October 2003 CainozoicResearch, 2(l-2)(2002), pp. 143-155,

First record of gorgonocephalid ophiuroids (Echinodermata)

from the Middle Miocene of the Central Paratethys

Andreas Kroh

Institut für Geologie undPaläontologie, Karl-Franzens-Universität Graz, Heinrichstraße 26, A-8010 Graz, Austria;

e-mail: [email protected]

Received 28 February 2003; revised version accepted 21 May 2003

The shallow-water (sublittoral) coarse sandy sediments ofthe Hartl Formation (Langhian, Middle Miocene) which outcrop north of Eisenstadt (Burgenland, Austria) have yielded abundant disarticulated ophiuroid material, mostly large streptospondylous vertebrae.

These are here assigned to the Gorgonocephalidae, on the basis of arm branching and the presence of an open ventral groove along the

vertebrae entire arm. Oral plates found associated with these are described as a new species, Astrodendrum?pilleri n. sp. The fossil

is the material the first record of this from record of gorgonocephalidophiuroids very poor; present represents unquestionable group

the Central Paratethys, and at that, the stratigraphically oldest. To date, only few ophiuroids have been recorded from Neogenestrata

ofthe Paratethys; these are listed in an Appendix.

KEY WORDS: Echinodermata, Ophiuroidea, Gorgonocephalidae, Miocene, Austria, CentralParatethys, new species.

Introduction side near Dunedin (IGNS, locality number GS 1890) (pers.

the basis of comm. J. Simes, February 2003). On photo-

The Gorgonocephalidae have a very poor fossil record, graphs I received of these specimens (three ‘non-

which is rather surprising since extant representatives are branching’ vertebrae), I am unable to either confirm or

found in a large variety of habitats, ranging from the sub- dismiss Fell’s identification.The third record pertains to a

polar to tropical climate zones (Doderlein, 1927; Morten- small collection of dissociated gorgonocephalid vertebrae

Hendler sen, 1933; Baker, 1980; et al, 1995). In addition, from Pliocene strata penetrated in borehole IJsselmuiden-1

be in shallow-water habitats Kroh in they may common (Baker, (eastern Netherlands; see & Jagt, press).

1980; Hendler et al., 1995) and generally possess numer- Apart from these three records that explicitly refer to

ous large and relatively sturdy ossicles. gorgonocephalids, there are several reports of euryalid-type

The and of fossil vertebrae from of Cainozoic first, only, unquestionable report deposits age (ie.g., Vadasz,

gorgonocephalids thus far is that by Pomel (1885-1887), 1915 - Miocene, Hungary; Valette, 1928 - Burdigalian

who described dissociated vertebrae from the ‘Sahelien’(= (Miocene), France, misidentifiedas brachials of the coma-

‘ Upper Miocene-Pliocene; see Durand-Delga, 1962) of tulidcrinoid Antedon’ rhodanicus; and Donovan & Paul,

Algeria, underthe name ofAstrophyton saheliensis Pomel, 1998 - Upper Pliocene, Jamaica). However, it is not clear

of 1885. One his illustrations (Pomel, 1885-1887,pi. D II, whether any ofthese records do indeed represent gorgono-

shows vertebra. fig. 7) clearly a branching Subsequent cephalids or are merely similar vertebrae ofother ophiuroid

authors have oftenoverlooked this record; a rare exception groups with streptospondylous vertebral type (e.g.,

is Sieverts-Doreck’s (1954) compilation. Anotherreport of Hemieuryalidae, Euryalidae, Ophiobyrsinae, Asterosche-

fossil gorgonocephalids is found in Spencer & Wright matidae and others).

(1966, p. U91), who noted, ‘An undescribed genus occurs The fossil record of euryalids, another extant family

in the of New Zealand with Oligocene (Fell, in lift.).’ Assess- branching, streptospondylous vertebral type, is simi-

ment of this statement is in the absence of illus- of liasica difficult, larly poor. An early report Euryale? Quenstedt,

trations and/or description of the material both in Spencer 1876 has subsequently been shown to consist of resting

far in traces stars & Wright (1966) and, as as I know, subsequent of indeterminatebrittle with unbranching arms,

literature sources. The material on which this record is with the impression of branched arms having been pro-

based is currently housed at the Institute ofGeological and duced by multiple traces of moving arm tips (Seilacher,

Nuclear Sciences (Lower Hutt, New Zealand), labelled 1953). Records of Trichasterl ornatus (Rasmussen, 1950)

vertebral ossicle - ‘Ophiurid Gorgonocephalus [sic]’; it is and Trichasterl sp. from the Campanian and Maastrichtian

from the Caversham Sandstone Bum- of the Netherlands and (Lower Miocene) at SE NE Belgium (Jagt, 2000, pp. 8, - 144-

of than 9; pi. 1, figs 5-10) and from the Maastrichtian Riigen more 150 years ago. This may be an artifact, however,

and & since earlier workers considered less (Germany) Mon (Denmark; see Jagt Kutscher, 1998; preferentially more or

Kutscher 2000, 51-53; and disarticu- & Jagt, pp. pi. 32, figs 1-7) com- complete well-preserved specimens. In fact, prise dissociated streptospondylous vertebraewith an open lated skeletal material was commonly ignored. Recent ventral attributedwith examination of bulk from in groove, originally a query to Astero- samples Neogene strata Aus-

Rasmussen tria has revealed that ossicles in fact nyx by (1950, 1972). Although superficially ophiuroid are a com- similar to vertebrae of the genus Trichaster, the attribution mon element in echinoderm palaeocommunities (Kroh, of these fossil ossicles to the Euryalidae and in particular research in progress). The Appendix lists all post-Eocene to this is all extant the from the most taxa in genus questionable. In euryalids, ophiuroids Paratethys; are urgent ventral furrow is closed by a ‘bridge-like’ structure in distal need of revision, having been based mostly on single

with little material for arm portions (Mortensen, 1933, pp. 3, 4). In many Recent specimens only or extant comparison

hand. taxa only a few proximal vertebrae have open grooves, e.g. at in 1879 the first and The is the third in series of studies of Asteroceras pergamenumLyman, only present paper a second vertebrae. In addition, neither Jagt (2000) nor Neogene echinoderms. Earlier papers were devoted to

echinoderm faunas from the Sands’ Mio- Kutscher & Jagt (2000) recorded any branching vertebrae, ‘Retz (Lower while extant Trichaster have arms branching up to ten cene), with emphasis on the palaeoecology of echinoid- times. bearing levels and on more global aspects, such as echinoid

The post-Eocene ophiuroid record in the Paratethys migration (Kroh & Harzhauser, 1999), and from the Lower

to be the fact that the first Badenian in the Molasse Zone appears extremely poor, despite (Langhian) as represented papers on echinoderm fossils from this area were published and the northernVienna Basin (Kroh, 2003).

Figure 1. Geographic and stratigraphic position of localities mentioned in the text. A: distributionof Lower Badenian (Langhian) sediments in Austria (slightly modified from Mandic etal., 2002 [courtesy of O. Mandic and F. Rdgl]; B: location of sections stud-

ied, 1 - Hartl-Lucke; 2 - Johannesgrotte (modified from Kroh et al., in press); C: Langhianpalaeogeographyafter Rdgl (1998).

vertebrae found the but Study area cephalid are throughout section, are

particularly abundant in the basal part. For detailed sec-

section ‘Hartl-Lucke’ microfacies and sedimen- Material studied comes from the type tions, lithological descriptions,

of the Hard Formation, north of Eisenstadt in Burgen- tological data, as well as a discussion of regional geology

50 reference is made to Kroh al. land/Austria (OK map, sheet 77) (Figure 1). Gorgono- et (in press). - 145 -

The the bulk study area belongs to Eisenstadt-Sopron Basin, Both samples and picked ophiuroid material are a small, strongly asymmetrical subbasin of the Vienna deposited at the Geologisch-Palaontologische Abteilung of

Basin complex, which is part of the Central Paratethys, the Naturhistorisches Museum Wien (NHMW); the thin

less displaying a more or trigonal size ofabout 20 x 20 km sections at the Institute fur Geologic und Palaontologie of in width(Filler & Vavra, 1991). In the north, it is bounded the Universitat Graz. by the NE-SW trending Leitha Mountains and the associ- A number of extant gorgonocephalid species housed in ated SE-dipping Eisenstadt fault (Fodor, 1992), in the east the collections of the United States National Museum

the and the by N-S trending Rust faults, Rust-Fertorakos (USNM) were examined for comparison with fossil speci-

which the basin from the Danube Basin. The thanks to the of viz. Astro- hills, separate mens, courtesy Cynthia Aheam, , southern margin ofthe Eisenstadt-Sopron is definedby the caneum spinosum (USNM E34453), Astrocladus dofleini

of the Rosalia which Astroboa nuda crystalline ridge Mountains, separates (USNM 15767), (USNM E1470), Astro- it from the Styrian Basin complex. dendrum sagaminum (USNM 25665), Astroglymma

Strata ofthe Haiti Formationhave recently been dated sculptum (USNM E1476), Astrochalcis micropus (USNM on foraminiferal evidence as Early Badenian (Late E1463), and Astrophyton muricatum (USNM E44551). A

Middle Kroh al. small of the Langhian, Miocene) by et (in press). piece middle portion ofan arm was removed

from each specimen, the soft tissue then dissolved using

sodium hypochlorite (30% solution). After a period in

clean the ossicles air dried and examinedunder Materialand methods water, were

a light microscope. Specimens used for SEM analysis were

Field work was carried out in the Spring/Summer of 2001 cleanedwith hydrogen peroxide and an ultrasonic bath, and

in of the both unconsolidated and the Spring 2002. In field, then mounted on stubs and gold coated. and consolidated sediments were sampled; ofthe former Higher-order systematics follow Smith et al. (1995),

250 g samples were first driedand then disintegrated using terminology of oral plates is after Murakami (1963) and

the were washed H202. Subsequently, samples using stan- that ofvertebrae follows Irimura & Fujita (2003). dard sieve-sets (0.063 mm, 0.125 mm, 0.25 mm, 0.5 mm,

1 mm, 2 mm, 4 mm, and 8 mm). Additional bulk samples

taken the of about2.5 kg each were to investigate macro- Systematic description fauna. These were washed on a 2 mm sieve and residues were handpicked under a stereo microscope. For quantita- Class Ophiuroidea Gray, 1840 tive thin sections ofthe consolidated Subclass 1840 component analysis, Ophiuridea Gray, sediments were prepared. Unconsolidatedsediments were Order Euryalina Lamarck, 1816 embedded in resin and also thin-sectioned to produce Family Gorgonocephalidae Ljungman, 1867 comparable results. Both the washed samples and the thin Genus Astrodendrum Doderlein, 1911

sections were subjected to quantitative analysis in order to assess the distributionof biogenic components within the Astrodendrum?pilleri n. sp.

sections. Detailed results of sieve, carbonate and thin

section analysis will be presented elsewhere (Kroh et al., in Figures 2, 3 press). To split the gorgonocephalid vertebrae into size

— oral is classes, a set of sieves from 1.0 to 5.0 mm size in 0.5 mm Types Holotype, an plate, NHMW

used. These sieves drill- another oral is gradation were were prepared by 2003z0010/0002;paratype, plate, NHMW

ing closely spaced holes into polystyrol boxes, with the size 2003z0010/0003.

of the holes controlled with an electronic caliper, not

than +/- with standard Locus - varying more 0.05 mm. Experiments typicus Eisenstadt, Burgenland (Austria).

analysis sieve sets had earlier shown to yield unreliable

results due to the rectangular shape ofthe holes. The round Stratum typicum - Hard Formation, Lower Badenian

holes of the sieves ensured splitting in equal size classes. (Langhian), Middle Miocene.

Sample Fraction (0 in mm) > 5.0 > 4.5 > 4.0 > 3.5 > 3.0 > 2.5 > 2.0 > 1,5 > 1.0 Sum

"normal" vertebrae 3 4 24 34 123 205 193 27 3 616

branching vertebrae (sym.) 1 2 4 5 2 2 4 - - 20

■C vertebrae - - 2 7 16 17 19 2 - 63 cu branching (asym.) I ratio (nV:bV) 1:3 1:2 1:4 1:2.8 1:6,9 1:10.8 1:8.4 1:13.5 - -

overall ratio 1:3.3 1:7.9 1:13.5 - 1:7.4

Table 1. Size distributionof dissociated gorgonocephalidvertebrae from sample Hartl-Lucke. - 146-

2. Astrodendrum? Eisenstadt basal Haiti Formation Figure pilleri n. sp.; Hartl-Lucke, (Burgenland, Austria), (Lower Badenian,

Langhian, Middle Miocene). Scale bars equal 1 mm.

1- oral plates, in abradial and adradial views, respectively; 1 - NHMW 2003z0010/0001;

2- NHMW 2003z0010/0002, holotype;

3 - NHMW 2003z0010/0003,paratype; 4 - NHMW 2003z0010/0004.

5 first vertebra, in proximal and distal views, respectively (NHMW 2003z0010/0005).

6 bifurcating vertebra; in proximal, distal and lateral (distal to left), aboral and oral views, respectively (NHMW 2003zOO 10/0006), - 147-

Figure 3. Astrodendrum?pilleri n. sp.; Hartl-Lucke, Eisenstadt (Burgenland, Austria), basal Hartl Formation (Lower Badenian,

Langhian, Middle Miocene). Scale bars equal 1 mm.

1 medial vertebra, in proximal, distal, lateral(distal to left), aboraland oral views, respectively (NHMW 2003z0010/0007);

2 medial vertebra, in proximal, distal, lateral (distal to left), aboral and oral views, respectively (NHMW 2003z0010/0008);

3 first vertebra, in proximal, distal, lateral(distal to left), aboral and oral views, respectively (NHMW 2003z0010/0009);

4 symmetrically bifurcating proximal vertebra, in proximal and distal views, respectively (NHMW 2003z0010/0010);

5 asymmetrically bifurcating proximal vertebra, in proximal and distal views, respectively (NHMW 2003z0010/0011);

6 vertebra fromjust proximal of bifurcation, in proximal and distal views, respectively (NHMW 2003z0010/00012);

7 proximal vertebra, in proximal and distal views, respectively (NHMW 2003zOO 10/00013). - 148-

Figure 4. Proximal and medialvertebrae of extant gorgonocephalids,all in proximal, distal, lateral (distal to left), aboral and oral views,

respectively. Scale bars equal 1 mm.

1 Astrodendrum sagaminum(Doderlein, 1902),USNM 25665, from medial portion of arm;

2 A. sagaminum, USNM 25665, from proximal portion of arm;

USNM from 3 Astroglymma sculptum (Doderlein, 1896), E1476, medial portion of arm;

4 Astrocladus dofleini Doderlein, 1911, USNM 15767, from proximal portion of arm;

5 USNM Astrophyton muricalum (Lamarck, 1816), E44551, from medialportion of arm;

6 Astroboa nuda (Lyman, 1874),USNM E1470, from medial portion ofarm;

7 Astrochalcis micropus Mortensen, 1912, USNM E1463, from medial portion ofarm. - 149-

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~ 1Euryale £ 14(16 mm) ’ 49CD 5-6S SS0-2908 ' Co 3 9c cb s S 9 fo o £ 30-35 36-714 Sthenocephalus§ ft sym3 7-9 50 8-33 16-24(10-34) 30-35 7 | N f Trichaster1 nchaster 3-4£ y 75-100 37 34-661 9t-n14-16 75-100 106-1468 i 1 I | H£_Z” 1 Sources:Lyman, 1865,1865, 1874, 1875,1875, 1877, 1878;1878: Studer, 1881, 1884; DPderlein,Ddderlein, 1902, 1911, 1927, 1930;Boone, 1933;Mortensen, 1933;Spencer & Wright, 1966; Baker, 19801980

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Table Arm in suborder 2. branching the Euryalina, compiled from literature sources.

Diagnosis - A gorgonocephalid with grossly rhomboidoral Taxonomicprocedure Most fossil ophiuroid material is

plate with a concave dorsal margin and a large, undivided, preserved in the form of dissociated ossicles. Although

adradial muscular like vertically elongated area, shaped a extant species usually are defined upon disc plating, oral withbent handle. spoon frame structure and presence/absence of oral and dental

papillae, provisional generic attributions of fossil brit-

Derivatio nominis- Named after friend and tlestars my colleague are possible (e.g., Jagt, 2000). In fossil material, head Werner E. Filler, of the Institut fur Geologic und lateral arm plates have been shown to produce the most

Palaontologie at the Karl-Franzens-University Graz. reliable results (Hess, 1962a, b, 1963, 1965, 1966, 1975a,

In various also b). papers, Hess demonstratedthat in most

Material studied - Ten oral plates (NHMW cases dissociated vertebrae could not be used, since similar

and ~ vertebrae in different 2003z0010/0001-4, 14-15) 1,000 (NHMW types occur genera and families. Moreover, Table 2003z0010/0005-13, 16-18; see 1), all these ossicles exhibit a wide of originating range morphological varia- from bulk taken from the basal HartlFormation samples at tion along the arm. In gorgonocephalids, however, lateral

the Hartl-Lucke Eisenstadt rather small section, (Burgenland, Austria). arm plates are and reduced; so far none have

Although this cannot be demonstratedbeyond doubt, oral been recognised in the bulk samples available. This is

plates and vertebrae are here considered to be conspecific. probably related to the fact that most ossicles smaller than

2 mm in diameter are obscured by epitaxial cement, often - 150-

beyond recognition. 1963, pi. 3) is absent here. The abradial muscular area is

In extant material borrowed from the gorgonocephalid barely visible, but appears to be small; the adradial area is

United States National Museum the variation in (USNM), undivided, large, vertically elongated and is shaped like a vertebral structure along the arm turns out to be less wide spoon with bent handle, the dorsal part being distinctly than in other ophiuroid groups. In addition, proximal and enlarged. Both ab- and adradial articulation areas are large medial vertebrae in the studied reveal marked and well The specimens developed. arrangement of depressions for differences in size of oral fossae and ornament the first and second oral tentacles is to that shape, very similar

(Figure 4). For the new species, an oral plate is here desig- seen in oral plates ofAstrodendrum sagaminum. nated holotype, since of this type of ossicle there is only The oral plates consist of fused first and second ambu- one (morpho)type, whereas vertebral morphology changes lacrals (Hendler, 1978; Stewart, 2000), rather than of first the is along arm. Besides, there considerable variation in adambulacral and second ambulacral as stated by Mura- the of these in the shape plates amongst genera family kami (1963).

Gorgonocephalidae (Murakami, 1963), which allows a provisional generic attribution of the fossils to be made.

Matsumoto (1917) noted that oral plates are of consider- Discussion able taxonomic importance in extant ophiuroids.

The vertebrae studied clearly are assignable to the Gor-

gonocephalidae, as this is the only family to include forms

with Description branching arms which have vertebrae with open

ventral furrow both in proximal and distal arm positions.

’ ‘Normal vertebrae — Vertebrae range in size from less The family Euryalidae can definitely be excluded since than 1 5.5 in and show characteristic there the to c. mm diameter, a ventral furrow is closed by a ‘bridge-like’ struc-

hourglass-shaped streptospondylous articulation. In lateral ture in all but the proximalmost vertebrae (Mortensen, view, most are relatively narrow; the lateral furrow between 1933, pp. 3, 4). Additional evidence comes from the oral the and distal insertion is which proximal areas rather narrow and plates, are very similar to those of extant gorgono-

traces of ornament could be found in ossicle. The no any cephalids (compare Murakami, 1963, pis 3, 4).

aboral groove is V-shaped, the oral groove U- to V-shaped. Generic placement ofthe present material is difficult. In

Where preserved, oral fossae are large and not well differ- shape, the oral plates compare well with those of extant

entiated from the ossicles similar Astrodendrum remaining surface; a sagaminum (compare Murakami, 1963, pi.

condition is found in vertebrae of extant 3, 35, in the of Astrodendrum, figs 36), differing only presence an undi- while those of other genera exhibit well-differentiatedoral vided adradial muscular area; this is divided in A. sagami- fossae Proximal (see Figures 4.3-4.7). vertebrae (ie.g. num. In overall shape, the vertebrae are similar in all

3.4 and to in examinationof of Figures 2.5, 3.7) are trapezoidal rectangular gorgonocephalids. Nevertheless, a range proximal/distal view; medial ones are more rounded in extant species in the USNM collections indicates that there

outline Similar in the distinct differences (e.g., Figures 3.1, 3.2). changes shape are in ornament, shape of the aboral

of vertebrae the have been observed in size and of the oral fossae well lateral along arm extant groove, shape as as material (see e.g., Figures 4.1, 4.2). Vertebrae just distal of profile. In view ofthe fact that larger ossicles usually are

a branching vertebra are smaller horizontally and have a better preserved than small ones in the fossil material vertically elongated outline (see Figure 3.6). The first arm available, a comparison between them and extant speci- vertebrae (see Figures 2.5 and 3.3) could also be identified mens cannot be but restricted to proximal and medial

in the material studied; these occur in the size classes down ossicles. In outline and size and shape of the oral fossae, to diameters of> the fossil 2.5 mm. species compares well with Astrodendrum sa-

gaminum (Figure 4.2), and it is here tentatively referred to

Branching vertebrae — Branching vertebrae are slightly that genus. The lack of ornament would support this attri- wider than ‘normal’ their faces since A. ones, proximal being bution, sagaminum shows only very faint, easily

but the distal face has two inclined articu- obscured whereas in other closely similar, ornament, genera {e g., Astro- lation surfaces. In the present material there are vertebrae glymma, Astrophyton or Astroboa) ornament is rather that are symmetrical in distal view as well as asymmetrical coarse.

ones in distal view. In larger (i.e., from proximal arm Assuming all vertebraeto be conspecific, as suggested

portions) size classes there are mainly (near-)symmetrical by the presence ofjust one type of oral plate, it shouldalso

branching vertebrae, whereas the smaller size classes be possible to use the abundance of branching vertebrae to

predominantly yield the asymmetrical type (see Table 1). assign the material to an extant genus or group of genera.

Of the extant and thirty-four gorgonocephalid genera

Oral plates Oral plates of Astrodendrum?pilleri n. sp. subgenera, thirteen have simple, unbranched arms (Astero-

rhomboidal in outline, with dorsal are grossly a concave porpa s. str., Asteroporpa (Astromoana), Astrochele,

margin; however, the distinct notch in the dorsal margin Astrocrius, Astrogomphus, Astrohamma, Astrohelix, Astro-

Astrocladus present in many gorgonocephalids {e.g., stephanus, Astrothamnus, Astrothorax, Astrothrombus,

be coniferus or Astrothamnus bellator; compare Murakami, Astrotomaand Astrozona) and may thus excluded from - 151 -

consideration. Schizostella, a diminutive genus from the family Gorgonocephalidae. Close extant relatives of echi-

Caribbean has arms which show usually only a single arm noderms from the Paratethys are usually found in the

branching or, more rarely, two (Clark, 1952; Hendler et al, Atlantic Ocean (including the Caribbean Sea), the Mediter-

The of also additional and the 1995). type branching provides ranean western Indian Ocean, whereas taxa cur-

information. In the branch restricted the Indo-Pacific have some genera, arms mainly rently to region only rare

symmetrically (Astrocaneum, Astrodictyum, Astrosierra, fossil relatives in the Paratethys/Mediterranean region. The

Conocladus(syn. Astroconus) and Gorgonocephalus). In palaeoenvironmental interpretation of the Haiti Formation

a of related Astroboa Kroh al. indicates group genera to (e.g., Astrocladus, presented by et (in press) a nearshore

Astroglymna, Astrophyton, Astrodendrum), the proximal- position and relatively shallow depositional depth for the

the first bifurcations that have most (usually to third) are symmetri- strata yielded the new gorgonocephalid material. the distal cal, more ones are markedly asymmetrical. The This does not contradict with the proposed classification

number ofvertebrae before the first bifurcation occurring and supports the exclusion of multibranched, deep-water

and betweenbifurcation in the and medial such points proximal taxa as Astracme, Astrochlamys or Astroclon.

also shows arm portions some systematic variation and

couldbe useful in classification offossil material. A sum-

of these features and additional data size mary on range, Implications for gorgonocephalid phylogeny

depth range, geographical and climatic distributionof the

withinthe order be found in genera Euryalina may Table 2. Doderlein (1927, p. 12) presented an ad hoc phylogeny,

In order the number of vertebrae based to analyse branching on morphological features, of extant representatives

and their distribution the all fossil vertebrae ofthe with Conocladus the basal along arms, group, as form of nearly

from a bulk sample were sorted into nine size classes all extant this is (using gorgonocephalids. However, genus en-

sieves with round holes between 1.0 and 5.0 demicto South ranging mm, Australiancoastal waters (Baker, 1980, p. in of grades 0.5 mm). Within the size classes, ‘normal’ 73) and is probably highly derived. Doderlein considered

vertebrae, symmetrical and asymmetrical branching verte- the large number of vertebrae occurring before the first

brae The ratio between and bifurcation were distinguished. branching in the generaAstrocnida, Astroclon and Cono-

‘normal’ vertebrae the data matrix cladus be was calculated; resulting to a paedomorphic [sic] feature. The material be found in Table may 1. from the Middle Miocene of central Europe discussed in The distribution of symmetrical vs asymmetrical the present paper is the oldest unquestionable record of

vertebrae in the different size classes branching suggests a gorgonocephalids from the fossil record and shows multi-

branching type similar to that ofthe extant Astroboa group. ple branching and branching close to the disc to have

Symmetrically branching vertebraeare found mainly in the occurred already in the Miocene. There is no fossil evi-

size of whereas dence that coarse classes, or proximal portions arms, to suggest a low number ofvertebrae before the the number of asymmetrically branching vertebrae in- first branching can really be considered as the ancestral

in the smaller size creases ranges, or the distal portions of stage. Since these animals are passive suspension feeders

the The situation is found arm. same in extant gorgono- (Fricke, 1968; Macurda, 1976; Hendler, 1982; Wolfe,

cephalids relatedto Astroboa (e.g. Astrocladus, Astroclon, 1982), bifurcations in proximal arm portions are much ,

and Table better suited increase the available for Astrodendrum, Astroglymna others; compare 2, to area feeding than

column The overall ratio between and bifurcations 5). branching ‘nor- in the distal parts. Additionally, the mesh

mal’ vertebrae inthe material is 1:7.3. In the differ- width of the fan present feeding formed by the arms is smaller

ent size classes this value is distinctly different. In the four when there are numerous proximal bifurcations. Smaller

> size classes 3.5 mm, the ratio is 1:3.3. In size classes mesh widths enable the animals to catch more and smaller

between 2.0 and 3.0 mm, it rises to 1:7.9 and further to prey than with larger widths.

1:13.5 in the size class >1.5 mm. Although the sedimentof

the bulk samples has clearly been sorted by currents, this

< has affected mainly the small fraction 1 mm grain size, as Conclusions

indicated by a grain size analysis (Kroh, unpublished data).

Moreover, sorting is unlikely to have affected the ratios The of remains of multibranched recognition gorgono- between and ‘normal’ since these branching vertebrae, are cephalids in the fossil record is important since these similarin size and gross morphology. Ifthe pattern result- animals are highly specialised suspension feeders and thus from the the distri- ing analysis to some extent represents are useful in palaeoecological reconstructions. Extant

bution ofbifurcations along the arm and is not a sampling species are often found in reef habitats or elevations on

artefact, it also indicates that the material is mud and associated assignable to sea grass flats, with gorgonians, stony the Astroboa The group. genera Astroboa, Astrocladus, corals, fire corals and sponges (Hendler et al, 1995). They

Astrodendrum,. and in ascend Astroglymna Astrophyton particular are nocturnal, to an elevated perch at dusk and

show structure Table a closely comparable arm (compare extend their feeding arms in a parabolic array in moderately columns 2, 8-10). strong water currents. The stomach content ofAstrophyton This result also fits well the current depth range, geo- muricatum (Lamarck, 1816) has been shown to consist

graphical and climatic distribution ofthe generawithin the mainly of small copepods (Davis, 1966; Fricke, 1968; - 152-

Wolfe, Clark, A.H. 1952. Schizostella a new ofbrittle-star 1982). , genus (Gor- gonocephalidae).Proceedings of the United States National In addition, it is important to have a sound fossil record Museum 102, 451-454, pi. 40. ofanimal groups to allow calibration with the outcome of Davis, W.P, 1966. Observations on the biology of the ophiuroid phylogenetic analyses. The fossil record of ophiuroids, and Astrophyton muricatum. Bulletinofmarine Science 16,435- of in is rather Smith al. euryalids particular, poor. et 444.

(1995, p. 227) stated that, the absence of Euryaline taxa Dbderlein, L. 1896. Bericht fiber die von Herm Prof. Semon bei in the earlier of the Mesozoic is, however, part very puz- Amboina und Thursday Island gesammelten Ophiuroidea. In: since that should zling, our analysis suggests they have a Semon, R.W. (ed.). Zoologische Forschungsreisen in Aus- much longer fossil record’. In fact, the only fossil euryaline tralien und dem Malayischen Archipel. Jenaische Denck- taxa listed in Smith et al. (1995) are those described by schriften 8, 279-300. Doderlein, L. 1902. Japanische ZoologischerAnzei- Rasmussen (1950, 1972), plus the cursory mention of Euryaliden.

ger 25, 320-326. gorgonocephalids by Fell (in lift.; see Spencer & Wright, DQderlein, L. 1911. Uber und andere Euryale. Bei- the japanische 1966, p. U91). In present paper, at least for one group tragezur Naturgeschichte Ostasiens, Herausgegeben von Dr. of the Euryalina an unquestionable fossil record is docu- F. Doflein. Abhandlungen der Bayerischen Akademie der mented.Hopefully, future studies will focus in more detail Wissenschaften, mathematisch-physikalische Klasse, Sup- on euryaline vertebral ossicles and their confident assign- plement-Band2, 1-123, 7 pis. ment at the family level. Doderlein, L. 1927. Indopacifische Euryalae, Abhandlungen der

Bayerischen Akademie der Wissenschaften, mathematisch-

physikalische Klasse 31, 1-105, lOpIs.

Doderlein, L. 1930. Die der Deutschen Tiefsee- Acknowledgements Ophiuren Expedition. 2. Euryale. In: Chun, C. (ed.). Wissenschaftliche

Ergebnisse der Deutschen Tiefsee-Expedition auf dem This study was supported by the Austrian Science Founda- Dampfer Valdivia" 1898-1899, 22, 348-396, pis 14-16. tion „ (Project no. P-14366-Bio) to W.E. Filler (Karl- Jena (Gustav Fischer). Franzens-Universitat, Graz). Facilities at the Geologisch- Donovan, S.K. & Paul, C.R.C. 1998. Echinodermsof the Plio- Palaontologische Abteilung of the Naturhistorisches Mu- cene Bowden shell bed, southeast Jamaica. In: Donovan, S.K. Wien and collections the Institut fur seum access to at (ed.). The Pliocene Bowden shell bed, southeast Jamaica.

Palaontologie at the Universitat Wien are gratefully ac- Contributions to Tertiary and Quaternary Geology 35, 129- 146, 4 knowledged. For valuable discussion and support, I wish to pis, M. 1962. International, thank Rosalinde Esberger, Mathias Harzhauser, Ortwin Durand-Delga, Lexique Stratigraphique 4. Afrique; lb, Algerie, 132 Paris (Imprimerie Louis- Schultz and Helmut Sattmann (all Naturhistorisches Mu- pp., Jean), seum Wien), Gordon Hendler (Natural History Museum of Fodor, L. 1992. Tectonic setting of the Eisenstadt area. In: Sauer, Los Angeles County), John W.M. Jagt (Natuurhistorisch R., Seifert, P. & Wessely, G. (eds). Guidebook to excursions Museum Maastricht), Daphne E. Lee (University of in the Vienna Basin and the adjacent Alpine-Carpathian Werner E. Filler John Simes Otago), (Universitat Graz), thrustbelt in Austria. Mitteilungen der Osterreichischen of (Institute Geological and Nuclear Sciences, Lower Hurt, Geologischen Gesellschaft 85, 124-126.

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Appendix

Post-Eocene ophiuroid taxa recorded from the Paratethys, arranged in alphabetical order. Note: Unless species have been reassigned in subsequent papers, generic attribution is that of the original publication.

Amphiura? badensis Ktipper, 1954

(Kiipper, 1954, p. 163, pi. 15, figs 1,2; Lower Badenian[Langhian], Baden, eastern Austria)

Amphiura? gigantiformis Kiipper, 1954

(Kiipper, 1954, pp. 163, 164, pi. 14, figs 1-8; Badenian[Langhian-Serravallian], Mannersdorf, eastern Austria)

(Kristan-Tollmann, 1964, p. 77; Badenian[Langhian-Serravallian], Miillendorf, eastern Austria)

(Kristan-Tollmann, 1966, p. 132; same locality and stratigraphy)

Amphioplus? kuehni (Binder & Steininger, 1967)

(Binder & Steininger, 1967, pp. 21-23, pi. 1, figs 1,2, as Amphiural kÜhni; Karpatian [Upper Burdigalian], Teiritzberg,

northeast Austria)

(Meyer, 2002, pp. 325-331, pi. 1, figs 1, 2; same locality and stratigraphy)

Asteronyx sp.

(Kiipper, 1954, p. 164, pi. 15, fig. 3a-e; Badenian[Langhian-Serravallian], Romerberg near Baden, eastern Austria)

Ephippiellum symmetricum Lomnicki, 1899

(Lomnicki, 1899, pp. 224-227, fig. 3a-d; Badenian [Langhian-Serravallian], Wieliczka, Poland])

(Szajnocha, 1899, pp. 387-389; Badenian[Langhian-Serravallian], Wieliczka, Poland])

Tschokra Kertsch marl with denuda- (Andrusov, 1899, pp. 248,249; Miocene, Limestone, Peninsula; Miocene, Pecten

tus, Cap Tarchan])

(Lomnicki, 1902, pp. 155-157)

Ophiural parviformis Kiipper, 1954

(Kiipper, 1954, pp. 161,162, pi. 15, figs 4-14; Badenian[Langhian-Serravallian], Rauchstallbrunngraben near Baden,

eastern Austria)

Ophiura? vindobonensis Kiipper, 1954

1954, 162, 14, 9-17; Badenian near eastern (Kiipper, p. pi. figs [Langhian-Serravallian], Romerberg Baden, Austria) Pseudaspidura hungarica Kolosvary, 1941

Kiscell (Kolosvary, 1941, pp. 307, 308, figs 1-3 [not an ophiuroid at all?]; Oligocene, Clay, Hungary indeterminateophiuroids

Middle and northern the (Vadasz, 1915, pp. 93, 94, figs 4, 5; Miocene; Matraverebely Kemence, Hungary; specimen

in fig. 5 is assignable to the Euryalae according to Sieverts-Doreck, 1954)

1928, 31, 32, 5.1a-e, 5.2a-e as brachials of (Valette, pp. figs [described ‘Antedon’ rhodanicus]; Burdigalian, Angeles (Card), France)

Badenian and (Kiihn, 1952, p. 124; Eggenburgian [Lower Burdigalian] and [Langhian-Serravallian], Horn, Brunn

Rauchstallbrunngraben near Baden, northeast Austria)

(Tollmann, 1955, table 1; Badenian [Langhian-Serravallian], Eisenstadt, GroBhoflein and Miillendorf, eastern Austria)

Badenian (Kristan-Tollmann, 1964, p. 79; Lower [Langhian], Eisenstadt, eastern Austria)

(Kroh & Harzhauser, 1999, pp. 153, 154, pi. 4, figs 5-13; Upper Eggenburgian [Lower Burdigalian], Untemalb, north-

east Austria)

ophiuroid trace fossils

(Prakfalvi, 1992, figs 9, 10; Eggenburgian [Lower Burdigalian], Matraszele, northern Hungary)

Asteriacites lumbricalis von Schlotheim, 1820 (interpreted as ophiuroid resting traces by Seilacher, 1953)

(Heer, 1865,p. 440 as .Seesterne’; Muschelsandstein [Burdigalian], Reiden near Luzern, Switzerland)

eines Muschelsandstein (Bachmann, 1868, p. 250 as ,Abdrticke Seestern’; [Burdigalian], Reiden near Luzern, Switzer- land)

(Kaufmann, 1872, pp. 279, 281 as ,Seesteme’ and ,Astropecten helveticus’; Burdigalian, Reiden, Switzerland)

(Mayer, 1872, p. 489 as ,Astropecten helveticus May.’; Muschelsandstein, Helvet II and III [Burdigalian], near Bern,

Switzerland)

(Seilacher, 1953, pp. 94-102; Molasse [Burdigalian], Switzerland)