KIMMERIDGIAN COMATULID CRINOIDS 277

cite, some with echinoid test fragments (see (SCHLOTHEIM, 1820) whose longer stem fragments and RADWANSKA & RAoWANSKI 2005), suggest a weak­ holdfasts are present (see Text-fig. 8.8-9), and as large­ ened structure of echinoid plates to which they sized specimens attributable to the genus Liliocrinus adhered and, moreover, a relatively high parasite (see Text-fig. 8.6-7) and/or Apiocrinites and MillCl'i­ pressure of cope pods upon echinoids from the crinus. Of the latter, a stem fragment infested by a Upper Oolite habitats. myzostomidan polychaete is discussed separately (RADWANSKA & RADWANSKI 2005), and a large hold­ Stalked crinoids. The most common remains of echino­ fast of Apiocrinites grazed by echinoids to produce derms in the alpheid burrows are from stalked crinoids, represented largely by isolated columnals, pluricolum­ COMATULID CRINOIDS Material naIs, holdfasts, as well as cirrals and radials, the latter not CC CD R distinguishable from those of comatulids. Nevertheless, 1 Comatulina peroni 4 5 - (DE LORIOL, 1889) two ecotypes of stalked crinoids are recognized (cf. 2 Comatulina cf. beltrel11ieuxi - 3 1 KLIKUSHIN 1996), viz. (i) long-stemmed ones, adorned (DE LORIOL, 1889) with cirri, living 'freely' on soft bottoms, and (ii) long- or 3 Comatulina l11alogostiana Sp.Il. 4 6 - short-stemmed ones, devoid of cirri, cemented to a hard 4 Palaeocomaster sp. - 1 - 5 Solanocrinites sanctacrllcensis sp.n. 6 4 1 substrate (organic skeletons, hardground). ECHINOIDS Material The first ecotype is represented (see Text-fig. 8.4-5) Tc Tf Sp by columnals of IsoC/inus amblyscalaris (THURMANN, 6 Rhabdocidaris orbignyana - 4 26 1861), the species according to HESS (1975a, p. 55) syn­ (L. AGASSIZ, 1840) 7 Plegiocidaris cruci/era - - 3 onymous with crowns of 1. pendulus VON MEYER, which (L. AGASSIZ, 1840) lived more or less freely, having been either 8 Acrosalenia anglliaris 2 1 - anchored/stilted with its cirri in a stable place (see HESS (L. AGASSIZ, 1840) 1975a, fig. 8; 1999a, p. 209 and fig. 212; 1999b, figs 235- 9 Pselldosalenia malogostiana 2 - - RADW ANSKA, 1999 236), or supported by cirri distributed along a longer 10 Hemicidaris interl71edia 2 44 22 stem that could float over the bottom (see WIENBERG (FLEMING, 1828) RASMUSSEN 1977, fig. 2; KLIKUSHIN 1996, fig. 3). Some 11 Pseudocidaris sanctacrllcensis - 10 present-day isocrinids are even able to relocate actively RADW ANSKA, 1999 12 Trochotiara kongieli 2 - - when irritated (see MESSING & al. 1988). The studied RADW ANSKA, 1999 species had certainly the ability to autotomize its parts 13 Phymosoma sllpracorallinllm 1 -- (see MLADENOV 1983, OJI 2001, On & OKAMOTO 1994) (COTTEAU, 1865) 14 Polycyphus distinctlls - 3 - when seriously disturbed by hydrodynamic agents (L. AGASSIZ, 1840) and/or predators. Its remains in the alpheid burrows 15 Pygaster l110rrisi - 3 should thus be regarded as having been swept into WRIGHT, 1851 those from more distant surroundings. STALKED CRINOIDS Material St Co H The second ecotype is characterized by smaller­ 16 Isocrinus amblyscalaris 23 6 - sized species, such as Angulocrinus echinatus (THURMANN,1861) 17 Angulocrinus echinatus 6 - 2 Fig. 6. A unique site of the Echil1odennenlagal·tiitte in the Lower (SCHLOTHEIM,1820) Kimmeridgian oolitic banks (Upper Oolite Member) at Malogoszcz. 18 LiliocrinlislApiocrinites sP. 55 10 Ii OPHIUROIDS Material A- Soft -bottom stage: tiered burrows of alpheid shrimp in highly bio­ Bv Ba turbated oolite become refuges for cryptic forms (living and/or hiding 19 Ophioderma? spectabilis - 12 at daytime), and taphonomic traps for others, either alive or dead, HESS, 1966 swept into them by agitated waters. B - Hard-bottom stage: the ero­ 20 Opi1iolJctra? oerflii HESS, 1966 4 - ASTEROIDS Material sionally truncated and lithified burrows of alpheids are still open at the Sm Sa sediment/water interface and encrusted by stalked crinoids and oysters, 21 Pentasteria longispina HESS, 1968 10 1 I to become the taphonomic traps for all biota (living and dead speci­ mens, disarticulated remains) swept in by agitated waters of storm ori­ Fig. 7. Echinoderm remains contained in alpheid burrows; Upper Oolite gin that had sabred or uprooted some stalked crinoids. The echino­ Member at Malogoszcz denn remains are distinguished as follows: f - Comatulid crinoids, Comatulid crinoids: CC - calyx, CD - eentrodorsal, R - radial; Echinoids: e - Empty tests of echinoids, s - Echinoid tests with attached spines, Te - complete test, Tf - test fragment, Sp - spine; Stalked crinoids: t - Columnalsipluricolumnals of stalked crinoids, h - Echinoderm hash Sf - stem fragment, Co - eolumnal, H - holdfast; Ophiuroids: Bv - vertebra, of isolated ossicles (crinoids, echinoids, ophiuroids. asteroids) Ba - ann plate; Asteroids: Sm - marginal ossicle, Sa - ambulacral o,,-,ide 278 URSZULA RADWANSKA traces Gnathichnus pentax BROMLEY, 1975, from ENVIRONMENTAL CONCLUSIONS another, hardgrounded ooid bed of the Upper Oolite Member, was presented earlier (RADWANSKA 1999, The studied assemblage of comatulid crinoids from pp. 355-356 and fig. l1.1-la). All remains of these Malogoszcz and Karsy comprises 39 calyces and/or cen­ massively stemmed crinoids (AnguloCl1nus, LilioCl1nus, trodorsals of seven taxa. Compared to these from mod­ Apiocrinites) are thought to have originated by storm ern habitats in the tropical Indo-Pacific the studied destruction of specimens that lived nearby having assemblage is very poor, both taxonomically and been cemented presumably to the hardgrounded por­ regarding the population density (see D.L. MEYER & tions of oolitic banks. Stronger agitation caused not MACURDA 1977: 26 species at a single locality at Lizard only breakage of some specimens into columnar parts, Island on the Great Barrier Reef; MACURDA & D.L. but also tearing-out ('uprooting') of their holdfasts MEYER 1983: 14 species on a coral block from that (see Text-fig. 8.9), which all were finally swept into the Lizard Island; see also MESSING 1997, p. 4); densely alpheid burrows. populated are also the Red Sea habitats (see MAGNUS 1963: up to 700 specimens on 1m2). The studied assem­ Ophiuroids. Isolated plates of ophiuroids comprise blage is close to the values for the Caribbean (see numerous vertebrae and various arm plates. Of these, HENDLER & al. 1995, p. 44: 8 species altogether). only some may be recognized taxonomically (see Text­ Nonetheless, the life habits and trophic requirements of fig. 8.2-3), to represent the two species established by the studied comatulids were probably similar to those HESS (1966): Ophioderma? spectabilis HESS, and domiciled in modern tropical/subtropical shallow Ophiopetra? oeltlii HESS. All ophiuroid ossicles, relative­ waters, with bottom topography full of nooks and/or ly well preserved, are presumably remains of specimens crevices favouring the development of cryptic refuges (live or dead) swept into burrows where their bodies/ for comatulids which preferred a nocturnal mode of skeletons decomposed. life. To note, this is the first occurrence site of identifi­ In the Upper Oolite Member at Malogoszcz, able Late Jurassic ophiuroids in Poland (see alpheid burrows formed such refuges, lithified more or RADWANSKA 2004b, pp. 127-128). In other countries, less early, and open at the sediment/water interface on notable is a mass occurrence of ophiuroid isolated ossi­ extreme shallow-water and/or emergent oolitic shoals cles in some Oxfordian strata of Switzerland (see HESS (comp. FURSICH & PALMER 1975). In these burrows, 1975b). comatulids could hide at daytime, and they could live there as long as the burrows' openings remained free, Asteroids. Isolated plates of asteroids, largely marginals not having been clogged by algae or sediment. (see Text-fig. 8.1a-1b) and ambulacral plates (see Associated to comatulids were dwarf-sized gastropods, RADWANSKA 2004b, pI. 2, fig. 8) are assigned to whose taxonomy remains as yet unclear (see Pentasteria longispina HESS, 1968, a species whose com­ RADWANSKA 2004b). The alpheid burrows discussed are plete skeletons are recorded from Oxfordian strata of thus interpreted as habitats for a selected crevice fauna Switzerland (see HESS 1968, 1975a; c.A. MEYER 1984). (comp. PALMER & FURSICH 1974). More or less complete skeletons of this species have Moreover, the alpheid burrows at Malogoszcz have recently been also found in oolite beds beyond the also acted as preservational traps for other echino­ alpheid burrows at Malogoszcz (RADWANSKA, unpub­ derms, the assemblage of which comprises ten echinoid lished). Because of the larger size of live specimens of taxa, three stalked crinoids, two ophiuroids, and one this species it is thought that their ossicles were swept asteroid (see Text-figs 7-8). Their diverse remains, in into the burrows after the disarticulation of the skele­ varying states of preservation (quite fresh, or heavily tons somewhere nearby. worn), indicate a multi-phase delivery into the burrows,

Fig. 8. Echinoderm associates of comatulid crinoids from alpheid burrows; Upper Oolite Member at Malogoszcz. Asteroids:Pentastelia iongispina HESS, 1968: la - Marginal ossicle in lateral view, 1b - same, in outer view; both x 7. Ophiuroids: 2 - Ophiopetra? oel1lii HESS, 1966, vertebra in proximal view, x 10; 3 - Ophiodel111a? spectabilis HESS, 1966, arm plate, x 10. Stalked crinoids: [soClinus amblyscaimis (THURMANN, 1861): 4 - Columnal (nodal), articular facet; 5 - Pluricolumnal composed of three internodals and one nodal (arrowed is a cirrus socket); both x 7; Liliocrinus sp.: 6 - Longer fragment of column, nat. size; 7 - Columnal, articular facet, x 5; Anguiocrinlls echinatus (SCHLOTHEIM, 1820): 8 - Longer fragment of column, with WaIt-like tubercles, 9 - Holdfast of a juvenile specimen, top view; both x 5. Echinoids: 10 - Phymosoma supracorallinwn (C01TEAU, 1865), test of a juvenile specimen, partially infilled with ooids, x 5; Pseudosalenia maiogostialla RADwANsKA, 1999: 11 - Juvenile specimen, oral view of empty test; 12 - Empty test of an adult specimen, aboral view; both x 10; Acrosaienia anguiwis (L. AGASSIZ, 1840): l3 - Empty test of a juvenile specimen, aboral view; 14 - Test partially infilled, with spine canopy preserved, aboral view; both x 5 KIMMERIDGIAN COMATULID CRINOIDS 279 280 URSZULA RADWANSKA most likely by storm-induced agitated water sweeping (DE LORIOL 1889, GISLEN 1924, SIEVERTS-DoRECK all its load into the burrows. The alpheid burrows were 1958), and Portugal (DE LORIOL 1880, 1891). These thus not only refuges for, but also preservational traps rather scarce occurrences of Jurassic comatulid crinoids of, all echinoderms that inhabited oolitic shoals. This is are thus in their frequency clearly different from those especially true of the smaller-sized echinoids, some of ageing Late Cretaceous (see WIENBERG RASMUSSEN which possess spine canopy in position, others having 1978, JAGT 1999) or Neogene (SIEVERTS-DoRECK 1960; their tests empty, both cases indicative of rapid entomb­ RADWANSKA 1987, 2003b). This evidences generally a ment of live specimens (see ASLIN 1968; SMITH 1984; phyletic radiation and adaptive plasticity of comatulids RADWANSKA 1999, 2004b). If so, then the alpheid bur­ to colonise effectively shallow-marine post-Triassic rows were acting as mortal traps for still alive echinoids habitats left by other crinoids that sincc the Late which could not escape from the burrows. Cretaceous till Early Palaeogene have migrated to The alpheid burrows of the Upper Oolite Member at greater depths (see D.L. MEYER & MACURDA 1977, Malogoszcz have long acted as traps, both during the D.L. MEYER & OJI 1993, MESSING 1997). soft-bottom stage, and later, at hard bottom phase, when they became lithified and erosionally truncated. Even then, when the hardground was overgrown by massively­ Acknowledgements stalked crinoids Apiocrinites, the detrital material was still swept into the alpheid burrows. Comatulid crinoids Dr. Marcin MACHALSKI (Institute of Palaeobiology, Polish studied are thought to have lived over the oolitic shoals Academy of Sciences, Warsaw) and Adrian KlN, M.Sc. successfully, to form local 'feather-star gardens'. Together (Department of Palaeontology, University of Warsaw) are with stalked crinoids, echinoids, ophiuroids, and aster­ thanked for kind donation of some specimens to this study. oids they largely contributed to the formation of an Dr. Hans HESS (Binningen, Switzerland) critically 'echinoderm graveyard' (in German Echinodermen­ reviewed the manuscript and suggested some alterations which lagerstiitte, see c.A. MEYER 1984). improved content of the text. Dr. John W.M. JAGT (Natuur­ Noteworthy is that in the whole section exposed at historisch Museum Maastricht, The Netherlands) not only Malogoszcz Quarry the comatulids appear only at some reviewed an early draft, but also trimmed the language conside­ intervals ( see Text-fig. 1) which bear COlmnon burrows, rably. and formed under extremely shallow-water conditions: The Project has been supported financially by the Polish oolitic shoals burrowed by alpheid shrimps (see State Committee for Scientific Research (KBN), Grant No.3 RADWANSKA & RADWANSKI 2004b) and oyster shellbeds of P04D 01925). supposedly storm-bar origin (see MACHALSKI 1996, RADWANSKA & RADWANSKI 2003). In both cases, an effec­ tive agcnt causing death and burial of comatnlids was REFERENCES stonn agitation. To this velY agent some other Jurassic echinoderm lags/graveyards (Echinodermenlagerstiitten) ASUN, C.J. 1968. 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ManusC/ipt submitted: lOth October 2004 Revised version accepted: 20th March 2005 l~LI· ACTA GEOLOGICA POLONICA, VOL. 55 U. RADWANSKA, J'L. I

PLATE 1

1- Comatulina cf. beliremiellxi (DE LORIOL, Ul89); la - Aboral vicw of centrodorsal; Ib - Lateral view of centrodorsal; both x 7; CKm/004; Ic - Radial plate, articular facet; CKm/005; x 10 2-4 - Comatulina peroni (DE L01<.10L, 1889); 2 - Lateral view of calyx; CKm/006; 3a - Oblique-lateral view of calyx, 3b - Aboral view of centro dorsal; CKm/007; 4a - Lateral view of calyx, to show basal plate; 4b - Lateral view of calyx, to show radial plate; CKm/008; all x 7 5 - Palaeocomaster sp., CKm/01 0; 5a - Adoral view of centro dorsal, 5b - Lateral view of centrodorsal; both x 7 ACTA GEOLOGICA POLONICA, VOL. 55 U. RADWANSKA, PL. 1 ACfA GEOLOGICA POLONICA, VOL. 55 U. RADWANSKA, PL. 2

PLATE 2

Comatulina malogostiana sp.nov.

1-2 - Paratypes: la - Lateral view of eentrodorsal. Ib - Aboral view of centrodorsal; CKm/003; 2a - Lateral view of calyx, to show radial plate; 2b - Lateral view of calyx, to show basal plate; 2c - Aboral view of centro dorsal; 2d - Adoral view of calyx; CKm/002; all x 7 3 - Holotype: 3a Adoral view of calyx, 3b - Lateral view of calyx, to show basal plate; 3c - Lateral view calyx, to show radial plate; 3d - Aboral view of centro dorsal; CKm/OOI; all x 7 ACTA GEOLOGICA POLONICA, VOL 55 U. RADWANSKA, PL 2 ACTA GEOLOGICA POLONICA, VOL. 55 U. RADWANSKA, PL. 3

PLATE 3

1 - Comatulina Sp., CKm/009; Ia - Lateral view of centrodorsal, Ib - Adoral view of cen­ trodorsal; all x 7 2 -Palaeocomaster karsensis sp.nov., CKm/O 11 , holotype: 2a - Lateral view of calyx, to show not observable basal plate; 2b Lateral view of calyx, to show radial plate; 2c - Oblique-lateral view of calyx, to show articular facet of radial plate; 2d - Adoral view of calyx; 2e - Aboral view of centrodorsal; all x 7 ACTA GEOLOGICA POLONICA, VOL 55 U. RADWANSKA, PL 3 AcrA GEOLOGICA POLONICA, VOL. 55 U. RADWANSKA, PL. 4

PLATE 4

Solanocrinites sanctacrucensis sp.nov.

1 - Holotype, CKm/012: Ia Articular facet of radial plate; Ib - Lateral view of calyx, to show basal plate; Ic Adoral view of calyx; Id - Lateral view of calyx, to show the overhanging radial ring 2 - Paratype, CKm/013: 2a - Lateral view of calyx, to show radial plate; 2b Lateral view of calyx, to show basal plate ACTA GEOLOGICA POLONICA, VOL. 55 U. RADWANSKA, PL. 4