GeolCarp_Vol54_No6_395

CRETACEOUS BRYOZOA FROM SCHARRERGRABEN (EASTERN ALPS) 395

GEOLOGICA CARPATHICA, 54, 6, BRATISLAVA, DECEMBER 2003

395407

CRETACEOUS BRYOZOA FROM SCHARRERGRABEN

(SANTONIAN, GOSAU GROUP, EASTERN ALPS)

KAMIL ZÁGOREK

and ANDREAS KROH

Department of Paleontology, National Museum, Vaclavské nám. 69, CZ-115 79 Prague, Czech Republic; kamil.zagorsek@nm.cz

Institute of Geology and Palaeontology, Karl-Franzens-Universität Graz, Heinrichstraße 26, A-8010 Graz, Austria; discometra@gmx.at

(Manuscript received November 18, 2002; accepted in revised form June 23, 2003)

Abstract: The bryozoan fauna of the famous coral-locality Scharrergraben near Piesting, Lower Austria consists of 18

taxa, 13 of which encrust corals or more rarely rudists. This fauna is one of the richest bryozoan faunas described from

the Late Santonian of Europe, a time slice in which few bryozoan faunas have been studied. Thus the fauna of

Scharrergraben is an important piece in the puzzle of Santonian bryozoan biogeography. The faunal composition indi-

cates a shallow marine, relatively strongly agitated habitat with a moderate sedimentation rate in a tropical setting.

Key words: Santonian, Austria, Northern Calcareous Alps, Gosau Group, Bryozoa, taxonomy, paleoecology.

Introduction

Scharrergraben near Piesting, Lower Austria is a famous and

important locality within the Gosau Group. Fossils from this

outcrop have been studied for more than 150 years and vari-

ous taxonomic groups have been reported. The coral fauna of

Scharrergraben is especially abundant and diverse. First noted

by Reuss (1854), the fauna includes 60 coral taxa (Beauvais

1982), placing it among the richest in the Eastern Alps. Apart

from corals, bivalves (Zittel 186566; Stur 1877; Bittner

1882), gastropods (Stur 1877; Bittner 1882), cephalopods

(Brinkmann 1935; Summesberger 1997) and a single bryozo-

an species (Stur 1877) have been reported from this outcrop.

Bryozoans are very rare and usually poorly preserved in the

Upper Cretaceous sediments of Austria. Only two papers

dealing with bryozoans from these strata have been published

up to now: one (Reuss 1854) concerns the Santonian bryozo-

an fauna from the locality Neffgraben near Gosau, Upper

Austria; the other considers a small fauna from the Santonian

strata of Kössen, Tyrol (Voigt 1928). Nearly all taxa known

from these two outcrops are encrusting species attached to

corals or rudists. Since bryozoans are generally rather rare in

the Santonian strata of southern Europe, where very few local-

ities with biogeographically important bryozoan faunas exist

(Voigt 1979, 1983), the bryozoan fauna recovered from the

locality Scharrergraben adds an important piece to the puzzle

of Tethyan bryozoans.

Study area and geological setting

The Gosau Group, which comprises Upper Cretaceous to

Paleogene sediments, lies unconformably on folded and fault-

ed rocks of Permian to Early Cretaceous age (Wagreich &

Faupl 1994; Sanders et al. 1997; Faupl & Wagreich 2000).

Whereas the Late Cretaceous transgression started during the

Late Turonian in the western part of the Northern Calcareous

Alps (NCA) (e.g. in the type area of the Gosau Group; Sum-

mesberger & Kennedy 1996), it began earlier in the south-east

of the NCA (Wagreich & Marschalko 1995).

In the GrünbachNeue WeltPiesting area, the sediments of

the Gosau Group form a continuous outcrop belt in the Grün-

bach Syncline (Plöchinger 1961, 1967). These sediments

were the subject of geological and paleontological studies in

the first part of the 19

century and are well known for their

Upper Cretaceous fossils (Zekeli 1852; Zittel 186566).

These studies profited from exploitation of coal seams within

the Grünbach Formation (Cjek 1851; Summesberger 1997),

during which also the only Austrian reptile fauna of Late Cre-

taceous age was discovered (Bunzel 1871; Seeley 1881). Al-

Fig. 1. Distribution of the Cretaceous sediments north of Piesting.

The position of the studied section is indicated by an : (based on

the geological map of Austria, Sheet 76, Brix & Plöchinger 1982).

Small inset in upper right corner shows the position of Piesting in

Austria (indicated by a star).

396 ZÁGOREK and KROH

though the existence of an overturned syncline had already

been recognized by Cjek (1851), and several studies on

litho- and biostratigraphy of this area have been published re-

cently (Summesberger 1997; Hradecká et al. 1999; Summes-

berger et al. 2000 and in print), the structural relationship to

the Upper Cretaceous sediments in the north-eastern part of

this area, around Piesting, is still poorly known. This is caused

partly by the lack of outcrops in this area and by Tertiary

strike-slip faulting along the E-W trending Piesting valley

(Decker & Peresson 1996), which complicates the already

complex situation.

The locality Scharrergraben is situated northeast of Piesting

in a small valley between the two hills Steinkogel and Kranz-

kogel. Outcrop conditions are very poor and no section could

be taken. However, a schematic cross-section and short pro-

file of the locality can be found in Karrer (1877: 87), Stur

(1877: 172), Bittner (1882: 239) and Summesberger (1997:

8687). Exposed are fossiliferous sandy marls with a rich cor-

al fauna, interbedded between marls and sandstones with con-

glomerate intercalations. The age of the section is Late Santo-

nian according to Summesberger (1997: 86), who reported

Placenticeras polyopsis (Dujardin) from beds just below the

coral layer. The occurrence of Hemitissotia randoi Gerth in

a neighbouring valley indicates Early Santonian for the basal

part of the succession of Piesting (Summesberger 1997: 86).

Material and methods

The specimens were obtained by collecting fragments of

corals from weathered material in the outcrop. The corals

were cleaned using H

and ultrasonically. After drying all

specimens were carefully scanned for encrusting bryozoans.

Additionally, two bulk samples of approximately 5 kg each

were taken and washed using H

to macerate the samples.

From the washed material bryozoans (erect and encrusting)

were picked using a binocular microscope. Specimens for mi-

crophotography were further cleaned using treatments of

, ultrasonic and Quaternary O (see Zingula 1968). To

obtain further material unweathered sediment was dissolved

in concentrated acetic acid (for a detailed description of the

method see Zágorek & Vávra 2000). However, the residue

did not yield any recognizable bryozoans.

All photos were taken with a Jeol type JSM-6400 SEM at

the Institute of Palaeontology, University of Vienna. All ma-

terial used in this study is deposited in the collection of the

Natural History Museum, Vienna (Department of Geology &

Palaeontology). Specimens which could not be determined

due to their poor preservation have been deposited under the

inventory number NHMW 2002z0119/0067.

Systematics

The systematics of Cyclostomata have been modified on the

basis of Bassler (1953) and Taylor (pers. com. 2001, 2003).

The systematics of Cheilostomata are modified according to

Gordon (1989, pers. com. 2002) and Zágorek (in print). All

measurements are given in µm in Table 1.

Class Stenolaemata Borg, 1926

Order Cyclostomata Busk, 1852

Suborder Tubuliporina Milne Edwards, 1838

Family Diastoporidae Gregory, 1899

Genus Berenicea Lamouroux, 1821

Berenicea sp.

Fig. 2a

?v*1877 Berenicea radians nov. sp.; Novák p. 22, Pl. 4, Figs. 1518

Material: 3 specimens from the locality Scharrergraben

(NHMW 2002z0119/0007, 0008a, 0009).

Description: The colony is encrusting, very thin, uniseri-

al and fan-shaped. The autozooecia are tubular and grow radi-

ally from the centre of the colony. They develop a long peris-

tome which protrudes from the colonial margin. The distal

fringe of the basal lamina is very narrow. Brood chambers

were not observed.

Remarks and notes: External features are identical

with the specimens described by Novák (1877) as Berenicea

radians, stored in the National Museum in Prague (Inv. No.

ÈL 2429), but none of the colonies developed brood-chambers

and identification to species level is impossible. As gonozooe-

cia are necessary even for generic identification of bryozoans

of this type, the studied specimens can only be referred to the

form-genus Berenicea following the protocol suggested by

Taylor & Sequeiros (1982).

Distribution in time and space: Early Turonian (?)

to Late Santonian. (?) Kamajka (Czech) Early Turonian.

Scharrergraben (Austria) Late Santonian.

Family Oncousoeciidae Canu, 1918

Genus Proboscina Audouin, 1826

Proboscina bohemica Novák, 1877

Fig. 2b

v*1877 Proboscina bohemica nov. sp.; Novák p. 25, Pl. 5, Figs. 24, 25

Material: 7 specimens from the locality Scharrergraben

(NHMW 2002z0119/0055a, 0055b, 0056, 0057a, 0058,

0059a, 0060).

Description: The colonies are multiserial encrusting, cor-

responding to the runner type. Usually 47 parallel longitu-

dinal zooecial rows are developed. The outer shape of the colo-

nies is irregular, with one, rarely two, branches situated on

opposite sides of the ancestrula. Young colonies show the so-

called Stomatopora stadium (Hillmer 1971), when the colo-

ny is uniserial encrusting. The autozooecia are tubular, rela-

tively short (in comparison with other species of Proboscina),

with large, circular, terminal apertures. Gonozooecia were not

observed.

Remarks and notes: In most of the studied specimens

only one branch leading away from the ancestrula is devel-

oped. However, the type material stored in the National Muse-

um in Prague (Inv. No. ÈL 6297) usually developed two

branches. Apart from this, all other features are identical. Char-

acteristic features of this species are short autozooecial tubes,

CRETACEOUS BRYOZOA FROM SCHARRERGRABEN (EASTERN ALPS) 397

the obscured shape of the autozooecia and the irregular shape

of the colonial branches.

Pitt & Taylor (1990) argued that Proboscina could be used

only informally as Proboscina because of doubt over the

identity of the type species (the type material of the type spe-

cies is unknown).

Distribution in time and space: Coniacian to Late

Santonian. Velký Újezd by Kolín (Czech) Coniacian.

Scharrergraben (Austria) Late Santonian.

Family Eleidae dOrbigny, 1852

Genus Reptomultelea dOrbigny, 1853

Reptomultelea sp. aff. Elea lamellosa (dOrbigny, 1850)

Fig. 2ij

? 1850 Bidiastopora lamellosa dOrb., 1847; dOrbigny p. 266

?v1922 Meliceritites lamellosa dOrbigny; Canu & Bassler p. 85, Pl.

14, Fig. 13

? 1994 Elea lamellosa (dOrbigny 1852), Taylor p. 14, Figs 1728

(cum. syn)

Material: 7 specimens from the locality Scharrergraben

(NHMW 2002z0119/00260032).

Description: The colony is encrusting or unilamellar.

The autozooecia have a slightly porous frontal wall and a

semilunar to oval aperture usually with a straight proximal

margin. The distal fringe of the basal lamina is very narrow,

sometimes not preserved. Gonozooecia and avicularia (eleo-

zooecia) were not observed.

Remarks and notes: The studied specimens very

much resemble Elea lamellosa (dOrbigny) in shape and size

of autozooecia and pattern of distribution of pseudopores.

Elea has, however, erect colonies, the large, globular gonozo-

oecia and E. lamellosa has a terminal oeciopore (Canu &

Bassler 1922; Taylor 1994). The encrusting grown form of the

studied specimens suggests placement in Reptomultelea

dOrbigny, 1853 as revised in Taylor (1994). Although we

have seven specimens in our collection, none of them show

gonozooecia or avicularia (eleozooecia) and therefore spe-

cies determination remains uncertain. The studied specimens

represent, with large probability, a new species, but more and

better-preserved specimens are needed to establish this.

Taxa

measured individual

Aperture

Ovicell

Inv. No.

length width length width length width

Akatopora subwintonensis (Voigt, 1967)

Autozooecium+ovicell

383

163

141

NHMW 2002z0119/0012b

Autozooecium

331

215

183

131

NHMW 2002z0119/0012b

Avicularium (adventitious)

174

119

NHMW 2002z0119/0012b

Avicularium (vicarious)

136

128

NHMW 2002z0119/0012a

Alderina dilatata (Reuss, 1872)

Autozooecium+ovicell

527

301

353

217

149

165

NHMW 2002z0119/0003

Autozooecium

438

303

357

220

NHMW 2002z0119/0003

Avicularium

603

343

452

196

NHMW 2002z0119/0003

"Berenicea" sp.

Autozooecium

502

306

NHMW 2002z0119/0008a

Autozooecium

437

257

106

NHMW 2002z0119/0008a

Biflustra cf. teres (Novák, 1877)

Autozooecium

580

341

230

157

NHMW 2002z0119/0010

Disporella cf. confluens (Roemer, 1841)

Autozooecium

155

111

NHMW 2002z0119/0022

Marginaria ostiolata Reuss, 1846

Autozooecium+ovicell

537

236

203

136

160

140

NHMW 2002z0119/0024

with vicariuos avicularia

Autozooecium

455

247

275

159

NHMW 2002z0119/0024

Avicularium (vicarious)

301

201

119

NHMW 2002z0119/0024

Avicularium (adventitious)

173

NHMW 2002z0119/0024

Marginaria ostiolata Reuss, 1846

Autozooecium+ovicell

392

283

169

124

155

129

NHMW 2002z0119/0025

Autozooecium

321

271

177

148

NHMW 2002z0119/0025

Avicularium (adventitious)

186

118

NHMW 2002z0119/0025

Membraniporidra bohemica (Prantl, 1938)

Autozooecium

323

243

211

150

NHMW 2002z0119/0035

Autozooecium

292

238

177

137

NHMW 2002z0119/0035

Onychocella cyclostoma (Goldfuss, 1826)

Autozooecium

624

453

293

251

NHMW 2002z0119/0036

Autozooecium

563

448

262

240

NHMW 2002z0119/0036

Avicularium

902

425

600

188

NHMW 2002z0119/0036

Onychocella depressa (v. Hagenow, 1851)

Autozooecium

543

441

117

219

NHMW 2002z0119/0038

Autozooecium

494

426

158

185

NHMW 2002z0119/0038

Avicularium

707

340

264

211

NHMW 2002z0119/0038

Onychocella michaudiana (d'Orbigny,1850)

Autozooecium+ovicell

337

223

109

147

NHMW 2002z0119/0041a

Autozooecium

292

265

111

NHMW 2002z0119/0041a

Avicularium

299

160

NHMW 2002z0119/0041a

Onychocella pseudoirregularis Voigt,1924

Autozooecium+ovicell

381

292

121

119

150

NHMW 2002z0119/0039b

Autozooecium

342

206

NHMW 2002z0119/0039b

Avicularium

428

111

NHMW 2002z0119/0039b

Onychocella reussi Prantl,1938

Autozooecium+ovicell

512

194

100

115

134

NHMW 2002z0119/0041b

Autozooecium

388

233

101

105

NHMW 2002z0119/0041b

Avicularium

370

122

NHMW 2002z0119/0041b

Osculipora truncata Goldfuss,1826

Autozooecium

437

108

NHMW 2002z0119/0046

Autozooecium

661

122

NHMW 2002z0119/0048

"Proboscina" bohemica Novák, 1877

Autozooecium

235

163

NHMW 2002z0119/0060

Reptomultelea sp. aff. Elea lamellosa

Autozooecium

337

285

119

102

NHMW 2002z0119/0031

(dOrbigny,1850)

Autozooecium

341

252

129

106

NHMW 2002z0119/0031

Table 1

398 ZÁGOREK and KROH

Distribution in time and space: Late Santonian.

Scharrergraben (Austria) Santonian.

Suborder Cancellata Gregory, 1899

Family Cytididae dOrbigny, 1854

Genus Osculipora dOrbigny, 1849

Osculipora truncata Goldfuss, 1826

Fig. 2de

*1826 Retepora truncata nobis.; Goldfuss p. 29, Pl. 9, Fig. 14ad

1922 Osculipora truncata Goldfuss; Canu & Bassler p. 57, Pl. 23,

Figs. 16

Material: 9 specimens from the locality Scharrergraben

(NHMW 2002z0119/00430051).

Description: The colony is erect, sometimes dichoto-

mously branching with autozooecial apertures opening on one

side only. The autozooecia are tubular with apertures opening

in cluster (fascicles). Each cluster comprises 610 autozooe-

cial apertures. The clusters are circular to oval, slightly elevat-

ed from the frontal surface of the colony. The dorsal side of

the colony is slightly ribbed and porous due to the presence of

short kenozooecia (so-called nematopores of Canu & Bassler

1922). Gonozooecia have not been observed.

Remarks and notes: According to Canu & Bassler

(1922), the gonozooecia are globular and developed on the

dorsal or the frontal side of the colony, with a non-porous,

slightly granular frontal wall. The studied specimens do not

have gonozooecia, but in all other features are identical with

the type material of O. truncata. Therefore we believe that the

studied specimens and the type material are conspecific. This

species is very common in the Campanian strata of France.

Distribution in time and space: Late Santonian to

Maastrichtian. Scharrergraben (Austria) Late Santonian.

Many localities in France Campanian. Maastricht (Nether-

lands) Maastrichtian.

Family Petaloporidae Gregory, 1899

Genus Petalopora Lonsdale, 1850

Petalopora? sp.

Fig. 2f

? 1953 Petalopora Lonsdale; Bassler p. 63, Fig. 30/1

Material: 3 specimens from the locality Scharrergraben

(NHMW 2002z0119/00520054).

Description: The colony is erect, sometimes dichoto-

mously branching with cylindrical branches. The autozooecia

are long, tubular and arranged around the colonial stems.

Kenozooecia (mesopores) are abundant and arranged in 25

longitudinal rows proximally from the autozooecial apertures.

Gonozooecia were not observed.

Remarks and notes: Species of Petalopora typically

have a fixed number of rows of kenozooids (Taylor 2002).

The studied specimens have, however, different numbers of

rows of kenozooecia, and probably represent more than one

species. Due to the poor preservation of the studied specimens

and the lack of gonozooecia, more precise generic and species

determination is not possible.

Distribution in time and space: Late Santonian.

Scharrergraben (Austria) Late Santonian.

Genus Reteporidea dOrbigny, 1849

Reteporidea? sp.

Fig. 2h

? 1953 Reteporidea dOrbigny; Bassler p. 63, Fig. 30/1

Material: 2 specimens from the locality Scharrergraben

(NHMW 2002z0119/00610062).

Description: The colony is erect and branching. Auto-

zooecia are arranged in uniserial fascicles on one side of the

colony only. Kenozooecia (mesopores) are very abundant, sit-

uated around the entire branch circumference. Gonozooecia

were not observed.

Remarks and notes: On the basis of the presence of

numerous mesopores and the arrangement of the autozooecia

in fascicles, these specimens are tentatively referred to Rete-

poridea. Precise determination is impossible due to the very

poor preservation.

Distribution in time and space: Late Santonian.

Scharrergraben (Austria) Late Santonian.

Suborder Rectangulata Waters, 1887

Family Lichenoporidae Smitt, 1866

Genus Disporella Gray,1848

Disporella cf. confluens (Roemer, 1841)

Fig. 2g

*1841 Rosacilla confluens N; Roemer p. 19

v?1877 Berenicea confluens Roemer; Novák p. 22, Pl. 4, Figs. 1922

Material: 2 specimens from the locality Scharrergraben

(NHMW 2002z0119/0021, 0022).

Description: The colony is encrusting, discoidal with

central maculae. The maculae are surrounded by a relatively

steeply sloping margin and have non-porous and slightly

granular surface, possibly the remnants of gonozooecia. The

autozooecia are short with oval to circular apertures without

peristomes. The apertures are arranged in uniserial, slightly

visible radial ribs. Rarely, an autozooecial aperture may occur

also in the central macula. Kenozooecia are slightly smaller

than autozooecia. The distal fringe of the basal lamina is very

narrow and usually not preserved. Gonozooecia are possibly

developed at maculae centres, but no frontal walls are pre-

served.

Remarks and notes: The studied specimens are al-

most identical with Nováks (1877) specimens stored in the

National Museum in Prague except for the less steeply sloping

margin of the macular centre. The main characteristic feature

of this species is the occurrence of autozooecial apertures in-

side the maculae centre.

Disporella Gray, 1848 has non-stalked colonies, with a

non-porous basal part, and autozooecia arranged on the fron-

tal surface in radial fascicles with kenozooecia situated be-

tween them (Brood 1972). Due to the presence of gonozooe-

cia in the maculae centre and arrangement of autozooecia in

400 ZÁGOREK and KROH

radial ribs with development of smaller kenozooecia between

them, this species is listed under Disporella.

Roemer (1841) described Berenicea confluens with gono-

zooecia, but unfortunately, he did not illustrate them. Accord-

ing to Voigt (pers. com., 1980) the type material is lost. Be-

cause Nováks (1877) specimens developed no gonozooecia

and the studied specimens have only remnants of the gonozo-

oecia, the correct species determination of the present speci-

mens remains uncertain.

Distribution in time and space: Cenomanian to

Late Santonian. Kamajka and Zbislav (Czech) Cenoma-

nian. Scharrergraben (Austria) Late Santonian.

Disporella sp.

Fig. 2c

*1929 Lichenopora perneri sp.n.; Prantl p. 250, Pl. 1, Figs. 12

?v1938 Lichenopora perneri Prantl 1929; Prantl p. 29, (64), Pl. 1,

Fig. 20

Material: 1 specimen from the locality Scharrergraben

(NHMW 2002z0119/0023).

Description: The colony is encrusting, discoidal with

central, circular, porous maculae. Autozooecia are slightly

raised, oval to circular in shape without a peristome. Autozoo-

ecia are arranged chaotically and not in radial ribs and have

almost the same size as kenozooecia. The kenozooecia are

slightly larger in the maculae centre. The distal fringe of the

basal lamina is narrow, but always developed. Gonozooecia

are unknown.

Remarks and notes: The studied specimen is very

similar to Lichenopora perneri Prantl, 1929 in general shape

and development of the colony. We have only one specimen of

this species, which developed no gonozooecia. Therefore spe-

cific determination is impossible.

According to Gordon & Taylor (1997), Lichenopora should

be restricted to species characterized by conical/stalked (pe-

dunculate) colonies, with apertures only on the top of the

cone. The side of the cone is formed by exterior wall and the

surface shows a smooth texture. Disporella Gray, 1848, in

contrast, has non-stalked colonies, with autozooecia arranged

in radial fascicles on the frontal part and kenozooecia situated

between the autozooecia (Brood 1972). The specimen from

Scharrergraben shows these features and is therefore placed in

the genus Disporella.

Distribution in time and space: Early Turonian to

Late Santonian. (?) Pøedboj (Czech) Early Turonian.

Scharrergraben (Austria) Late Santonian.

Class Gymnolaemata Allman, 1896

Order Cheilostomata Busk, 1852

Suborder Malacostegina Levinsen, 1902

Superfamily Membraniporoidea Busk, 1852

Family Membraniporidae Busk, 1852

Genus Biflustra dOrbigny, 1852

Biflustra cf. teres (Novák, 1877)

Fig. 2k

?v*1877 Semieschara teres nov. sp.; Novák p. Pl. 3, Figs. 811

Material: 1 specimen from the locality Scharrergraben

(NHMW 2002z0119/0010).

Description: The colony is erect, developing cylindrical

branches of almost circular cross-section. Autozooecia are ar-

ranged in parallel, longitudinal rows. They are rhomboidal

with an elongated opesia and a slightly depressed cryptocyst.

Ovicells, avicularia and kenozooecia were not observed.

Remarks and notes: The shape and size of autozooe-

cia of the studied specimen are very similar to Semieschara

teres (Novák, 1877) stored in the National Museum in Prague

(Inv. No. ÈL 444). The size of the colonies of the type materi-

al is, however, much larger and differs, furthermore, in the

presence of ovicells distally of the autozooecia. On the basis

of the presence of features characteristic for Biflustra, (i.e.

large opesia with flat cryptocyst and erect growth form of the

colony) this species is placed into this genus.

Distribution in time and space: (?) Turonian to

Late Santonian. (?) Kamajka (Czech) Early Turonian.

Scharrergraben (Austria) Late Santonian.

Suborder Flustrina Smitt, 1868

Superfamily Calloporoidea Norman, 1903

Family Calloporidae Norman, 1903

Genus Akatopora Davis, 1934

Akatopora subwintonensis (Voigt, 1967)

Fig. 3ac

*1967 Membranipora subwintonensis sp.n.; Voigt p. 28, Pl. 4, Figs. 46

Material: 6 specimens from the locality Scharrergraben

(NHMW 2002z0119/0011, 0012a, 0012b, 0013, 0020, 0066).

Description: The colony is encrusting. Autozooecia are

oval to triangular with distal margin slightly narrower than

proximal margin. The opesium is large, almost triangular (nar-

rower distally); the mural rim is narrow, granular or slightly

radially ribbed. Two pairs of oral spines are situated on the

distalmost margin of mural rim. The distal pair of the oral

spines is always smaller than the proximal one. Two types of

avicularia occur: large, vicarious rhomboidal avicularia,

which are rare and have a narrow, oval opesium; and smaller

oval to polygonal avicularia. This second type of the avicular-

ium is interzooecial, situated between 2 or 3 autozooecia and

often shows a long process (peristome-like structure). The ov-

icells are hyperstomial, globular and are deeply immersed in-

side the secondary calcification of the colony.

Remarks and notes: The studied specimens have

smaller vicarious avicularia than the specimens described by

Voigt (1967). In our material the vicarious avicularia are

about half the size of the autozooecia and the opesium is sub-

circular. Other features, especially type and position of ovice-

lls, are identical with the type material as described by Voigt

(1967). The differences observed are therefore considered to

represent intraspecific variation.

Based on the presence of interzooecial avicularia and deep-

ly immersed hyperstomial ovicells, this species is listed under

the genus Akatopora (as revised in Gordon 1986).

Distribution in time and space: Late Santonian to

Campanian. Scharrergraben (Austria) Late Santonian. Lo-

cality unknown (west Kazakhstan) Campanian.

402 ZÁGOREK and KROH

Genus Alderina Norman, 1903

Alderina dilatata (Reuss, 1872)

Fig. 3df

*1872 Membranipora dilatata n. sp.; Reuss p. 100; Pl. 24, Fig. 2

v1877 Membranipora perisparsa nov. spec.; Novák p. 15, Pl. 2, Figs.

1989 Alderina dilatata (Reuss), Voigt p. 19, Pl. 1, Fig. 1-4, Pl. 2,

Figs. 35

Material: 6 specimens from the locality Scharrergraben

(NHMW 2002z0119/0001a, 00020006).

Description: The colony is encrusting. The zooecia are

oval with large oval to irregular opesia and without gymno-

cyst. The mural rim is narrow, rarely with intramural buds,

and without any traces of spines. A small tubular kenozooeci-

um is sometimes developed inside the narrow interzooecial

space. Avicularia are vicarious, as large as the autozooecia

and have a well developed pivotal bar. Ovicell globular, hy-

perstomial, slightly immersed with a smooth, non-porous

frontal wall.

Remarks and notes: The ovicells of the studied speci-

mens are identical with those described by Voigt (1989). The

avicularia, however, are slightly larger in the present material,

probably demonstrating intraspecific variability in this fea-

ture. According to Voigt (1989), Membranipora perisparsa

Novák, 1877 is a junior synonym of Alderina dilatata (Reuss,

1872).

Distribution in time and space: Early Cenoma-

nian to Late Santonian. Mülheim-Broich (Germany) Early

Cenomanian. St. Calais (France) Late Cenomanian. Kama-

jka (Czech) Early Turonian. Scharrergraben (Austria)

Late Santonian.

Genus Membraniporidra Canu & Bassler, 1917

Membraniporidra bohemica (Prantl, 1938)

Fig. 3g

v1877 Membranipora elliptica Reuss; Novák p. 84, Pl. 2, Figs. 1114

non1877 Membranipora elliptica Reuss; Novák p. 84, Pl. 2, Figs. 1516

1938 Membranipora bohemica nov. sp.; Prantl p. 5 (41)

1989 Membraniporidra bohemica (Prantl); Voigt p. 25, Pl. 2, Figs.

12, Pl. 7, Fig. 7

Material: 4 specimens from the locality Scharrergraben

(NHMW 2002z0119/00330035, 0008b).

Description: The colony is encrusting. The autozooecia

are oval with very large opesia and short, but well developed,

gymnocyst. No cryptocyst is developed. The mural rim is nar-

row, usually smooth and only rarely with traces of oral spines.

Avicularia absent, ovicells not observed.

Remarks and notes: Prantl (1938) established the

species M. bohemica on the basis of the material of Novák

(1877) described as Membranipora elliptica Reuss (Novák

1877, p. 84, Pl. 2, Figs. 1112). According to Voigt (1989)

specimens of Membranipora elliptica Reuss as illustrated

by Novák (1877, Pl. 2, Figs. 1516) do not belong to this

species.

The studied specimens are almost identical with the type

material stored in the National Museum in Prague (Inv. No.

ÈL 457). A minor difference is the narrower mural rim.

Distribution in time and space: Turonian to Late

Santonian. Many localities in Czech Turonian. Schar-

rergraben (Austria) Late Santonian.

Genus Marginaria Roemer, 1841

Marginaria ostiolata Reuss, 1846

Fig. 3hj

*1846 Marginaria ostiolata Reuss; Reuss p. 69, Pl. 15, Fig. 14

1989 Marginaria ostiolata (Reuss); Voigt p. 33, Pl. 5, Figs. 15, Pl.

13, Fig. 2

Material: 2 specimens from the locality Scharrergraben

(NHMW 2002z0119/0025 and (NHMW 2002z0119/0024).

Description: The colony is encrusting and multilamel-

lar. The autozooecia are oval to triangular with a large opesi-

um. The gymnocyst is short or well developed, smooth and

slightly granular. Mural rim wide, sometimes with an intramu-

ral bud, without any traces of mural spines. Two types of avic-

ularia are present: interzooecial and vicarious. The interzooe-

cial avicularia are drop-like with an oval opesium, sometimes

with pivotal bar and tapering distally. Usually, among four

autozooecia, there are one or two avicularia. The vicarious

avicularia are as large as an autozooecium, rare and rhomboi-

dal with a small oval opesium. The ovicells are hyperstomial,

globular with smooth frontal wall.

Remarks and notes: The studied specimen developed

fewer avicularia than the specimens described and illustrated

by Voigt (1989), which show up to 5 avicularia around each

autozooecium. On the basis of the development of the ovicells

and the general features of the colony, the specimen is deter-

mined as Marginaria ostiolata Reuss, 1846. Although vicari-

ous avicularia are previously unknown in Marginaria ostiola-

ta, according to Taylor (pers. comm. 2003) it is not unusual,

for calloporid species to develop occasional larger avicularia.

On the other hand, the size of the autozooecia and avicularia

shows differences between specimens with or without large

vicarious avicularia (see Table 1). More and better preserved

specimens are needed to solve the question whether one or

two species are represented.

Distribution in time and space: Early Cenoma-

nian to Late Santonian. Mülheim-Broich (Germany) Early

Cenomanian. St. Calais (France), Bílina (Czech) Late Cen-

omanian. Kamajka (Czech) Early Turonian. Scharrergra-

ben (Austria) Late Santonian.

Superfamily Microporoidea Gray, 1848

Family Onychocellidae Jullien, 1881

Genus Onychocella Jullien, 1882 (=Semieschara dOrbigny,

1852)

Onychocella cyclostoma (Goldfuss, 1826)

Fig. 4a

*1826 Eschara cyclostomata nobis: Goldfuss p. 23, Pl. 8, Fig. 9ac

1985 Onychocella cyclostoma (Goldfuss): Voigt p. 63, Figs. 34

404 ZÁGOREK and KROH

rial described by Voigt (1985). In some cases, our material

shows a slightly larger cryptocyst than that of Voigt (1985).

Apart from this feature the specimens are very similar.

The characteristic feature of Onychocella cyclostoma

(Goldfuss, 1826) is a very short cryptocyst, which may be ab-

sent altogether.

Distribution in time and space: Late Santonian to

Maastrichtian. Scharrergraben (Austria) Late Santonian.

Maastricht (Netherlands) Maastrichtian.

Onychocella depressa (v. Hagenow, 1851)

Fig. 4b

*1851 Cellepora (Discop.) depressa, Hag.: von Hagenow p. 93; Pl. 11,

Fig. 13ab

1989 Onychocella depressa (v. Hagenow); Voigt p. 57, Pl.13, Figs. 36

Material: 1 specimen from the locality Scharrergraben

(NHMW 2002z0119/0038).

Description: The colony is encrusting or unilamellar.

The autozooecia are rhomboidal to oval and are arranged in

regular longitudinal rows. The opesia occupy almost the en-

tire distal halves of the autozooecia and have enlarged proxi-

mo-lateral corners for parietal muscles. The cryptocyst is well

developed, almost flat to concave, and slightly granular. Ad-

ventitious avicularia (onychocellaria) are large, longitudinally

narrow about half of the width of the autozooecia and tapering

on both sides. The opesia of avicularia are small and oval.

Remarks and notes: The studied specimen is encrust-

ing and shows wider avicularia than specimens described by

Voigt (1989). All other features are identical, suggesting that

these specimens are conspecific.

Onychocella depressa (v. Hagenow, 1851) differs from oth-

er similar species of Onychocella in developing a large con-

cave cryptocyst.

Distribution in time and space: Late Santonian to

Maastrichtian. Scharrergraben (Austria) Late Santonian.

St. Pietersberg by Maastricht Maastrichtian.

Onychocella michaudiana (dOrbigny, 1850)

Fig. 4c

*1850 Escharina michaudiana, dOrb. 1847; dOrbigny p. 175

v1938 Aechmella michaudiana (dOrbigny); Prantl p. 8, Pl. 1, Fig. 1

1989 Onychocella michaudiana (dOrbigny); Voigt p. 63, Pl. 10,

Figs. 110

Material: 3 specimens from the locality Scharrergraben

(NHMW 2002z0119/0039a, 0040a, 0041a).

Description: The colony is encrusting and small. The

autozooecia are rhomboidal to hexagonal with a distally de-

veloped mural rim. The cryptocyst is well developed, concave

and slightly granular. The opesia are small, oval, sometimes

with enlarged proximo-lateral corners, or with a straight prox-

imal margin. The avicularia (onychocellaria) are usually as

long as the autozooecia, but are only half as wide and are

sharply tapering distally. The distalmost end of the aviculari-

um (palate) is sometimes very narrow and slightly curved lat-

erally. The avicularian opesia are oval to semicircular and nar-

row. Ovicells are very rare, globular, probably endozooidal,

deeply immersed, thereby slightly deforming the distal neigh-

bour autozooecia.

Remarks and notes: The avicularia are distinctly ta-

pering distally in the studied specimens, whereas in the mate-

rial described by Voigt (1989) the avicularia do not taper. The

specimens described by Prantl (1938), however, have identi-

cal avicularia to the specimens from Scharrergraben. Other

features visible on the studied specimens are identical with

those described by Voigt (1989) and Prantl (1938).

The characteristic features of Onychocella michaudiana

(dOrbigny, 1850) are a distal mural rim around the autozooe-

cia and the presence of small, immersed ovicells.

Distribution in time and space: Cenomanian to

Late Santonian. Le Mans (France) middle Cenomanian. St.

Calais (France) Late Cenomanian. Pøedboj and Velim by

Kolín (Czech) Turonian. Scharrergraben (Austria) Late

Santonian.

Onychocella pseudoirregularis Voigt, 1924

Fig. 4d

1851 Cellepora irregularis; v. Hagenow p. 92; Pl. 11, Fig. 14

*1924 Onychocella pseudoirregularis nom. nov.; Voigt p. 203205;

Pl. 7, Figs. 1314

1989 Onychocella pseudoirregularis Voigt; Voigt p. 53, Pl. 11, Figs.

Material: 3 specimens from the locality Scharrergraben

(NHMW 2002z0119/0040b, 0042, 0039b).

Description: The colony is encrusting with autozooecia

arranged in chaotic rows. The autozooecia are oval with a

well-developed distal part of the mural rim and a flat or slight-

ly convex, granular cryptocyst. The autozooecial opesia are

oval to elongated oval, very small. Adventitious avicularia

(onychocellaria) are abundant and have a long, straight and

very narrow distal part (palate). The ovicells are globular,

hardly visible because they are deeply immersed in the direc-

tion of the distal neighbouring autozooecium.

Remarks and notes: The studied specimens are iden-

tical with those described by Voigt (1989), except that the dis-

tal part of the avicularia is narrower. This slight difference in

the development of the avicularia may be considered as within

species variability.

The main differences between Onychocella pseudoirregu-

laris Voigt, 1924 and Onychocella michaudiana (dOrbigny,

1850) are in the size of the autozooecial opesia and the fact

that O. michaudiana has avicularia with curved palates.

Distribution in time and space: Late Santonian to

Maastrichtian. Scharrergraben (Austria) Late Santonian.

Many localities around Maastricht (Netherlands) Maas-

trichtian.

Onychocella reussi Prantl, 1938

Fig. 4e

v*1938 Onychocella reussi sp.n.; Prantl, p. 7 and 43

1989 Onychocella reussi Prantl; Voigt p. 54, Pl. 6, Figs. 69

Material: 1 specimen from the locality Scharrergraben

(NHMW 2002z0119/0041b).

CRETACEOUS BRYOZOA FROM SCHARRERGRABEN (EASTERN ALPS) 405

Description: The colony is encrusting. The autozooecia

are elongated with comparatively small oval opesia which

usually have straight distal margins. Sometimes enlarged

proximo-lateral corners for parietal muscles are developed.

The cryptocyst is flat, non-porous and smooth (not granular).

Adventitious avicularia (onychocellaria) are abundant and

have a short distal part (palate), which is sometimes slightly

curved. The avicularian opesia are oval, sometimes slightly

tapering proximally. The ovicells are small, hyperstomial

deeply immersed to the distal autozooecium. The frontal wall

of the ovicell is smooth, non-porous and slightly granular.

Remarks and notes: Apart from the less prominent

enlarged proximo-lateral corners of opesia in the type materi-

al, all other features, including the elongated shape of the au-

tozooecia and the shape and size of the avicularian palate, are

identical.

This species differs from other species of Onychocella

mainly in its elongated autozooecia, and the small opesia with

straight distal margin.

Distribution in time and space: Late Cenomanian

to Late Santonian. St. Calais (France) Late Cenomanian.

Many localities in Czech Turonian. Scharrergraben (Aus-

tria) Late Santonian.

Conclusions

The studied bryozoan assemblage represents one of the

most diverse bryozoan faunas reported from the Late Santo-

nian of Europe. A total of 18 species were identified, 8 of

which are Cyclostomata and 10 Cheilostomata.

Nearly all of the species found developed encrusting colo-

nies (13 of 18 species); other growth forms occur only rarely.

Four species developed delicate branching colonies and one

species robust branching colonies.

The encrusting bryozoans are nearly exclusively found on

the thecae of branched corals, only very few specimens are lo-

cated on the underside of massive coral colonies and on mol-

lusc shells (rudists). The dominance of encrusting bryozoans

with additional erect delicate branching forms in areas with

suitable substrates indicates an inter- to subtidal position with

moderate sedimentation rate and relatively strong wave action

according to McKinney & Jackson (1989) and Smith (1995).

The highly diverse coral fauna (more than 60 taxa; Beau-

vais 1982) of the locality Scharrergraben indicates optimal

living conditions for hermatypic corals, as found today in the

tropical climate zone. The dominance of encrusting growth

forms of Bryozoa indicates shallow water and warm temper-

ate conditions (McKinney & Jackson 1989) and therefore,

does not contradict the data of the corals. According to Smith

(1995), diverse bryozoan faunas are characteristic of nontropi-

cal settings. However, the importance of Bryozoa might be

underestimated in extant tropical settings, as were, for exam-

ple, brachiopods (Kowalewski et al. 2002).

Recent studies on the Campanian Grünbach flora (Herman

& Kvaèek 2002), from nearby localities suggested a humid

subtropical to marine mesothermal climate for the plant bear-

ing strata. However, the lithostratigraphic correlation and tec-

tonic relation of the Neue Welt Basin (including the Campa-

nian sediments bearing the Grünbach flora) with the strata

north of Piesting (Scharrergraben) is unclear (Summesberger

1997; Summesberger et al. in print). Paleomagnetic data for

the Gosau Basin (Preisinger et al. 1986), which indicate a pa-

leolatitude of 32°N for the Late Maastrichtian/Early Paleo-

gene of the Elendgraben-Section (Salzburg) are controversial

according to Herman & Kvaèek (2002). Paleomagnetic data

for the Neue Welt Basin (including the locality Scharrergra-

ben) indicate a paleolatitude of 20.6°±5.8° in the Late Santo-

nian to Early Maastrichtian (calculated by R. Scholger from

the Neue Welt area mean data presented in Haubold et al.

1999; pers. com. March 2003). Thus, the question on the pale-

oclimate of the marine strata exposed at Scharrergraben has to

remain open, but evidence suggests that it was tropical.

Although bryozoans are often abundant and well preserved

in Santonian chalk deposits of north-western Europe (e.g.

Taylor 2002) few faunas have been systematically studied.

Contrary to these faunas, which belong to the Kalaian paleo-

realm (new name for Boreal Province, see Kollmann 2002),

the Gosau Group belongs to the Theian paleorealm (new

name for Tethyan Province, see Kollmann 2002). Santonian

Bryozoa from the Gosau Group, however, are rather rare and

have been reported only in four other localities. Detailed in-

vestigations on their taxonomy have been done for just two of

these outcrops (Voigt 1979). Reuss (1854) was the first to de-

scribe the bryozoans from Neffgraben (Austria) in a mono-

graph on the fauna of Cretaceous sediments of the Eastern

Alps. Later, Voigt (1928) described another Santonian bryo-

zoan fauna from Kössen, Tyrol, Austria (originally described

as the locality Dichtleralm).

The bryozoan fauna from Neffgraben (Austria) is similar to

the fauna from Scharrergraben; in both localities mainly en-

crusting bryozoans attached to corals are found. Reuss (1854)

described seven genera, from which five also occur in the

studied section. A revision of the Neffgraben fauna is, howev-

er, needed for more detailed comparison.

Voigt (1928) reported six species in six different genera

from the locality Dichtleralm (Kössen in Tyrol). This bryozo-

an fauna is different from that of Scharrergraben; only one ge-

nus is present in both localities. Moreover, at Dichtleralm the

bryozoans are associated with molluscs and serpulids and rep-

resent, therefore, a different paleoecological setting.

Beside these two known bryozoan associations, Voigt

(1979) reported two more localities where bryozoans occur in

the Santonian of the Gosau Group: Edelbachgraben bei Gosau

(Austria) and Baumgarten near Rosenheim (Germany). These

faunas are similar to that of Scharrergraben in the respect that

here the bryozoans also encrusted corals and rudists. Howev-

er, since no systematic investigations on these faunas have

been published, a comparison is not possible.

Acknowledgments: We would like to express our thanks to:

C. Latal (Univ. Graz), R. Scholger (Univ. Leoben), H. Sum-

mesberger (NHM Vienna), V. Turek (Prague), M. Wagreich

(Univ. Vienna), the Geological-Palaeontological Department

of the Vienna Natural History Museum, Paleontological Insti-

tute of the National Museum in Prague and the Department of

Palaeontology, University of Vienna. We are greatly appreci-

ated the kind comments of N. Vávra (Univ. Vienna), E. Voigt

406 ZÁGOREK and KROH

(Hamburg) and P.D. Taylor (NHM London), which improved

the quality of this paper. The Ministry of Culture of Czech

Republic Project MK0CEZ99F0201 and Austrian Science

Fund Project No. P-14366-Bio covered parts of the costs.

References

Bassler R.S. 1953: Treatise on Invertebrate Paleontology, Part G

Bryozoa. Univ. Kan. Press, Lawrence KS, 1253.

Beauvais M. 1982: Révision systématique des Madréporaires des

Couches de Gosau (Crétacé supérieur, Autriche). Trav. Lab.

Paléont. Invertébres Univ. Pierre et Marie Curie 1: 1256, 2:

1277, 3: 1177, 4, 5.

Bittner A. 1882: Die Geologischen Verhältnisse von Hernstein in

Niederösterreich und der weiteren Umgebung. In: Becker M.A.

(Ed.): Hernstein in Niederösterreich, sein Gutsgebiet und das

Land im weiteren Umkreise. I. Theil. Die Geologischen Ver-

hältnisse, Adolf Holzhausen, Wien IXI, 1309.

Brinkmann R. 1935: Beiträge zur Kenntnis der alpinen Oberkreide

Nr. 2. Die Ammoniten der Gosau und des Flysch in den nördli-

chen Ostalpen. Mitt. Geol. St.-Inst. Hamburg 15, 114.

Brix F. & Plöchinger B. 1982: Geologische Karte der Republik Ös-

terreich 1:50,000. Blatt 76 Wiener Neustadt. Geol. Bundesan-

stalt, Wien.

Brood K. 1972: Cyclostomatous Bryozoa from the Upper Creta-

ceous and Danian in Scandinavia. Stockholm Contr. Geol. 26,

1464.

Bunzel E. 1871: Die Reptilienfauna der Gosauformation in der Neuen

Welt bei Wr. Neustadt. Abh. K.-Kön. Geol. Reichsanst. 5, 118.

Canu F. & Bassler R.S. 1922: Studies on the Cyclostomatous Bryo-

zoa. Proc. U.S. Nat. Mus. Bull. 61, 22, 1160.

Cjek J. 1851: Die Kohle in der Kreideablagerung bei Grünbach.

Jb. K.-Kön. Geol. Reichsanst. 2, 107123.

Decker K. & Peresson H. 1996: Tertiary kinematics in the Alpine-

Carpathian-Pannonian system: links between thrusting, trans-

form faulting and crustal extension. In: Wessely G. & Liebl W.

(Eds.): Oil and gas in Alpidic thrustbelts and basins of central

and eastern Europe. EAGE Spec. Publ. 5, 6977.

Faupl P. & Wagreich M. 2000: Late Jurassic to Eocene palaeogeog-

raphy and geodynamic evolution of the Eastern Alps. Mitt. Ös-

terr. Geol. Gesell. 92, 7994.

Goldfuss A. 182644: Petrefacta Germaniæ tam ea, quae in Museo

Universitatis Regiae Borussicae Fridericiae Wilhelmiae

Rhenanae servantur quam alia quaecunque in Museis Hoening-

husiano Muensteriano aliisque extant, Iconibus et Descriptioni-

bus illustrata. Arnz & Comp., Düsseldorf, 1(1): iviii+176;

pls. 125 (1826); 1(2): 77164; pls. 2650 (1829); 1(3): 165

240; pls. 5170 (1831); 1(4): 241252 (1833); 2(1): iiii+168;

pls. 7196 (1834); 2(2): 69140; pls. 97121 (1835); 2(3):

141224; pls. 122146 (1837); 2(4): 225312; pls. 147165

(1840); 3(1): iiv+120; pls. 166171 (1841); 3(2): 2128; pls.

172195 (1844); 3(3): 29128; pls. 196200 (1844).

Gordon D.P. 1984: The Marine Fauna of New Zealand:Bryozoa

Gymnolaemata from the Kermadec Ridge. N. Z. Oceanograph-

ic Inst. Mem. 91, 1198.

Gordon D.P. 1986: The marine fauna of New Zealand: Bryozoa

Gymnolaemata (Ctenostomata and Cheilostomata Anasca)

from the Western South Island Continental Shelf and Slope. N.

Z. Oceanographic Inst. Mem. 95, 1121.

Gordon D. & Taylor P.D. 1997: The Cretaceous-Miocene genus Li-

chenopora (Bryozoa) with a description of a new species from

New Zealand. Bull. Nat. Hist. Mus. (Geology) 53, 1, 7178.

Haubold H., Scholger R., Summesberger H. & Mauritsch H.J. 1999:

Reconstruction of geodynamic evolution of the Northern Cal-

careous Alps by means of paleomagnetism. Phys. Chem. Earth

(A) 24, 8, 697703.

Hagenow F. von 1851: Die Bryozoen der Maastrichter Kreidebil-

dung. Theodor Fischer, Cassel, ixv, 1111.

Herman A. & Kvaèek J. 2002: Campanian Grünbach Flora of Lower

Austria: preliminary floristics and palaeoclimatology. Ann.

Naturhist. Mus. Wien 103A, 121.

Hillmer G. 1971: Bryozoen (Cyclostomata) aus dem Unter-Hau-

terive von Nordwestdeutschland. Mitt. Geol. St.-Inst. Hamburg

40, 1106.

Hradecká L., Lobitzer H., Ottner F., vábenická L. & Svoboda M.

1999: Biostratigraphy and facies of selected exposures in the

GrünbachNeue Welt Gosau Group (Coal-bearing Series, In-

oceramus-Marl and Zweiersdorf-Formation, Late Cretaceous

and Paleocene, Lower Austria). Abh. Geol. Bundesanstalt 56,

519551.

Karrer F. 1877: Geologie der Kaiser Franz Josefs Hochquellen-

Wasserleitung. Abh. K.-Kön. Geol. Reichsanst. 9, 1420.

Kollmann H.A. 2002: Theia and Kalais Paleobiogeographic

terms replacing Tethys and Boreal. In: Michalik J. (Ed.):

Tethyan/Boreal Cretaceous Correlation. Publ. House Slovak

Acad. Sci. (VEDA), Bratislava, 285291.

Kowalewski M., Simões M.G., Carrol M. & Rodland D.L. 2002:

Abundant brachiopods on a tropical upwelling-influenced

shelf (Southeast Brazilian Bight, South Atlantic). Palaios 17,

277286.

McKinney F.K. & Jackson J.B.C. 1989: Bryozoan evolution. Unwin

Hyman, Boston 1238.

Novák O. 1877: Beitrag zur Kenntnisse der Bryozoen der böhmis-

chen Kreideformation. Denkschr. Österr. Akad. Wiss. Math.-

Naturwiss. Kl. 2, 37, 150.

dOrbigny A. 1850: Prodrome de Paléontologie stratigraphique uni-

verselle des Animaux Mollusques & Rayonnés faisant suite au

cours élémentaire de Paléontologie et de Géologie strati-

graphiques. Deuxieme Volume. Victor Masson, Paris, 1427.

dOrbigny A. 1852: Prodrome de Paléontologie stratigraphique uni-

verselle des Animaux Mollusques & Rayonnés faisant suite au

cours élémentaire de Paléontologie et de Géologie strati-

graphiques. Troisième Volume. Victor Masson, Paris, 1191.

Pitt L.J. & Taylor P.D. 1990: Cretaceous Bryozoa from the

Faringt(d)on Sponge Gravel (Aptian) of Oxfordshire. Bull.

Brit. Mus. Natur. Hist. (Geology) 46, 1, 61152.

Plöchinger B. 1961: Die Gosaumulde von Grünbach und der Neuen

Welt (Niederösterreich). J. Geol. Bundesanstalt 104, 359441.

Plöchinger B. 1967: Erläuterungen zur Geologischen Karte des

Hohe-Wand-Gebietes (Niederösterreich). Geol. Bundesanstalt,

Wien, 1142

Prantl F. 1929: Contribution a la connaisance du genre Lichenopora

Defr. Vìst. Stát. Geol. Úst. 5, 247257 (in Czech and France).

Prantl F. 1938: Lower Turonian Bryozoa from Pøedboj (Bohemia).

Rozpr. Stát. Geol. Úst. Ès. Republ. 8, 171 (in Czech and En-

glish).

Preisinger A., Zobetz E., Gratz A.J., Lahodynsky R., Becke M.,

Mauritsch H.J., Eder G., Grass F., Rögl F., Stradner H. & Sure-

nian R. 1986: The Cretaceous/Tertiary boundary in the Gosau

Basin, Austria. Nature 322, 6082, 794799.

Reuss A.E. 184546: Die Versteinerungen der böhmischen Kreide-

formation. Schweizerbartsche Verlagsbuchhandlung, Stut-

tgart, Abt. I: iiv+158, pls. 113; Abt. II: iiv+59148, pls.

1451.

Reuss A.E. 1854: Beiträge zur Charakteristik der Kreideschichten in

den Ostalpen, besonders im Gosauthale und am Wolfgangsee.

Denkschr. Österr. Akad. Wiss., Math.-Naturwiss. Kl. 7, 1, 1156.

Reuss A.E. 1872: Die Bryozoen und Foraminiferen des unteren

Pläners. In: Geinitz H.B. (Ed.): Das Elbthalgebirge in Sachsen.

Erster Theil. Palaeontographica 20, 1, 95144.

CRETACEOUS BRYOZOA FROM SCHARRERGRABEN (EASTERN ALPS) 407

Roemer F.A. 1841: Die Versteinerungen des Norddeutschen Kre-

idegebirges. Verlage der Hahnschen Hofbuchhandlung, Han-

nover, 1145.

Sanders D., Kollmann H. & Wagreich M. 1997: Sequence develop-

ment and biotic assemblages on an active continental margin:

the Turonian-Campanian of the Northern Calcareous Alps,

Austria. Bull. Soc. Géol. France 168, 351372.

Seeley H.G. 1881: The reptilian fauna of the Gosau Formation pre-

served in the Geological Museum of the University of Vienna;

with a note on the geological horizon of the fossils at Neue

Welt, west of Wiener Neustadt, by Prof. Eduard Suess. Quart.

J. Geol. Soc. London 37, 620714.

Smith A. 1995: Palaeoenvironmental interpretation using bryozo-

ans: a review. In: Bosence D.W.J. & Allison P.A. (Eds.): Ma-

rine palaeoenvironmental analysis from fossils. Geol. Soc.

Spec. Publ. 83, 231243.

Stur D. 1877: Skizze über die Gosauformation in der Neuen Welt

und deren Umgebung. In: Hauer F. & Neumayr M. (Eds.):

Führer zu den Excursionen der Deutschen Geologischen Ge-

sellschaft nach der Allgemeinen Versammlung in Wien 1877.

Geol. Reichsanstalt., Wien, 154184.

Summesberger H. 1997: Postconference Excursion 2. The Creta-

ceous of the Grünbach Neue Welt Basin. In: Kollmann H.A.

& Hubmann B. (Eds.): Climates: past, present and future. Sec-

ond European Palaeontological Congress. Excursion Guides,

Wien, 7789.

Summesberger H. & Kennedy W.J. 1996: Turonian ammonites from

the Gosau Group (Upper Cretaceous; Northern Calcareous

Alps; Austria) with a revision of Barroisiceras haberfellneri

(Hauer, 1866). Beitr. Paläont. Österr. 21, 105177.

Summesberger H., Wagreich M., Tröger K.-A. & Scholger R. 2000:

Piesting-Formation, Grünbach-Formation und Maiersdorf-For-

mation drei neue lithostratigraphische Termini in der Gosau

Gruppe (Oberkreide) von Grünbach und der Neuen Welt

(Niederösterreich). Ber. Inst. Geol. Paläont. Karl-Franzens-

Univ. Graz 2, 23.

Summesberger H., Wagreich M., Tröger K.-A. & Scholger R. in

print: The Upper Cretaceous of Piesting (Austria): Integrated

stratigraphy of the Piesting Formation (Gosau Group). In:

Wagreich M. (Ed.): Aspects of Cretaceous stratigraphy and

palaeobiogeography. Schriftenreihe der Erdwissenschaftlichen

Kommissionen.

Taylor P.D. & Sequeiros L. 1982: Toarcian bryozoans from Bel-

chite in north-east Spain. Bull. Brit. Mus. Natur. Hist. Geol.

Ser. 36, 117129.

Taylor P.D. 1994: Systematics of the melicerititid cyclostome bryo-

zoans; introduction and the genera Elea, Semielea and Repto-

multelea. Bull. Natur. Hist. Mus. Geol. Ser. 50, 1103.

Taylor P.D. 2002: Bryozoans. In: Smith A.B & Batten D.J. (Eds.):

Fossils of the Chalk. Second edition. Palaeont. Assoc. London

5375.

Voigt E. 1924: Beiträge zur Kenntnis der Bryozoenfauna der sub-

herzynen Kreidemulde. Paläont. Z. 6, 93172, 191242.

Voigt E. 1928: Bryozoen aus dem Gosauvorkommen am Taubensee

bei Kössen in den Nordtiroler Kalkalpen. Cbl. Mineral. Geol.

Paläont., Abt. B 1928, 7, 443448.

Voigt E. 1967: Oberkreide-Bryozoen aus den asiatischen Gebieten

der UdSSR. Mitt. Geol. Staatsinst. Hamburg 36, 595.

Voigt E. 1979: Vorkommen, Geschichte und Stand der Erforschung

der Bryozoen des Kreidesystems in Deutschland und benach-

barten Gebieten. Aspekte der Kreide Europas. IUGS Series A 6,

171210.

Voigt E. 1983: Zur Biogeographie der europäischen Oberkreide-

Bryozoenfauna. Zitteliana 10, 317347.

Voigt E. 1985: Regressive Astogenese bei Nudonychocella n.g.n.sp.

und anderen Bryozoen aus der Tuffkreide von Maastricht.

Paläont. Z. 59, 1/2, 4, 5773.

Voigt E. 1989: Beitrag zur Bryozoen-Fauna des sächsischen Cen-

omaniums. Abh. Staatl. Mus. Mineral. Geol. Dresden 36, 887.

Wagreich M. & Faupl P. 1994: Palaeogeography and geodynamic

evolution of the Gosau Group of the Northern Calcareous Alps

(Late Cretaceous, Eastern Alps, Austria). Palaeogeogr. Palae-

oclimatol. Palaeoecol. 110, 235254.

Wagreich M. & Marschalko R. 1995: Late Cretaceous to Early Ter-

tiary palaeogeography of the Western Carpathians (Slovakia)

and the Eastern Alps (Austria): implications from heavy miner-

al data. Geol. Rdsch. 84, 187199.

Zágorek K. in print: Bryozoa from the Upper Eocene of the

Waschberg Zone (Austria). Beitr. Paläont.

Zágorek K. & Vávra N. 2000: A new method for the extraction of

bryozoans from Hard Rocks from the Eocene of Austria. Jb.

Geol. Bundesanstalt 142, 2, 249258.

Zekeli F. 1852: Die Gasteropoden der Gosaugebilde. Abh. K.-Kön.

Geol. Reichsanst. 1, 2, III, 1124.

Zingula R.P. 1968: A new breakthrough in sample washing. J. Pale-

ont. 42, 4, 1092.

Zittel K.A. 18651966: Die Bivalven der Gosaugebilde in den nor-

döstlichen Alpen. Ein Beitrag zur Charakteristik der Kreidefor-

mation in Österreich. Denkschr. Österr. Akad. Wiss.,

Math.-Naturwiss. Kl. 242: 105178, 252: 77198.