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Contr. Tert. Quatern. Geol. 28(1) 35-43 3 figs, 1 pi. Leiden, August 1991

Early Miocene luidiid asteroids

(Echinodermata, Asteroidea)

from Winterswijk-Miste (The Netherlands)

John W.M. Jagt

Venlo, The Netherlands

Jagt, John W.M. Early Miocene luidiid asteroids (Echinodermata, Asteroidea) from Winterswijk-Miste (The Netherlands). — Contr

Tert. Quatern. Geol., 28(1): 35-43, 3 figs, 1 pi. Leiden, August 1991.

Dissociated ossicles (inferomarginals, ambulacrals and adambulacrals) of luidiid asteroids (Luidiidae) from early Miocene Hemmoorian

(Oxlundian) strata at Winterswijk-Miste (province ofGelderland, The Netherlands) are illustrated and briefly described. The material is

with ciliaris for the time left in nomenclature. It a new compared(Philippi,contemporary 1837), but, being, open may represent

forma within the Ciliaris ofthe Luidia Forbes, 1839. The occurrence to subspecies or Group (sensu Döderlein, 1920) present appears

be stratigraphically older than records of Miocene luidiids from central Poland.

- - Key words Echinodermata, Asteroidea, Luidiidae, Miocene, Hemmoorian (Oxlundian), The Netherlands.

J.W.M. Jagt, Tweede Maasveldstraat 47, 5921 JN Venlo, The Netherlands.

Contents

Introduction 35 p.

36 Systematic description p.

Future studies p. 39

Acknowledgements p. 40

References 40 p.

Introduction

During processing of a rather small sediment sam-

taken from ple a temporary outcrop at Winterswijk-

Miste (Fig. 1) in 1988, dissociated ossicles of the asteroid genus Luidia were recognised amongst the generally diverse remains. Asteroids

found in this material isolated ossicles are as only

and comprise at least nine . Astropectinids are the commonest asteroids, and include three or four of which be still undes- species, some appear to cribed. Figure 2 illustrates morphological differen-

Some of the ces between astropectinids and luidiids.

well asteroid ossicles available for study are fairly

but preserved, most are corroded and/or abraded, location of Fig. 1. Locality map showing temporary exposure and often fragmentary. Echinoids comprise cida- (black dot) at Winterswijk-Miste. Abbreviations: W - rids, and clypeasteroids spatangoids. Ophiurid Winterswijk, M - Miste, B - Bredevoort (modified after ophiuroids are apparently extremely rare as only Janssen, 1984). 36

lateral arm plates and vertebrae have so far been

recorded.

From coeval or slightly younger strata (Hem-

moorian-Reinbekian) in NW Europe luidiids were

unknown up to now. R. Janssen (1972) described

from the northwest German Middle and Late

Miocene several species of and

Goniasteridae, but no Luidiidae. Polish occurrences

of luidiids of middle Miocene (Badenian) age were

recorded by Kaczmarska (1987). The Polish mate-

rial (referred by Kaczmarska to two modern taxa,

but probably representing only one species) appears

to be conspecific with the Dutch material, which is

here compared with (Philippi, 1837).

Over the few well past years, contemporary as as

fossil luidiidasteroids have received ample attention

in various papers, especially those by Blake (1972,

1973, 1982) and Blake & Guensburg (1989).

SYSTEMATIC DESCRIPTION

Classification follows Blake (1987, 1989) and Gale

(1987a).

Class Asteroidea de Blainville, 1830

Superorder Valvatacea Perrier, 1884

Order Perrier, 1884

Suborder Diplozonina Spencer & Wright in

Moore, 1966

Family Luidiidae Sladen, 1889

Genus Luidia Forbes, 1839

Type species — Luidia fragilissima Forbes, 1839 (=

Asterias ciliaris Fig. 2a-c. Cross sections of arms of luidiid (A, C) and astropec- Philippi, 1837).

tinid (B) asteroids to show position ofindividual ossi-

— Ciliaris Hemicnemis cles (modified after Clark, 1982, Döderlein, 1920, Species group Group (subgenus Miiller and Fell, 1963). & Troschel, 1840) sensu Doderlein, 1920. 37

Luidia aff. illustrated herein Nationaal sp. ciliaris (Philippi, 1837) mens (PI. 1),

PI. 1 Natuurhistorisch Museum Collections, Leiden,

Compare: RGM registration numbers 383 547-383 554.

Description — Blake (1973) presented detailed 1837 Asterias ciliaris 194. Philippi, p. of the ossicles of various luidiid 1920 Luidia ciliaris Doderlein, 287, 288, 18, descriptions species Philippi pp. pi. used and indicated the fig. 8; pi. 19, fig. 17; pi. 20, fig. 34. (for terminology see Fig. 3)

ciliaris — 39a. 1927 Luidia (Philippi) Mortensen, p. 70, fig. following features to be characteristic of the Ciliaris

1953 Luidia ciliaris — 380. (Philippi) Clark, pp. 379, Group sensu Doderlein (1920): — 1973 Luidia ciliaris 1837 Blake, 40, 10, (Philippi), p. pi. figs - inferomarginals: outline crescentic, without a dis- 1-26. boss mod- tinct inner face step; superambulacral 1982 Luidia ciliaris — 170, 3k (Philippi) Clark, p. fig. (with distal articulation synonymy). erately prominent; ridge parallel

1987 Luidia ciliaris (Philippi, 1837)— Kaczmarska, 139, pi. outline wide p. to curvature, rectangular, very

5, figs 6-8. with articulation and long, two marginal processes;

1987 Luidia alternata — 139 (Say, 1825) Kaczmarska, p. (? pars), proximal articulation ridge shortand wide. pi. 5, figs 10, 11 (? non 9, 12).

- ambulacrals: ambulacral body triangular, asym-

— Unt G Material Dissociated ossicles including inferomar- metrical; dentition weak, with medial gap;

ambulacrals and adambulacrals oral GG ginals {c. 40), ( c. 10) small; apophyse prominent; subsym-

(c. 10), J.W.M. Jagt Collection, and the speci- metrical; aboral ridge low and rounded.

neozelanica 1925; Ciliaris Fig. 3. Diagrammatic representations of various ossicles of a luidiid asteroid Luidia Mortensen, Group)

illustrating morphological terms used in the text (modified after Blake, 1973). A - left adambulacral, B - right adambulacral, C -

- face surface - inferomarginals, D - ambulacrals; 1 - spine bases, 2 - dml (distal face muscle 1), 3 psr1 (proximal 1), 4

- - - 6 - distal articulation 7 - articulation 8 adambulacral 9 superambulacral boss, 5 inner face step, ridge, proximal ridge, notch,

- - oral 10 - aboral 11 - ambulacral 12 - medial 13 14 Unt G Insertion des apophyse, ridge, body, gap, dentition, (“Grube zur

- 16 - face muscle unteren Quermuskels”), 15 GG (“Flügel zur Verbindung mit den Adambulacralia”), pm1 (proximal

depression 1). German terminology is after Müller (1953). 38

A comparison of Blake's (1973) detailed descrip- from southern Norway, the Skagerrak, Kieler

tion of ossicle morphology of Luidia ciliaris with the Bucht, Shetland Islands, Faeroe Channel, south to

Miste material makes it clear that this should be the Canaries and Azores (? Cape Verde Islands),

classed in the Ciliaris Group. It shows the following and the Mediterranean at depths between 1-400

typical features: metres (Doderlein, 1920; Mortensen, 1927[1977];

Dons, 1937; Blake, 1973; Rodriguez, 1980; Tor-

1980; Clark, 1982). Inferomarginals tonese, Discussion — The stratigraphically oldest luidiid to

- outline crescentic 1, 13, far Luidia inferomarginal (PI. Figs have been recorded so is hungarica, based

I 4 from , 16) on a fragmentary specimen Nagybatony (Hun-

- distinct inner face no step (PI. 1, Figs 13, 14, 16) gary) and described by Rakusz (1927). Spencer &

- boss prominent (PI. did not superambulacral moderately Wright (in Moore, 1966, p. U 43, fig. 42/3)

1, and it is this Fig. 14) question this generic assignment on

- distal ossicle articulation ridge parallel to curva- material that based the of the Hungarian they range and outline ture (PI. 1, Figs 10, 14) very angular Luidia Dr D.B. Blake genus (Miocene-Recent); (let-

with distinct notch 1, Figs 9, 10, 12) considers the marginal (PI. ter of March 28, 1989) tells me that he

- three moderate-sized spine bases (PL 1, Figs 9-14) with which specimen to be incorrectly identified,

- short and wide proximal ridge (PI. 1, Figs 9, 12, conclusion I Hess also concur. (1955, p. 68) briefly

15) discussed this record and stated literally, Luidia

dieser hungarica aus dem Miozan diirfte hingegen

that the Ambulacrals Gattung angehoren.' He remarked occur-

of rence a 'modern' species of luidiid was quite

- ambulacral body triangular and asymmetrical (PI. possible, since Mio-Pliocene Italian astropectinid 1, Figs 4, 7) species displayed modern features and were refer-

- dentition weak, with medial 1, 4, gap (PI. Figs 5) Another able to the genus Gray, 1840. - Unt G small (PI. 1, Fig. 4) fossil asteroid to have previously been referred to

- oral apophyse prominent (PI. 1, Figs 5, 6) the Luidiidae is the Liassic (early ) Asterias

- GG subsymmetrical (PI. 1, Figs 4, 7) murchisoni which Hess Williamson, 1836, (1955, p.

- aboral ridge low and rounded (PI. 1, Figs 5, 6) 68) referred with a query to Plumaster Wright, 1861

(? Tropidasteridae Wright, 1880). Adambulacrals The Miste specimens have been collected from

the Aalten Member, Miste Bed, Hiatella arctica - pml moderately wide (PI. 1, Fig. 3) Acme Zone and/or base of Astarte radiata Acme

- psrl relatively large (PL 1, Fig. 3) Zone den and of ( sensu van Bosch et al., 1975) are - spine bases weak (PI. 1, Figs 1, 2) Hemmoorian(Oxlundian) age (A.W. Janssen, 1984; - dml large (PI. 1, Figs 1, 2) also A.W. Janssen & King, 1988; see Rogl, 1990).

would the In contrast with Blake (1973), who concentrated on This correspond to proposed global early

descriptive ossicle morphology, Clark (1982) Miocene Burdigalian Stage (Cowie & Bassett, 1989;

described external features and tabulated the fol- Hinsch, 1990; Jenkins, 1990). The Polish luidiids

Kaczmarska lowing characteristics for Luidia ciliaris: described by (1987) are apparently

of consecutive lateral and of middle Miocene = Lan- Arm number 7 or 8; number younger (Badenian

10 in international paxillae corresponding to superomarginal ones ghian/Serravallian nomenclature;

from 15 to 18; paxillar armament consisting of central Haq & van Eysinga, 1987) age. Judging

illustrations the Polish material spinelets distinctly coarser than peripheral ones; Kaczmarska's (pi. 5)

less best of shape of superomarginal paxillae more or elon- seems referred to a single species, instead to

gate; alignment of inferomarginal plates lateral; two taxa.

consecutive The here described inferomarginal spines alternating on luidiid ossicles are provi-

Luidia plates; number of large erect inferomarginal spines sionally assigned to the contemporary species,

of ciliaris. asteroids 3 to 5; number of adambulacralspines 2; number Many post-Palaeozoic are very

3 when similar indeed from modern valves on actinal pedicellariae up to present; to or inseparable spe- oral furrow pedicellariae absent. cies at the generic level (see discussion in Blake,

Modern Luidia ciliaris lives in the NE and the level Atlantic, 1986), plausibly at specific as well, 39

Cainozoic FUTURE which seems especially true for species. STUDIES

Yet, in view of the fact that the present material is

form's of varia- As stated above, the asteroid fauna from Win- insufficient to determine this range

nine Astro- to leave it in nomenclature. at least tion, it is preferred open terswijk-Miste comprises species. four defi- Additional material from Winterswijk-Miste, and pectinids include three or species, two

to 1840, and preferably from as many coeval occurrences as pos- nitely assignable Astropecten Gray, Brasiliensis of Doderlein, sible, will certainly show their exact relationship possibly to the Group

reminiscent of aff. L. with L. ciliaris of which it well be a one Lophidiaster pyg- may precursor 1917, sp.

maeus 1913 1972, 3, distinct at the subspecific or forma level. It belongs Spencer, (Rasmussen, pi. figs

and another the Ciliaris of Doder- one without any doubt to Group 26, 27), strongly resembling repre-

sentatives of the Coulonia de Loriol, 1874 lein (1920) as pointed out above. genus (=

be the strati- Cuneaster Cuneaster hauteriviensis Hess, Most importantly, this appears to Hess, 1955; of Coulonia neocomiensis graphically oldest record of fossil luidiids for NW 1955 being a junior synonym

luidiid This is a of Europe. Blake (1973) described five fossil de Loriol, 1874; see Hess, 1970). genus

Pliocene that from the Cre- species from California (U.S.A.): a late or astropectinids ranges Early

referred to the mod- taceous to the Eocene Rasmussen, early Pleistocene specimen was Early (see 1972).

L. A terminal in the collection available for ern L. foliolata Grube, 1866; two new species, single ossicle referred Kaczmarska etchegoinensis Blake, 1973 and L. sanjoaquinensis Blake, study resembles the by

Pleistocene to 1973, of Pliocene and early age, respec- (1987, pi. 3, fig. 2a-b) Astropecten granulatus from and three additional a first described tively, were erected species, Rasmussen, 1972, species

of which Luidia of the remain- British Bartonian strata. one ( sp. A) Miocene, (Eocene) include left in Goniasterids a new ing of Pliocene age, were open nomenclature, possibly representa-

tive 1899 related but all assigned to the Alternata Group of Doderlein of Verrill, or a closely

also recorded that it Rasmussen see 1950) (1920). Blake (1973, p. 55) also pointed out genus. (1972,

Ciliaris three Cainozoic of this all of which might be desirable to treat the Group as a species genus,

in from the ossicle of more differ several details Miste distinct genus, once morphology specimens. with better known. Kutscher referred a dissoci- luidiid species were (1980, 1985) query

ated of Rasmussen's of this Both Luidia and Astropecten are active predators marginals to one species

Another to be that hunt during certain periods and then bury genus. goniasterid species appears which related to Teichaster 1913, themselves to digest prey (Clark, 1968; Lawrence, closely Spencer,

in with Crateraster 1987; Nichols, 1966). They are found on and Gale (1987b) synonymised Spencer,

rock and shell 1913. Yet another form resem- unconsolidated sediment, but also on displays a superficial

blance the Late Caletaster bottoms, in shallow, turbulent environments. to genus

Amongst the paxillosidans, both Luidia and Astro- (Breton, 1979) and to the Late Jurassic-Miocene

and Recent Sladen, 1889 Blake, pecten are offensive specialists (Blake, 1988, 1989, Paragonaster (see

and other A few aboral ossicles are 1990) that feed on molluscs, 1973, pi. 18, figs 14-25). that invertebrates; the remaining families referred to the like those of Stimpson, 1857, a genus North American Paxillosida being small particle feeders. Deposition includes Cretaceous (Blake, 1986)

Miocene New Zealand of the Miste Bed of the Aalten Member must have and Early representatives.

in favourable environment for The last is characterised a curious taken place a astro- goniasterid by which pectinids and luidiids, in view of the profuse mollus- ornament on the marginals, may, however,

and the be induced can assemblages (A.W. Janssen, 1984), gen- by poor preservation.

invertebrate It is unfortunate that the Cainozoic strata in erally varied echinoderm and other very

faunas. Contemporary Luidia ciliaris feeds primarily the North Sea Basin have not yet yielded any more

lesser or less asteroids. Finds of on other echinoderms and, to a extent, on completely preserved

molluscs (Brun, 1972). remarkably well-preserved specimens of goniaste-

Luidia rids and the Baluk & Rad- Blake (1989) noted that Astropecten and are astropectinids as ones

the successful of modern of wanski recorded from the Tortonian of amongst most genera (1968) early

asteroids and that their origin and adaptation to central Poland deserve the closest possible atten-

of shallow, turbulent environments might have been tion. In general, Polish occurrences Cainozoic

from the North Sea late, considering their comparatively recent (Eocene are similar to the ones

Basin in dissociated ossicles and Miocene, respectively) first known occurrence. comprising only (see e.g. 40

Baluk, 1970; Baluk & Radwanski, 1977; Gutowski, Blake, D.B., 1988. Paxillosidans are not primitive asteroids: A based functional considerations. In: R.D. hypothesis on 1984). Burke, P.V. Mladenov, P. Lambert & R.L. Parsley (eds). Future studies will hopefully comprise additional Echinoderm Biology. Proceedings of the sixth international material of in addition extensive discus- luidiids, to echinoderm conference, Victoria/23-28 August 1987. Rot-

terdam & Brookfield 4 sions of the species mentioned above. (A.A. Balkema): 309-314, figs. Blake, D.B., 1989. Asteroidea: Functional morphology, classi-

fication and phylogeny. In: M. Jangoux & J.M. Lawrence ACKNOWLEDGEMENTS (eds). Echinoderm Studies 3. Rotterdam & Brookfield (A.A.

I wish extend best thanks Dr B.D. Blake Balkema): 179-223, 3 figs, 3 tabs, pis 1-3. to my to Blake, D.B., 1990. Adaptive zones ofthe class Asteroidea (Echi- (University of Illinois, Urbana, U.S.A.) for items of nodermata). — Bulletin ofMarine Science, 46(3): 701-718, 3 literature, for critical reading of the manuscript and figs, 5 tabs. for his views the material described in expressing on Blake, D.B., & T.E. Guensburg, 1989. Illusioluidia teneryi n.

this and Dr U. Radwanska and (Asteroidea: Echinodermata) from the Pennsyl- paper to (Uniwersytet gen. sp. vanian of and its with the Texas, homeomorphy extant genus Warszawski, Instytut Geologii Podstawowej, Wars- Luidia Forbes. —Journal of Paleontology, 63(5): 662-668, 4 for of Polish zawa) supplying an offprint a paper. figs. The SEM taken Dr M. micrographs were by van Bosch, M. van den, M.C. Cadee & A.W. Janssen, 1975.

and subdivision of den Boogaard with the JEOL JSM-840A unit of Lithostratigraphical biostratigraphical Tertiary deposits (Oligocene-Pliocene) in the Winterswijk- Leiden University and Mrs C. Pepermans (Leiden of — Almelo region (eastern part the Netherlands). Scripta item of University library) provided an literature; Geologica, 29: 1-167, 37 figs, 10 tabs, 2 encl., 23 pis. — their assistance is gratefully acknowledged. Fellow Breton, G., 1979. Les asteries du Cretace de Normandie.

Bulletin de la Societe de Normandie et des Amis members of the Werkgroep voor Tertiaire en Kwar- geologique du Museum du Havre, 65(4)( 1978): 5-87, 32 figs, pis 1-8. taire Geologie (Cor Karnekamp, Nico Dekker, Wim Brun, E., 1972. Food and feeding habits of Luidia ciliaris Echi- Groeneveld, Jac Parren, Andre Jansen, Wim in 't nodermata: Asteroidea. —Journal of the Marine Biological

Hout, Jan Boes, Bert Kokmeijer and Freek Rheber- Association ofthe United Kingdom, 52: 225-236, 5 figs,pi. 1.

Notes asteroids in the British Museum Clark, A.M., 1953. on gen) are thanked for making available echinoderm — (Natural History). III. Luidia. Bulletin of the British material from Winterswijk-Miste. Dr P.A.M. Gae- Museum (Natural History), Zoology, 1: 379-396, 12 figs, pis mers is thanked for my attention (Leiden) drawing 39-41. Polish to a paper. Clark, A.M., 1968. and their relations (2nd edition).

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Rotterdam (Middle Miocene; Holy Cross Mountains, Central Poland). 141-149. (A.A. Balkema).

— Acta geologica polonica, 37(3-4): 131-144, 3 figs, pis 1-8.

Kutscher, M., 1980. Die Echinodermen des Oberoiigozan von

Sternberg. — Zeitschrift der geologischen Wissenschaften,

8(10): 221-239, 1 fig., 4 pis.

Kutscher, M., 1985. Die Echinodermen des MagdeburgerGriin-

— Berichte sandes (Mittel-Oligozan. Abhandlungenund zur November, revised version Naturkunde und Vorgeschichte (Magdeburg), 12(6): 3-14, 3 Manuscript received 28 accepted

23 1991 pis. January 42

PLATE 1

Dissociated ossicles of Luidia aff. ciliaris from Miocene sp. (Philippi, 1837) temporary exposure at Winterswijk-Miste. Early (Hemmoorian,

Pteropod Zone 18 of Janssen & King, 1988), Breda Formation, Aalten Member, Miste Bed (van den Bosch et al., 1975). All specimens illustrated have been deposited in the collections of the Nationaal Natuurhistorisch Museum at Leiden (formerly: Rijksmuseum van

Geologie en Mineralogie), RGM registration numbers (leg. J.W.M. Jagt):

Figs 1, 2. right adambulacral, oblique distal and oblique lateral views, respectively, RGM 383 547, x 22 and x 26, respectively.

Fig. 3. left adambulacral, oblique proximal view, RGM 383 548, x 21.

4-6. and lateral RGM 383 18. Figs right ambulacral, oral, two views, respectively, 549, x

Figs 7, 8. left ambulacral, aboral and oblique lateral view, respectively, RGM 383 550, x 26.

Figs 9-11. right inferomarginal, oral, oblique distal view and detail of spine base, respectively, RGM 383 551, x 23 and c. 50,

respectively.

Figs 12, 13. right inferomarginal, oral and proximal view, respectively, RGM 383 552, x 18.

14. left distal RGM 383 16. Fig. inferomarginal, view, 553, x

Figs 15, 16. right inferomarginal, oral and proximal view, respectively, RGM 383 554, x 16. 43\1

PLATE 1