Cretaceous Research 87 (2018) 126e144

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Cretaceous Research
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Comptoniaster michaelisi nom. nov. (Asteroidea, Goniasteridae):

Revision of a starfish species from the lower Upper Cretaceous of
central Europe previously described as Pentagonaster semilunatus and
Asterias schulzii
Birgit Niebuhr a, *, Ekbert Seibertz b
a
Senckenberg Naturhistorische Sammlungen Dresden, Museum für Mineralogie und Geologie, Ko€nigsbrücker Landstraße 159, 01109 Dresden, Germany
b €kologie, TU Braunschweig, Langer Kamp 19c, 38106 Braunschweig, Germany
Institut für Geoo

a r t i c l e i n f o a b s t r a c t

Article history: The goniasterid starfish Comptoniaster michaelisi nom. nov., previously known under the names of As-
Received 9 December 2016 terias schulzii or Stellaster schulzei, is re-examined in terms of nomenclature and taxonomic significance.
Accepted in revised form 16 May 2017 The species was described and illustrated by Schulze in 1760 as Pentagonaster semilunatus, a name that
Available online 18 May 2017
referred to an extant species and did not conform to the rules of binomial nomenclature. The species-
group name schulzii is invalid, so that the replacement name michaelisi is suggested. This early Late
Keywords:
Cretaceous species can now be accommodated in the genus Comptoniaster. A lectotype for the species is
Echinodermata
designated (i.e., Schulze's figured specimen), reillustrated and described in detail herein. Additional data
Taxonomy
Taphonomy
provided include other occurrences of the species in middle Turonian to middle Coniacian strata in the
Germany Saxonian, North Sudetic and Intrasudetic Cretaceous subbasins (Germany, Poland and Czech Republic).
Poland Eighteen moulds of well-preserved, articulated specimens were available for this study, and 14 of them
Czech Republic are illustrated. The skeletal morphology of the species is reconstructed and single elements are
described, characterising C. michaelisi as a medium-sized asteroid with an outer radius of up to 90 mm,
45e50 paired supero- and inferomarginals and an arm-to-disc ratio of 2.2e2.5. Striking morphological
affinities with C. comptoni from the upper Albianelower Cenomanian indicate that species as a possible
ancestor of C. michaelisi. The species lived on shallow-marine, medium- to coarse-grained sandy sea
floors. Taphonomic pathways suggest that both death and rapid burial of these asteroids were most likely
induced by tempestite deposition.
© 2017 Elsevier Ltd. All rights reserved.

1. Introduction
Asteroids rank amongst the diverse and ecologically important
members of marine ecosystems in all present-day oceans, encom-
passing nearly 1900 species, in approximately 370 genera and 36
families (Mah and Blake, 2012). They occur at all depths, from the
intertidal to the abyssal (approximately 6000 m) and in all tem-
perature zones, but are most diverse in tropical regions of the
Atlantic and Indo-Pacific oceans (Hyman, 1955; Clark and Rowe,
1971; Blake, 1983, 1990; Clark and Downey, 1992). All extant star-
fish have been considered to be members of the post-Palaeozoic
Asteroidea (crown-group sensu Blake, 1987, 2000; Neoasteroidea
* Corresponding author.
E-mail address: birgit.niebuhr@senckenberg.de (B. Niebuhr).
sensu Gale, 1987), first occurring in the middle Triassic (Blake and
Hagdorn, 2003). Since their first appearance in Ordovician strata
(Blake, 2007; Blake and Rozhnov, 2007), certain asteroids (stem-
group sensu Blake, 1987; Palaeoasteroidea sensu Gale, 1987; out-
group sensu Gale, 2011) show both similar and intermediate mor-
phologies with the crown-group, and it is these similarities that
have been treated differently by authors (see e.g., Blake, 1987; Gale,
1987; Blake and Hagdorn, 2003).
The classification of the Asteroidea has had a history of con-
troversy since the nineteenth century, having been purely
morphology based and relying on comparison of mostly dissociated
ossicles in extinct taxa with complete extant starfish (see discus-
sions in Gale, 1987; Blake and Portell, 2009). More or less complete
asteroids are unusual finds, even in otherwise fossiliferous strata,
because the skeleton is composed of numerous individual ossicles

http://dx.doi.org/10.1016/j.cretres.2017.05.018
0195-6671/© 2017 Elsevier Ltd. All rights reserved.
 B. Niebuhr, E. Seibertz / Cretaceous Research 87 (2018) 126e144
connected by soft tissues that decay rapidly after death. Many
studies and reconstructions have relied solely on such ossicles (e.g.,
Spencer, 1913; Wright and Wright, 1940; Rasmussen, 1950). Occa-
sionally, assemblages of abundant ossicles of one or more species
are observed but, in general, asteroids are relatively rare fossils and
well-preserved specimens are exceptional. Some examples of the
latter have been known from the thick-bedded sandstones (i.e.,
‘Quader sandstones’) in the Saxonian Cretaceous Basin (SCB) since
the beginning of the eighteenth century (see Niebuhr, 2017). These
have recently been documented in detail (Niebuhr and Seibertz,
2016).
From the Upper Cretaceous (lower Cenomanianemiddle Con-
iacian) of the SCB, twelve asteroid species, in ten genera, are on
record to date (Niebuhr and Seibertz, 2016). Seven of these (Table 1)
are known only from isolated calcitic ossicles from the marly-
calcareous facies in the northwestern part of the SCB, between
Meißen and Dresden. Moulds of near-completely preserved star-
fishes from quartzose Quader sandstones are definitely the most
beautiful fossils of the SCB, and comprise five taxa (Table 1).
However, all of these are rare finds, with the exception of Comp-
toniaster michaelisi nom. nov., of which more than 18 individuals
are now known. The aim of the present paper is to revise this taxon,
previously described and figured as Pentagonaster semilunatus by
Schulze (1760) and Asterias schulzii Cotta by Roemer (1840, 1841),
and to provide a better picture of skeletal morphologies and mode
of life of C. michaelisi nom. nov.
2. Geological setting and stratigraphy
The Mid-European Island is one of the main palaeogeographical
features of the Cretaceous in central Europe, consisting of the
western Rhenish and eastern Bohemian massifs (Rhenobohemia)
as emergent areas (Fig. 1A). Striking roughly WNWeESE, this island
separates the temperate Boreal shelf sea in the north from the
Tethyan warm-water settings in the south (compare Ziegler, 1990;
Janetschke et al., 2015). The Saxonian (SCB), Bohemian (BCB),
North Sudetic (NSCB) and Intrasudetic (ISCB) Cretaceous basins
Table 1
Late Cretaceous (early Cenomanianemiddle Coniacian) asteroids from the Saxonian
Cretaceous Basin in stratigraphic order (Niebuhr and Seibertz, 2016) preserved
either as isolated ossicles in marly-calcareous facies (*) or as moulds of more or less
complete articulated specimens in Quader sandstones.
Lophidiaster scupini (Andert, 1934)
TuronianeConiacian boundary interval
*Chomataster coombii (Forbes, 1848)
mid-upper Turonian
Nymphaster albensis (Geinitz, 1872)
middle Turonianelower Coniacian
Comptoniaster michaelisi nom. nov.
middle Turonianemiddle Coniacian
Calliderma lindneri Niebuhr and Seibertz, 2016
basal middle Turonian
*Manfredaster praebulbiferus Niebuhr and Seibertz, 2016
upper upper Cenomanianebasal Coniacian
*Crateraster quinqueloba (Goldfuss, 1829)
upper upper Cenomanianemid-upper Turonian
*Metopaster parkinsoni (Forbes, 1848)
upper upper Cenomanianemid-upper Turonian
*Metopaster thoracifer (Geinitz, 1871)
upper upper Cenomanian
*Hadranderaster simplex (Geinitz, 1871)
upper upper Cenomanian
*Geinitzaster decoratus (Geinitz, 1871)
upper upper Cenomanian
Calliderma ottoi (Geinitz, 1871)
lower upper Cenomanian
127
were closely connected marginal subbasins of the wide central
European carbonate-dominated shelf sea. In all four subbasins
marine sedimentation commenced during the Cenomanian. The
oldest Cretaceous marine deposits are proximal red conglomerates
of early to middle Cenomanian age (Meißen Formation) that are
confined to the Meißen area, northwest of Dresden (Niebuhr et al.,
2007). During this time interval, the land areas of the future West-
and East-Sudetic islands (see Fig. 1A) were still connected with the
large Mid-European Island in the southwest (compare Voigt, 2009).
The Variscan basement structures of the Mid-European Island
dominated the palaeogeography during the middle to late Cen-
omanian. A major relative sea level rise in the lower Metoicoceras
geslinianum Zone (midelate Cenomanian) led to inundation of
many small palaeohighs and the establishment of fully marine
conditions in all four subbasins (Fig. 1A). Successively, rising sea
level resulted in narrow marine straits at the interface of Boreal and
Tethyan domains between the Mid-European and West-Sudetic
islands (SCB and northwestern BCB), as well as East- and West-
Sudetic islands (NSCB and ISCB). The maximum flooding of the
SCB is indicated by the Strehlen and Weinbo € hla limestones (mid-
eupper Turonian), the carbonate-richest sediment of the Elbtal
Group (see Wilmsen and Niebuhr, 2014). Since the middle Turo-
nian, inversion tectonics governed the depositional regime. The
major clastic source area for the quartzose Quader sandstones of
the SCB (Saxonian Switzerland) and northwestern part of the BCB
(Bohemian Switzerland) was the West-Sudetic Island. The
composition of both sandstones and clasts reveal that sedimentary
rocks of PermotriassiceCretaceous age were eroded (Voigt, 2009).
The ‘Heuschergebirge’ of the ISCB is composed of erosional material
from the East-Sudetic Island in the northeast, while the south-
eastern part of the BCB was filled up from the Bohemian Massif in
the southwest (for a summary of palaeogeographical and strati-
graphical data, see Uli
cný et al., 2008; Voigt, 2009; Janetschke and
Wilmsen, 2014; Wilmsen and Niebuhr, 2014; Janetschke et al.,
2015). Today, the northeastern margins of the SCB and the BCB, as
well as the NSCB, ISCB and the Opole Cretaceous Basin (Fig. 1B), are
characterised by faults; only the southwestern margins of SCB and
BCB show the onlap pattern of the Upper Cretaceous
transgressions.
3. Localities and material
Regional occurrences of Comptoniaster michaelisi nom. nov. in
the four subbasins are shown in Fig. 1B, indicated by stars. It is
obvious from the fossil material that these asteroids are especially
found in shallow-marine, quartzose Quader sandstones of the SCB,
NSCB and ISCB. The reason why C. michaelisi nom. nov. was not yet
found in Quader sandstones of Bohemian Switzerland in the
northwestern part of the BCB and the southeastern part of the BCB
is not known. Apart from a single specimen (MB.E.5313, in a fine-
grained sandstone), all specimens (15 in number) known to date
are moulds in weakly silicified, medium- to coarse-grained
sandstones.
Most of the specimens stem from the SCB (for detailed stratig-
raphy see Niebuhr et al., 2007; Wilmsen and Niebuhr, 2014); all are
historical finds, recovered in the eighteenth and nineteenth cen-
turies during large-scale quarrying of freestones for buildings in the
old town of Dresden and elsewhere. The oldest quarries are located
near Pirna (locality 1 in Fig. 1B) where the Pirnaer Oberquader of
the upper Postelwitz Formation was exploited, of early late Turo-
nian age. The Pirnaer Oberquader, and the time-equivalent Sand-
stone c which crops out in southeastern direction in Wehlen,
Rathen and the Saxonian Switzerland, are weakly silicified, poorly
sorted, medium- to coarse-grained quartzose sandstones, white
and yellowish variegated, with large, isolated detritic quartz grains
128 B. Niebuhr, E. Seibertz / Cretaceous Research 87 (2018) 126e144

Fig. 1. A, Turonianeearly Coniacian palaeogeography with position of subbasins around the Mid-European Island (modified after Voigt, 1994). DCB ¼ Danubian Cretaceous Basin,
SCB ¼ Saxonian Cretaceous Basin, BCB ¼ Bohemian Cretaceous Basin, NSCB ¼ North Sudetic Cretaceous Basin, ISCB ¼ Intrasudetic Cretaceous Basin. B, Uncovered geological map of
the border triangle of Germany, Poland and the Czech Republic with facies distribution during Turonianeearly Coniacian times (simplified after Ulicný et al., 2008; Voigt, 2009,
2015); stars 1e5 mark occurrences of Comptoniaster michaelisi nom. nov. [1] Germany, SCB, Pirnaer Oberquader and Sandstone c of the upper Postelwitz Formation, lower up-
per Turonian. [2] Germany, SCB, Sandstone a of the Postelwitz Formation, middle Turonian. [3] Poland, NSCB, Hockenau Sandstone a of the Rakowice Wielkie Formation (‘Ober-
quadersandstein der Hockenberge’), loweremiddle Coniacian. [4] Czech Republic and Poland, ISCB, Radko w Bluff Sandstone of the Karło
w Formation (‘Heuscheuergebirge’), middle
w Member (‘Kieslingswalder Ton’), loweremiddle Coniacian.
Turonian. [5] Poland, Nysa Graben of the ISCB, Lower Idziko

in yellow, white and pink colours. Exteriors of buildings such as the
old Frauenkirche and the Zwinger were constructed in this free-
stone. The following specimens are known from this lithology
(MMG: SaK 12512 [lectotype], old MMG no. 7431, FG 193/2, FG 193/
2a, MB.E.5316, MB.E.5322 and NHMW 1865/0010/0382). An indi-
vidual from Pirna-Obervogelgesang (see Geinitz, 1849a, pl. 12,
caption of fig. 5) has never been illustrated and is presumed lost.
Locality 2 in Fig. 1B marks Sandstone a of the lower Postelwitz
Formation of middle Turonian age. This slightly more strongly
silicified and matrix-free, medium-grained quartzose sandstone is
well sorted and white to light grey in colour. It is the fossil-richest
Quader sandstone of the SCB. This unit crops out between
Ko€nigstein and Bad Schandau-Schmilka, the latter locality being at
the GermaneCzech border. The largest quarries are those at Post-
elwitz on the right bank of the Elbe River, below the Schrammstein
Massif. Five specimens are known from Sandstone a, namely MMG:
SaK 6804, MMG: SaK 6805, MMG: SaK 6713, LfULG: RS 9878 and
MB.E.5315.
One typical, well-preserved specimen from the NSCB (locality 3
in Fig. 1B) was described and figured by Scupin (1912e1913, p. 255,
text-fig. 47). It is from the loweremiddle Coniacian ‘Oberqua-
dersandstein der Hockenberge’, i.e., the Hockenau Sandstone of the
Rakowice Wielkie Formation at Czaple (Powiat Złotoryjski, Poland;
for detailed stratigraphy see Walaszczyk and Tro € ger, 1996). The
original specimen could not be traced; we do not reproduce Scu-
pin's illustration here because it lacks a scale.
 B. Niebuhr, E. Seibertz / Cretaceous Research 87 (2018) 126e144
Two specimens are known from the Radko w Bluff Sandstone of
the Karłow Formation, a poorly sorted, medium- to coarse-grained
sandstone which an argillaceous matrix, of middle Turonian age
and part of the ‘Heuscheuergebirge’ in the ISCB at the CzechePolish
border (locality 4 in Fig. 1B; for detailed stratigraphy see Ulicný
et al., 2008). A single specimen from Hejsovina (Czech Republic)
is known, also found at a Quader sandstone quarry and now housed
in the collections of the National Museum Prague (NM-O3939).
Another individual was found near Radko  w, Poland (German
‘Wünschelburg’) and is now part of the collections of the Natural
History Museum, London (NHMUK E.13538).
A third specimen (MB.E.5313) of the ISCB stems from Idziko w
(Bystrzyca Kłodzka, Poland) (locality 5 in Fig. 1B), first mentioned
by Geinitz (1872, p. II.16) from the ‘Grünsande in Kieslingswalda’,
which is now the Lower Idziko  w Member of an earlyemiddle
Coniacian date. This unit was deposited in an active tectonic
structure (Nysa Graben); it comprises claystones and clayey silt-
stones, intercalated by sandstone layers (Jerzykiewicz, 1971; Ulicný
et al., 2008; for detailed stratigraphy see; Halamski and Kva cek,
2015). Furthermore, Langenhan and Grundey (1891, p. 8, pl. 1, figs
19e20) described and figured two specimens from the ‘Kieslings-
walder Gestein’ south of Mie˛ dzylesie (German ‘Mittelwalde’), both
of which are Coniacian in age. However, their original specimens
could not be traced and the original illustrations lack a scale.
4. Asteroid taphonomy in Quader sandstones
All complete, articulated starfish fossils of the SCB, NSCB and
ISCB, inclusive of Comptoniaster michaelisi nom. nov. are preserved
as moulds in Quader sandstones. These sandstones imply a habitat
above fair-weather wave base (Wilmsen and Niebuhr, 2014).
Sedimentological analyses of the nearshore, medium- to coarse-
grained Quader sandstones of the SCB that have yielded these
starfish, have implied frequent deposition of storm-induced sedi-
ments (Voigt, 1994, 2011; Wilmsen and Niebuhr, 2014; Wilmsen,
2017). The only way to preserve an articulated, complete asteroid
in such a turbulent environment is rapid burial, e.g., by tempestites
(e.g., Scha€fer, 1962). During storms, great quantities of sand can be
eroded and redeposited in a very short time-span of hours or days
(e.g., Myrow and Southard, 1996). This can be seen in tubular
tempestites and up to 1.5-m-long vertical escape structures of in-
vertebrates in the Quader sandstones of Saxonian Switzerland
(Voigt, 1994; Niebuhr and Wilmsen, 2016). The vigour of current
traction on a living asteroid would first result in damage of the arm
tips; however, in all specimens studied arm tips are retained. Such
impact may be seen in Fig. 6B and in Scupin (1912e1913) specimen
in which arms are current aligned, indicating that a pressure wave
preceded a tempestite.
In living asteroids, the ossicles (of a fine, sponge-like and porose
network of calcium carbonate, named stereom) are only loosely
bound. These ossicles may have had c. 50 per cent of live cells; only
after burial was this first transformed to compact calcitic ossicles.
The sandstones of the SCB, NSCB and ISCB had already completely
stabilised when calcitic ossicles were dissolved. The resultant hol-
low moulds were never filled by sediment, nor were they
deformed. Further compaction of sandstones after dissolution of
ossicles can thus be ruled out.
This kind of preservation as moulds in medium- to coarse-
grained sandstones has both advantages and disadvantages (see
also discussion in Blake, 2000). For instance, advantages include
appreciation of general outline and arm-to-disc ratio of complete
individuals, as well as assessment of the number, length-to-breadth
ratio and size of marginals and configuration of arm tips, actinals
and abactinals. However, a major drawback is the lack of most of
the fine surface details of the former ossicles, such as overall
129
appearance, density and distribution of granules and spine pits,
lateral profile and articulation facets of marginals. Such features are
often used in the taxonomy of extant Asteroidea (Gale, 2011).
Definitions of fossil taxa on the basis of isolated ossicles cannot be
applied to moulds in Quader sandstones.
This becomes clear from Fig. 4C2 and 4C3, which illustrate
plaster casts of the specimen illustrated in Fig. 4C1. The external
surface of marginals might appear to show a dense cover of small,
granular spinelets. Greater enlargement, however, shows only
single quartz grains with irregular open pores between them
(compare also Fig. 11A6eA8). Most of the Quader sandstones with
asteroids are matrix-free and consist exclusively of quartz grains
that are bound together by weak silicification. The viscous latex of
the casts penetrated the irregular open pores and thus contami-
nated the original specimens (all white spots in Fig. 4C1 are latex
remains). This method cannot be applied safety in carbonate- and
clay-free, medium- to coarse-grained quartzose sandstones of the
localities 1, 2 and 5 (see Fig. 1B). Microscopical small spinelets were
not preserved; larger spines of asteroids in Quader sandstones are
known, albeit in silicified preservation only (Niebuhr and Seibertz,
2016, fig. 12b4), or can be seen as well-defined, rounded holes in
contact with the surface of ossicles (Fig. 11A4; Niebuhr and
Seibertz, 2016, figs 12b2, 13b).
In contrast, the Radko  w Bluff Sandstone of the ISCB (locality 4 in
Fig. 1B) has a slightly larger amount of argillaceous matrix. Pores
between single quartz grains are partially filled by bright flakes of
clay (Figs. 4B, 10A1), and for this reason fine structures are better
preserved. However, only few surfaces of marginals of the cast in
Fig. 10 appear to come close to primary preservation of calcareous
ossicle surfaces and could actually retain granule pits of spinelets
(Fig. 10A4).
5. Collections
Most specimens listed here have their own multifarious history,
in particular the lectotype (for details see Niebuhr, 2017). The ma-
terial is contained in the following collections: FG, TU Bergakade-
mie Freiberg, Geoscience Collections (FG 193/2 and FG 193/2a);
LfULG, Sa €chsisches Landesamt für Umwelt, Landwirtschaft und
Geologie, Freiberg, regional collection (RS) (RS 9878, not illustrated
here; see Friebe, 2010; Niebuhr and Seibertz, 2016); MB, Museum
für Naturkunde Berlin, Leibniz-Institut für Evolutions- und Bio-
diversita€tsforschung, echinoderm collections (E) (MB.E.5313,
E.5315, E.5316 and E.5322); MMG, Senckenberg Naturhistorische
Sammlungen Dresden, Museum für Mineralogie und Geologie,
palaeozoological section, Cretaceous of Saxony (SaK) (MMG: Sak
6713, SaK 6804, SaK 6805, SaK 12512 [lectotype] and old MMG no.
7431, http://www.deutschefotothek.de/documents/obj/30114572,
photograph of Fig. 6B was taken in 1927, the original specimen is
lost); NHMUK, Natural History Museum, Department of Palae-
ontology, London, echinoderm collections (E) (NHMUK E.13538);
NHMW, Naturhistorisches Museum Wien (NHMW 1865/0010/
0382); NM, National Museum Prague (NM-03939).
Silicone rubber casts have been made from MMG: Sak 6713 and
NHMUK E.13538 so as to obtain a better insight into true ossicle
dimensions.
6. Systematic palaeontology
Terminology follows Spencer and Wright (1966), Ne raudeau and
Breton (1993) as well as Blake and Hagdorn (2003). Oral and aboral
are used for the lower and upper surfaces of the complete animal.
The radial distance from the disc centre to the arm tip (outer radius)
is abbreviated ‘R’; from the disc centre to the middle of the inter-
brachial arc (inner radius) is abbreviated ‘r’; both abbreviations
130 B. Niebuhr, E. Seibertz / Cretaceous Research 87 (2018) 126e144
being used as a quotient, i.e., the arm-to-disc ratio. The disc is
bordered by the concave interbrachial arc; proximal is towards the
disc centre, and distal is away from it. At the edge of the body is a
double row of marginal ossicles, the superomarginals above and
inferomarginals below. The height of marginals is vertical, the
length is parallel to the outline of the asteroid, and the breadth
perpendicular to this. Abactinals are platelets of the aboral body
side, actinals lie between the marginals and the ambulacral column
on the oral side. The ambulacral skeleton includes adambulacrals,
which line the margins of the ambulacral furrows, and ambulacrals,
which rest on the adambulacrals on their inner face. Mouth angle
ossicles are the comparatively large ossicles at the end of the
adambulacral column bordering the peristome (mouth), whereas
first ambulacrals head the ambulacral column and overlie and abut
the mouth angle ossicles. The first ambulacrals are thickened, but
superficially are ambulacral like, whereas the mouth angle ossicles
are adambulacral like. The odontophore is an unpaired, typically T-
shaped ossicle between and aboral to the distal side of each mouth
angle pair. The madreporite is a sieve plate on the aboral side
within the interbrachial arc, connecting the water vascular system
to the environment.
6.1. Classification
Class Asteroidea de Blainville, 1830
Clade Neoasteroidea Gale, 1987
Order Valvatida Perrier, 1884
Family Goniasteridae Forbes, 1841
Comptoniaster-Tylasteria Group sensu Breton, 1992
Genus Comptoniaster Breton, 1983
Type species: Goniaster (Stellaster) comptoni Forbes, 1848; by orig-
inal designation.
Remarks. Pentagonal to narrowly stellate forms generally with large
disc. Arms are long and tapering, with arm-to-disc ratios of 2e3
(Breton, 1992). Short, wide marginals, massive, well differentiated
and opposite. Separation of marginals along the dorsal midlines of
the arms is typical. Madreporite is in a central position. Ped-
icellariae and fascioles may be developed.
Comptoniaster michaelisi nom. nov.
(¼ Pentagonaster semilunatus sensu Schulze, 1760 [non Linck, 1733];
Asterias schulzii Cotta sensu Roemer 1840, 1841; and Stellaster
schulzei Cotta and Reich sensu Geinitz, 1872)
Figs. 2e11
1730 fünfeckigter See-Stern; Keyßler, p. 1306.
non 1733 Pentagonaster semilunatus. Das gestirnte Fünff-Eck mit
ausgerundeten Seiten; Linck in Linck et al., p. 21, pl. 23,
fig. 37; pl. 24, fig. 39; pl. 27, fig. 45.
1749 Seestern; Helf, p. 535.
1755 fünfeckigter Seestern; Eilenburg, p. 25.
1760 Pentagonaster semilunatus; Schulze, pp. 50, 54, pl. 2,
fig. 6.
1762 Stellae marinae, versteinerte Seesterne; Walch, p. 76
(pars), pl. 2, fig. 1d [non pl. 2, fig. 1aec].
1783 fünfeckigter Seestern; Hasche, p. 297.
1840 Asterias schulzii Roemer, pl. 6, fig. 21.
1841 Asterias schulzii Cotta; Roemer, pp. 28, 137.
1842 Asterias schulzii Reich; Geinitz, pp. xix, 89, 111.
1843 Asterias schulzii C.; Bronn, p. 87.
1844 Asterias schulzii Reich.; Reuss, p. 16.
1846 Asterias schulzii Cotta; Geinitz, p. 536, pl. 23, fig. 16.
1848 ‘Asterias schulzii’ [in Goniaster (Stellaster) elegans Gray];
Forbes, p. 476.
1849a Asterias schulzii Cotta & Reich, Asterias Schulzi Cotta;
Geinitz, p. 228, pl. 12, fig. 5.
1849b Asterias schulzii; Geinitz, p. 295.
1850 Asterias Schultzii of Roemer [in Stellaster Comptoni
Gray]; Forbes, p. 336.
1850 Pentetagonaster Schulzii d'Orb., 1847; d'Orbigny, p. 180,
no. 677.
1857 Asterias Schultzii Cotta [¼ Stellaster? Schultzii]; Pictet,
p. 269.
1858 Asterias schulzii Cotta und Reich; von Gutbier, p. 8,
text-fig. 3.
1861 Asterias schulzii Cotta und Reich; von Cotta, p. 191,
unnumbered figure.
1862 Stellaster schultzii; Dujardin and Hupe , p. 408.
non 1863 Asterias schulzii Cotta; Drescher, p. 359, pl. 8, fig. 5 [only
32 marginals per side].
1872 Stellaster schulzei Cotta & Reich; Geinitz, p. II.15, pl. II.5,
figs 3e4.
non 1876a Asterias schulzii; Quenstedt, p. 61 [¼ Comptoniaster
comptoni (Forbes, 1848)].
non 1876b Asterias schulzii; Quenstedt, pl. 92, fig. 14, 14m, s, x, z [¼
Comptoniaster comptoni (Forbes, 1848)].
1891 Asterias schulzii Cotta; Langenhan and Grundey, p. 8, pl.
1, figs 19e20.
?1897 Asterias Schulzei Cotta et Reich, Asterias schulzii Cotta;
Fri
c, pp. 34, 72.
1907 Asterias schulzii Cotta; Spencer, p. 109.
1912e1913 Stellaster schulzei Cotta; Scupin, p. 255 (pars), text-fig.
47 [non Drescher (1863)].
1914 Stellaster schultzii Pictet ¼ Asterias schultzii colta
Roemer ¼ Pentegonaster schultzii d'Orb.; Goto, p. 685.
1927 Stellaster schulzei Cotta und Reichenb.; http://www.
deutschefotothek.de/documents/obj/30114572
1934 Stellaster schulzei Cotta & Reich sp.; Andert, p. 71 (pars).
1939 Stellaster schulzei Cotta & Reichenbach; Fischer,
comment on p. 522, pl. 22 below, right side.
non 1939 Stellaster schulzei (Cotta et Reich); Soukup, p. 1, text-fig.
1 [¼ Nymphaster albensis (Geinitz, 1872)].
1959 Stellaster schulzei Cotta et Reich; Prescher, pl. 2, fig. 4.
1979 Stellaster schulzei; Prescher, p. 52, text-fig. 1.
2010 Stellaster sp.; Friebe, p. 8, lower figure.
2016 Comptoniaster michaelisi nom. nov.; Niebuhr and
Seibertz, p. 130, text-figs 10e11 [rationale not dis-
cussed in detail].
Derivatio nominis. In honour of the Dresden court art chamberlain
Johannes Gottlieb Michaelis (1704e1740), who collected the spec-
imen that was later described and figured by Schulze (1760).
Type. Lectotype is MMG: SaK 12512, Senckenberg Naturhistorische
Sammlungen Dresden, Museum für Mineralogie und Geologie
(MMG), palaeozoological section (see Fig. 2B1). The specimen,
discovered in 1727 by J.G. Michaelis, was on exhibit at the former
Ko€ nigliches Naturalien-Cabinet of the Dresden Zwinger between
1728 and 1849. After the fire that broke out during the revolution in
May 1849, the specimen was thought to have been lost (Geinitz,
1849b). During the reconstruction of the Zwinger it was salvaged
from the rubble after the fire and inventoried by H.B. Geinitz in
1854. However, Geinitz never noted that the original had been
retrieved. In the summer of 2016, during the first revision of
Cretaceous starfishes from Saxony since Geinitz's days (1872),
Schulze's original (1760) was rediscovered in the MMG collections.
It is one of the oldest palaeozoological objects and the only one to
left from the Michaelis Collection of 1727 (Niebuhr, 2017).
Stratum typicum. Pirnaer Oberquader of the Postelwitz Formation;
lower upper Turonian (see Wilmsen and Niebuhr, 2014). The
 B. Niebuhr, E. Seibertz / Cretaceous Research 87 (2018) 126e144 131

Fig. 2. Comptoniaster michaelisi nom. nov. A, Reproduction of the first illustration of a fossil starfish from the Cretaceous of Saxony (Schulze, 1760, pl. 2, fig. 6). B1, MMG: SaK 12512,
lectotype; inner view of oral side on upper bedding plane, surrounded by imprints of a continuous row of inferomarginals and partially discontinuous row of superomarginals, the
latter preserved in two interbrachial arcs, one central bulb of faeces is visible; note inner views of sunken abactinals; lower upper Turonian, locality 1 in Fig. 1B, Pirna-Posta (original
of Schulze, 1760, pl. 2, fig. 6, and of Walch 1762, pl. 2, fig. 1d). B2, Close-up of the upper ambulacral furrow with moulds of ambulacral ossicles; width of photograph 8 mm. B3, Close-
up of the lower right arm, showing a healed predation scar, with imprints of adambulacral ossicles; width of photograph 50 mm.

original colour of the sandstone is white and yellowish variegated,
as noted by Schulze (1760, p. 54). The lectotype is now of a
brownish hue as a result of fire damage (see Fig. 2B1).
Locus typicus. Quader sandstone quarries at Pirna-Posta on the right
banks of the Elbe River, Saxony, Germany.
6.2. Description
Of the eighteen specimens of Comptoniaster michaelisi nom. nov.
known to us, fifteen present the aboral side, namely MMG: SaK
6805, MMG: Sak 6713, old MMG no. 7431, FG 193/2, FG 193/2a,
132 B. Niebuhr, E. Seibertz / Cretaceous Research 87 (2018) 126e144

Fig. 3. Reconstruction of Comptoniaster michaelisi nom. nov., presenting a compilation of skeletal elements observed in the specimens illustrated here; dimensions are taken from
specimen MMG: SaK 6713 (see Fig. 4C); SM ¼ superomarginals, IM ¼ inferomarginals, AP ¼ abactinal plates, MAO ¼ mouth angle ossicles, Amb ¼ ambulacrals,
Adamb ¼ adambulacrals, Amb F ¼ ambulacral furrow. A, Oral side with inferomarginals, adambulacrals and mouth angle ossicles. B, Aboral side with superomarginals, abactinals
and madreporite. C, Proximal position of a transverse view of an arm showing stout marginals, flat, abutting abactinals and upright ambulacrals with a narrow furrow (compare
Blake, 1987, fig. 2f); stippled area ¼ sandy bottom.

MB.E.5313, MB.E.5316, MB.E.5322, NHMW 1865/0010/0382, NM-
03939, NHMUK E.13538 (see Figs. 4, 6, 7BeC, 8e11), LfULG: RS
9878, Langenhan and Grundey (1891, pl. 1, figs 19e20) and Scupin
(1912e1913, text-fig. 47). Three others, inclusive of the lectotype,
show the oral side; MMG: SaK 12512, MMG: SaK 6804 and
MB.E.5315 (see Figs. 2, 5, 7A; Table 2).
General outline. The oral side is flat (Fig. 5), the aboral side slightly
convex, pillow like (Figs. 4A, 7B). The centre of the aboral side is
elevated as a result of lesser compaction of the body above the oral
plates (Figs. 6A2, 10A5). The five moderately long arms are trian-
gular with straight sides distally, but broadening proximally into
rounded interbrachial arcs; arm tips are pointed (Fig. 3). Arm-to-
disc ratios vary between 2.2 and 2.5 (Table 2). The largest known
specimen has an outer radius (R) of 89 mm (NHMUK E.13538; see
Fig. 10). Nearly all specimens that show the aboral sides have some
arm tips bent downwards into the underlying sediment (best seen
in Figs. 4B1, 6A1, 6A4eA6, 6B, 8A1e2).
Ornament. Outer surfaces of all visible ossicles appear similar as a
dense cover of small granular pits of spinelets. Greater enlarge-
ment, however, shows in fact single quartz grains with irregular
open pores between them (compare Fig. 11A6eA8), which cannot
be linked to dissolved spines. Exceptionally, in the cast of NHMUK E.
13538, outer surfaces of a few marginals bear a cover of regularly
arranged, medium-sized granule pits which may have borne
spinelets (Fig. 10A4).
Marginal plates. In all specimens, ossicles are dissolved. The rows of
supero- and inferomarginals are opposed, and are arranged closely
paired (see Figs. 4A, 8e11). The number of marginals varies be-
tween 46 and 50 per side; most of the figured specimens in which
the rows between arm tips are complete have 48. Central marginals
of the interbrachial arc and proximal marginals of the arms have
length-to-breath ratios of less than 0.5 (2.3 : 5 mm in the largest
specimen, Fig. 10), and can be slightly trapeziform in outline
(Figs. 4C1, 10A2); the height can be estimated in artificial at c. 4 mm
(Figs. 4C2e3, 10A2eA4). Distal marginals in the arms are also
rectangular in outline with a rounded outer edge and near-parallel
sides (Figs. 4C2, 10A3), becoming abruptly shorter distally in some
specimens between ossicles 6e8 (counted from the arm tip) that
coincides with the area of downward bend (compare Fig. 3). At the
pointed arm tips of both oral and aboral sides, there are pairs of
small, elongated, triangular end plates (Figs. 2B3, 5A3, 6A4e6,
11A2eA4) which bear some prominent spines (Fig. 11A4). The
outer surface of marginals is vaulted and rimless; examples of
 B. Niebuhr, E. Seibertz / Cretaceous Research 87 (2018) 126e144 133

Fig. 4. Top views of Comptoniaster michaelisi nom. nov. on upper bedding planes. A1, MMG: SaK 6805; pillow-like, convex-up aboral side with madreporite in left interbrachial arc,
surrounded by imprints of closely approximated supero- and inferomarginals; middle Turonian, locality 2 in Fig. 1B, Saxonian Switzerland (original of Asterias schulzii Cotta of
Geinitz, 1846, pl. 23, fig. 16, and of Geinitz, 1849a, pl. 12, fig. 5). A2, Close-up of centre of disc with madreporite in lower interbrachial arc; width of photograph 15 mm. A3, Close-up
of outer continuous row of inferomarginals partially overlying sunken superomarginals or precisely opposing in inner interbrachial arcs; width of photograph 6 mm. B1, NM-O3939;
aboral side surrounded by imprints of continuous row of inferomarginals with madreporite in lower right interbrachial arc; middle Turonian, locality 4 in Fig. 1B, Hejsovina, Czech
Republic; photograph by Jan Sklena r, National Museum Prague. B2, Close-up of inner views of regularly arranged, tessellate abactinals; width of photograph 14 mm. C1, MMG: SaK
6713; aboral side surrounded by imprints of continuous row of inferomarginals; middle Turonian, locality 2 in Fig. 1B, Rietzschgrund near Ko €nigstein-Gohrisch (original of Stellaster
schulzei Cotta & Reich of Geinitz, 1872, pl. II.5, fig. 3). C2, Artificial cast of abactinals and inferomarginals of central upper arm; width of photograph 37 mm. C3, Artificial cast of outer
surfaces of ten inferomarginals of interbrachial arc; width of photograph 14 mm.
134 B. Niebuhr, E. Seibertz / Cretaceous Research 87 (2018) 126e144

Fig. 5. Comptoniaster michaelisi nom. nov. A1, MMG: SaK 6804; inner view of oral side on upper bedding plane, surrounded by imprints of continuous row of inferomarginals with
two lumps of faeces in left interbrachial arc; stratigraphically oldest specimen known, lower middle Turonian, locality 2 in Fig. 1B, Grenzbruch near Bad Schandau-Schmilka. A2,
Close-up of wide row of adambulacrals from upper left arm; width of photograph 37 mm. A3, Close-up of upper left arm tip showing moulds of two small end plates; width of
photograph 15 mm. A4, Close-up of outer surface of inferomaginals from upper left interbrachial area showing weak granulation; width of photograph 25 mm.

superomarginals are seen in Fig. 11A5eA7, of inferomaginals in
Figs. 2B, 5A4 and 7A1. Articular facets of marginals are not known,
due to preservation. Interspaces between dissolved marginals, fil-
led with quartz grains, are less than 1 mm wide. Whether they
represent pedicellarian openings or fasciolar grooves (e.g., Gale,
1987, p. 123) cannot be ascertained and is rather unlikely, because
all asteroids (c. 30 in total) of different taxa found in the SCB show
the same feature (Niebuhr and Seibertz, 2016).
Actinal and abactinal ossicles. Polygonal abactinal ossicles are very
small, have a hexagonal form, occasionally differentiated to
rhombical, pentagonal or heptagonal, and show a more or less
regular tessellate arrangement (Figs. 4B2, 8A4, 9, 10A1, 11A3,
11A5e6, 11A8). Impressions clearly show that outer surfaces were
flat (Fig. 11A6, A8) and ossicles tapered inwardly (Figs. 4B2, 8A4),
not showing any ornament such as granulation, spinelets or pax-
illae. In the same specimen abactinals can reach either into the arm
tips or end a few marginals before, counted from the end of the
arm. As far as actinal plates are concerned, nothing can be said.
Such appear in bottom-up views; however, all oral sides (Figs. 2B1,
5A1, 7A1) are preserved on upper bedding planes of the former
substrate and are visible in inner view (compare Fig. 12A1).
Ambulacral skeleton. Externally, the ambulacral system consists of
two rows of adambulacral ossicles (Figs. 2B2, 5A1e2, 7eA1),
opposed and with a narrow furrow between them (compare Fig. 3A)
and abutting the inferomarginals at the beginning of the arm be-
tween the 14th and 18th inferomaginals counted from the arm tip.
They have nearly the same dimensions as the central and proximal
marginals (length-to-breadth ratio c. 2 : 4 mm); the height cannot be
measured. From Fig. 2B2 it is clear that the adambulacrals possessed
scarcely scattered spinelets. Top views from aboral sides into the
arms (Figs. 6A3, 8A1, 10A1) show tops of ambulacrals, standing on
the adambulacrals in the same arrangement (compare Fig. 3C).
Oral plates. The ten paired mouth angle plates at the proximal end
of the adambulacrals are comparatively large (Fig. 7A1e2; compare
Blake, 1987, fig. 3b). They present a fusion of the last proximal
adambulacral ossicle and a triangular oral blade of the circumoral
mouth frame (see discussion in Blake and Hagdorn, 2003; Blake,
2007). They cover the mouth and thus protect the inner oral ossi-
cles such as first ambulacrals and odontophores (compare Blake,
1983, 1987; Gale, 2011, pl. 12, fig. 1). The first ambulacrals and un-
paired odontophores are best seen in Fig. 6A1 and A2, which pre-
sent a denuded aboral top view, rendering a view of the
arrangement of the oral ossicles. Another aboral top view (Fig. 10A1,
A5) shows the extruded ossicles; five strong knobs facing the
ambulacral furrows are of the first ambulacrals, and five weaker
knobs facing the interbrachial arcs are of the odontophores.
 B. Niebuhr, E. Seibertz / Cretaceous Research 87 (2018) 126e144 135

Fig. 6. Top views of Comptoniaster michaelisi nom. nov. on upper bedding planes. A1, NHMW 1865/0010/0382; aboral side surrounded by imprints of continuous row of infer-
omarginals with madreporite in upper interbrachial arc; arms above ambulacral furrows sunken and slightly corroded, as is apical area; lower upper Turonian, locality 1 in Fig. 1B,
Bad Schandau; photograph by Mathias Harzhauser, Naturhistorisches Museum Wien. A2, Close-up of inner view of odontophores, first ambulacrals and madreporite (lower right
corner); height of photograph 20 mm. A3, Close-up of inner view of corroded inner ambulacral plates of upper left arm; height of photograph 13 mm. A4eA6, Close-ups showing
downward bend of three arm tips; in A6, outer discontinuous row of superomarginals visible; height of photographs 20 mm. B, Old MMG no. 7431 (http://www.deutschefotothek.
de/documents/obj/30114572, original specimen lost); aboral side surrounded by imprints of continuous row of inferomarginals with madreporite in upper interbrachial arc; arms
above ambulacral furrows sunken; three lumps of faeces visible; arms current aligned from lower left to upper right; lower upper Turonian, locality 1 in Fig. 1B, Stadt Wehlen.

Madreporite. Thirteen specimens present aboral sides; of these,
eight retain the madreporite (Figs. 4AeB, 6AeB, 8e10). The sieve
plate is located approximately in the centre of the body and the
edge of the interbrachial arc, in most of the cases positioned slightly
nearer the centre which is typical of valvatidans (Blake, 1987). The
outline of the madreporite is well rounded, rectangular and convex,
the diameter varies between 6 and 8 mm. The internal side consists
of imprints of the inner face of the madreporite and thus shows the
typical convoluted structure differentiated by an elongated X
(Fig. 8A4).
Accessory ossicles. Pedicellariae, fascioles, spinelets and granules
that occur in abundance in most goniasterid asteroids are not
observed in Comptoniaster michaelisi nom. nov. This is undoubtedly
preservation induced.
6.3. Nomenclature and taxonomic assignment
The species here revised was first described and figured by
Christian Friedrich Schulze (1760, pp. 50, 54, pl. 2, fig. 6). Schulze's
specimen clearly represents a Cretaceous starfish from Saxony, and
this specimen survives (Fig. 2B); it is here designated lectotype. The
specimen was discovered in 1727 by Johannes Gottlieb Michaelis
and provided to the former Ko €nigliches Naturalien-Cabinet of the
Dresden Zwinger, where it remained between 1728 and 1849. The
specimen originated from the Pirnaer Oberquader of the Postelwitz
Formation (lower upper Turonian).
Schulze (1760) described the specimen under the name of
Pentagonaster semilunatus. Despite the fact that he appeared to
have recognised it as a new ‘form’, he applied a name used by Linck
(in Linck et al., 1733, p. 13) in a description of an extant starfish with
similarities in outline (half-moon shaped), but with only 12 mar-
ginals. Thus the name applied by Schulze (1760), without any
consideration of the greater number of marginals (>46, visible in
his figure of the specimen; Fig. 2A), is of no relevance for the species
studied. Moreover, it does not conform to the general rules of
binomial nomenclature based on the basic publication of Carl von
Linne (Linnaeus, 1735) according to article 11.4 of the ICZN (Ride
et al., 1999).
136 B. Niebuhr, E. Seibertz / Cretaceous Research 87 (2018) 126e144

Fig. 7. Comptoniaster michaelisi nom. nov.; von Cotta Collection, potential originals of Asterias schulzii Cotta of Roemer (1840, pl. 6, fig. 21). A1, MB.E.5316; inner view of oral side on
upper bedding plane, surrounded by imprints of continuous row of inferomarginals with three small lumps of faeces in lower, lower left and upper left interbrachial arc; lower
upper Turonian, locality 1 in Fig. 1B, ‘Oberer Quader, Sachsen’ sensu von Cotta. A2, Close-up of central area showing strong mouth angle ossicles; width of photograph 22 mm. B,
MB.E.5315; top view of pillow-like, convex-up aboral side on upper bedding plane surrounded by imprints of poorly preserved marginals with madreporite in upper interbrachial
arc, with arms above ambulacral furrows sunken, and two lumps of faeces in centre and right interbrachial arc; middle Turonian, locality 2 in Fig. 1B, ‘Obere Kreideformation,
Ko€ nigreich Sachsen’ sensu von Cotta. C, MB.E.5322; top view of poorly preserved aboral side on upper bedding plane, surrounded by imprints of marginals; lower upper Turonian,
€chsische Schweiz’ sensu von Cotta.
locality 1 in Fig. 1B, ‘Oberer Quader, Sa
 B. Niebuhr, E. Seibertz / Cretaceous Research 87 (2018) 126e144 137

Fig. 8. Top view of Comptoniaster michaelisi nom. nov. on upper bedding plane. A1, FG 193/2; aboral side surrounded by imprints of inner continuous row of inferomarginals and
partially preserved outer discontinuous row of superomarginals, with madreporite in lower right interbrachial arc and regularly arranged, tessellate abactinals; arms above
ambulacral furrows partially sunken; lower upper Turonian, locality 1 in Fig. 1B, Gans quarry in Stadt Wehlen (original of Stellaster schulzei Cotta & Reich of Geinitz, 1872, pl. II.5, fig.
4); photograph by Birgit Gaitzsch, TU Bergakademie Freiberg. A2, Close-up of downward bend of arm tip; two separate voids after paired end plates and both rows of marginals;
width of photograph 19 mm. A3, Close-up with both imprints of inner row of inferomarginals (below) and outer row of superomarginals (above) due to near-horizontal pressure;
width of photograph 19 mm. A4, Close-up showing inner surfaces of regularly arranged, tessellate abactinals and madreporite with convoluted suture in internal side, differentiated
by an elongated X; width of photograph 20 mm.

Eighty years later, Friedrich Adolf Roemer obtained at least one
Cretaceous asteroid from the collection of Carl Bernhard von Cotta
(probably MB.E.5315, MB.E.5316 and/or MB.E.5322; see Fig. 7),
which von Cotta had collected during mapping activities in the
Saxonian Switzerland that started in 1835. Roemer first illustrated
this specimen (Roemer, 1840, pl. 6, fig. 21) and later briefly
described it (Roemer, 1841, p. 28) as ‘Asterias schulzii Cotta’. The
specimen was noted to have come from the ‘Quader bei Tharand’,
southwest of Dresden, which is now recognised as the Unterquader
of the Oberha €slich Formation (lower upper Cenomanian; SCB; see
Wilmsen and Niebuhr, 2014). Roemer's illustration is a highly
schematised drawing of the aboral side which cannot be identified
with any of the specimens at hand. The original labels of von Cotta
for the specimens now housed in the Museum für Naturkunde
(Berlin) indicate that Roemer's (1841) assignment to the upper
Cenomanian locality ‘Tharand’ is incorrect. All specimens of
Comptoniaster michaelisi nom. nov. known to date are not older
than middle Turonian, and all specimens from Saxony (including
those collected by von Cotta) originate from the quartzose Quader
sandstones of Saxonian Switzerland southeast of Dresden. Geinitz
(1872, p. 15) already noted that Roemer's putative findings of the
species at Tharandt was erroneous and that the species ‘is an index
fossil of the upper Quader sandstone of the Saxonian Switzerland’.
Roemer's 1840 and 1841 papers provide a name and an illus-
tration for the species that has been used by subsequent authors.
However, the name Asterias schulzii (subsequent ones following
taxonomic revisions, such as Pentetagonaster schulzii and Stellaster
schulzii), plus corrected names such as Asterias schulzei and
Stellaster schulzei must be regarded invalid for the following rea-
sons. Roemer (1840, 1841) referred to the species as ‘Asterias schulzii
Cotta’, which clearly indicates that Roemer did not intend to
introduce this species name himself, but ascribed it to von Cotta. In
fact, only Forbes (1848, 1850) cited Roemer as the author of the
species. Von Cotta did not publish any paper in which the species
name is introduced prior to Roemer's articles.
Geinitz (1842) listed the species as ‘Asterias schulzii Reich’,
which suggests that he regarded a note by Ferdinand Reich as the
source of the name, but later mentioned and figured the same as
‘Asterias schulzii Cotta & Reich’ (Geinitz, 1849b). Von Cotta himself
illustrated the species and referred to it as ‘Asterias schulzii Cotta
und Reich’ (von Cotta, 1861, unnumbered figure on p. 191; reillus-
tration of von Gutbier, 1858, text-fig. 3), thereby contradicting
Roemer's suggested authorship and referring to a joint study with
Reich. Such a study, however, was never published in printed form,
and it is assumed that von Cotta in fact referred to a list of fossils, for
internal use in the collections of the Akademisches Mineralien-
138 B. Niebuhr, E. Seibertz / Cretaceous Research 87 (2018) 126e144

Fig. 9. Top view of Comptoniaster michaelisi nom. nov. on upper bedding plane, FG 193/2a; aboral side with madreporite in lower right interbrachial arc, surrounded by imprints of
inner continuous row of inferomarginals and outer discontinuous row of superomarginals; end of lower arm curved left, indicating pressure direction from lower left to upper right;
lower upper Turonian, locality 1 in Fig. 1B, Gans quarry in Stadt Wehlen; photograph by Birgit Gaitzsch, TU Bergakademie Freiberg.

Cabinett in Freiberg for which he and Reich were responsible. Reich
reidentified all fossils in the Freiberg collections, starting in 1822,
thereby applying binomial nomenclature. Subsequently, von Cotta
organised a comprehensive catalogue, and the name Asterias
schulzii obviously originated from a provisional version of this
catalogue. Both von Cotta and Reich announced the publication of a
monograph on Saxonian Cretaceous fossils (Prescher, 1979), which
would have included the asteroids. However, this monograph was
never published. This, in turn, indicates that the name ‘Asterias
schulzii’ is a nomen invalidum because it does not fulfil the criteria of
article 8.1 of the ICZN (Ride et al., 1999).
At times, the authorship was assigned to Heinrich Gottlieb
Ludwig Reichenbach (e.g., Deutsche Fotothek, 1927; Fischer, 1939),
which possibly resulted from an unintended period following the
author's name ‘Reich’ in some of the early publications (e.g., Reuss,
1844).
The names ‘Asterias Schultzii’ in Forbes (1850) and Pictet (1857)
and ‘Stellaster schultzii’ in Goto (1914) must be regarded as typo-
graphical errors and are thus nomina nulla. The species-group name
‘schulzii’ certainly refers to the material figured and described by
Schulze (1760), so that the correct spelling would have been
‘schulzei’. The first (legitimate) correction of the species-group
name is that by Geinitz (1872), but this act has no further
nomenclatural relevance.
In view of the fact that Schulze's (1760) name for the species is
not available and Roemer's assignment is invalid, a replacement
name is called for. We here propose the specific epithet michaelisi.
This name has already been used in two recent papers (Niebuhr and
Seibertz, 2016; Niebuhr, 2017), but the rationale was not discussed
in detail.
Earlier there was no type for this species. Roemer's crude figure
(1840, pl. 6, fig. 21) is insufficient to recognise any particular
specimen in the present lot, but the illustration provided by Schulze
(1760, pl. 2, fig. 6) does show details of an arm with healed pred-
atory injuries in the lower right corner of the figure to permit
recognition of MMG: SaK 12512 (compare Fig. 2A and 2B). For that
reason, this informative specimen is here designated lectotype of
the species.
6.4. Morphological affinities
Comptoniaster michaelisi nom. nov. exhibits close affinity with
the type species of the genus Comptoniaster, C. comptoni. Forbes
(1848, p. 476) first considered ‘Asterias Schultzii’ to be a synonym
of Goniaster (Stellaster) elegans Gray [¼ Comptonia elegans Gray,
1840, p. 278]. Later, Forbes (1850, p. 336) and Spencer (1913, p.
131) placed the species in synonymy with Stellaster comptoni Gray/
Comptonia comptoni Forbes, without providing a differential
 B. Niebuhr, E. Seibertz / Cretaceous Research 87 (2018) 126e144 139

Fig. 10. Top view of Comptoniaster michaelisi nom. nov. on upper bedding plane. A1, NHMUK E.13538; aboral side surrounded by imprints of inner continuous row of infer-
omarginals and outer discontinuous row of superomarginals in all five interbrachial arcs, with well-preserved inner surfaces of polygonal abactinals and madreporite in upper right
interbrachial area; largest specimen known (R ¼ 89 mm); middle Turonian, locality 4 in Fig. 1B, near Radkow, Poland; photograph by Loïc Villier, Pierre et Marie Curie Universite
,
Paris. A2, Artificial cast of the lower right arm and marginals of upper interbrachial arc; width of photograph 12 mm. A3, Artificial cast showing normal preservation of outer
surfaces of marginals; height of photograph 7 mm. A4, Artificial cast showing best preservation of granulation on outer surfaces of few marginals; height of photograph 7 mm. A5,
Close-up showing extruded oral ossicles, five strong knobs facing ambulacral furrows representing first ambulacrals, and five weaker knobs facing interbrachial arcs representing
odontophores; width of photograph 28 mm.

diagnosis. Both species were integrated by Breton (1983) in the new
genus Comptoniaster. Diagnostic characters of Comptoniaster are
regarded here to include (1) the pentagonal to narrowly stellate
general outline, with a large disc; (2) an arm-to-disc ratio of 2e3;
(3) short, wide marginals; massive, well differentiated and oppo-
site; (4) separation of marginals along the dorsal midlines of the
arms, as well as (5) the central position of the madreporite. All of
these can be seen in C. michaelisi nom. nov.
Comptoniaster comptoni is a rare species that occurs in upper
Albianelower Cenomanian strata of England (Forbes, 1848, 1850;
Spencer, 1905, 1907, 1913; Gale, 1988) and France (Breton, 1992).
Only two individuals are near complete; see Spencer (1905, pl. 17,
fig. 3 ¼ NHMUK E.34311, oral side up; pl. 18, fig. 2, Northampton
Museum, aboral side up). These specimens are closely similar to C.
michaelisi nom. nov., especially in general outline, size, number of
marginals per side, arm-to-disc ratio, arrangement of abactinals as
well as pointed arm tips with paired small, elongated, triangular
end plates (see upper arm tip in Spencer, 1905, pl. 18, fig. 2, in
comparison with Figs. 6A4, 11A2e4 here).
However, Comptoniaster michaelisi nom. nov. has slightly wider
adambulacrals and thus wider arms, which are triangular and not
elongated; in addition, it appears to lack the characteristic ped-
icellarian pits in the marginals, frequently seen in C. comptoni.
Forbes (1848, p. 476; 1850, p. 336) also realised this, noting,
‘Asterias Schultzii… comes so very near Goniaster elegans, that were
it not for the absence of all traces of pedicellarian sulcations on the
marginal plates represented in his figure, I should consider it
identical’ and ‘ … appears to be this (Stellaster Comptoni) species’,
respectively. All known Late Cretaceous species of Comptoniaster
are characterised by the presence of large oval pedicellarian pits on
the marginal plates (Breton, 1983, 1992; Jagt, 2000; Villier et al.,
2004; Blake, 2010). All five Late Cretaceous starfish species of the
SCB, however, lack pedicellarian pits on marginals and abactinals
(Niebuhr and Seibertz, 2016). Comptoniaster (sensu Breton, 1992) is
a genus that probably represents a paraphyletic or even poly-
phyletic cluster of species sharing a ‘generalized gonisterid
morphology’ (Villier et al., 2004; Blake 2010). Morphological sim-
ilarities of completely preserved Comptoniaster comptoni to moulds
of completely preserved C. michaelisi nom. nov., are such that they
must be congeneric. In view of the fact that no comptoni- and/or
michaelisi-like asteroid was ever found in underlying strata (middle
Cenomanian to lower Turonian Quader sandstones) of the four
subbasins considered here, nor elsewhere, we consider C. michaelisi
nom. nov. to be a distinct species and interpret C. comptoni to a late
Albianeearly Cenomanian precursor.
In the SCB, Comptoniaster michaelisi nom. nov. can be differen-
tiated from the late Cenomanian Calliderma ottoi and the basal
middle Turonian Calliderma lindneri by having narrower ambulacral
furrows, a greater arm-to-disc ratio (2.2e2.5 vs 1.8 and 1.2,
respectively) and showing a conspicuously higher number of
140 B. Niebuhr, E. Seibertz / Cretaceous Research 87 (2018) 126e144

Fig. 11. Comptoniaster michaelisi nom. nov. A1, MB.E.5313; inner view of aboral side on lower bedding plane of tempestite layer, partially surrounded by imprints of superomarginals
with four arms preserved; convex, pillow-like disc missing; one of stratigraphically youngest specimens known, loweremiddle Coniacian, locality 5 in Fig. 1B, Idziko w (Bystrzyca
Kłodzka, Poland; ‘Grünsande von Kieslingswalda’). A2eA4, Close-ups of three preserved arm tips; note bisection of imprints of paired end plates; in A4, imprints of some prominent
spines; each width of photographs 6.5 mm A5, Close-up of left arm showing tessellate, regularly arranged abactinals; width of photograph 29 mm. A6, Close-up of upper right arm
showing outer surfaces of hexagonal abactinals and granulation of superomarginals; width of photograph 15 mm. A7, Close-up of granulation on outer surface of five super-
omarginals; width of photograph 7 mm. A8, Close-up of flat outer surface of small, rounded abactinals; width of photograph 7 mm.

marginals (45e50 vs 25 and 10, respectively). Lophidiaster scupini
and Nymphaster albensis (TuronianeConiacian boundary interval)
appear to be contemporaneous with C. michaelisi nom. nov. (middle
Turonianelower Coniacian). However, L. scupini has c. 90 marginals
and an arm-to-disc ratio of 5e6, while N. albensis shows c. 45
marginals, but has thin, elongated arms and an arm-to-disc ratio of
3.3e3.5 (Niebuhr and Seibertz, 2016).
6.5. Mode of life
These slightly convex (pillow-like) starfish lived on sandy bot-
toms as mobile epifaunal detritivores (compare Jangoux, 1982a),
which is substantiated by the narrow ambulacral furrow (Blake,
1983). The animals probably used their strong mouth angle plates
to grab food together with sand particles. These sand bulbs appear
 B. Niebuhr, E. Seibertz / Cretaceous Research 87 (2018) 126e144 141

Table 2
Oral/aboral positions preserved, arm and disc dimensions with the arm-to-disc ratio and the known stratigraphical position of all 16 well-preserved specimens studied;
w Bluff Sandstone, L.I. ¼ Lower Idziko
Sst.a ¼ Sandstone a, Sst.c ¼ Sandstone c, P.Q. ¼ Pirnaer Oberquader, R.B. ¼ Radko w Member; grey ¼ lectotype; asterisk ¼ no original
specimen known.

Specimen Fig. Side Arm (R) Disc (r) Arm-to-disc ratio Stratigraphical position

Scupin (1912e1913)* e aboral ? ? 2.4 L.I., low.emid. Coniacian

MB.E.5313 11 aboral 43 mm 19 mm 2.3 L.I., low.emid. Coniacian
MMG: SaK 12512 2B oral 68 mm 28 mm 2.4 P.Q., low.up. Turonian
MB.E.5316 7A aboral 55 mm 24 mm 2.3 P.Q., low.up. Turonian
old MMG no. 7431* 6B aboral 53 mm 23 mm 2.3 P.Q., low.up. Turonian
FG 193/2 8 aboral 68 mm 29 mm 2.3 P.Q., low.up. Turonian
FG 193/2a 9 aboral 75 mm 32 mm 2.3 P.Q., low.up. Turonian
MB.E.5322 7C aboral 52 mm 23 mm 2.3 Sst.c, low.up. Turonian
NHMW 1865/0010/0382 6A aboral 59 mm 26 mm 2.3 Sst.c, low.up. Turonian
BMNH E.13538 10 aboral 89 mm 35 mm 2.5 R.B., middle Turonian
NM-03939 4B aboral 63 mm 28 mm 2.2 R.B., middle Turonian
MMG: SaK 6713 4C aboral 64 mm 28 mm 2.3 Sst.a, middle Turonian
MB.E.5315 7B oral 60 mm 26 mm 2.3 Sst.a, middle Turonian
LfULG: RS 9878 e aboral 46 mm 21 mm 2.2 Sst.a, middle Turonian
MMG: SaK 6805 4A aboral 38 mm 17 mm 2.2 Sst.a, middle Turonian
MMG: SaK 6804 5 oral 75 mm 33 mm 2.3 basal Sst.a, low.mid.Tu.

Fig. 12. Taphonomic scenario for Cretaceous asteroids on sandy bottoms. A, Inner views of oral side on upper bedding planes, with continuous row of inferomarginals impressed
into former substrate (A1, below; e.g., Figs. 2B1, 5A1 and 7A1); aboral side on lower bedding plane, respectively, with continuous row of superomarginals and outer surfaces of
abactinal plates impressed into overlying tempestite layer (A2, above; e.g., Fig. 11). B, Top view of less compacted aboral side on upper bedding plane, with outer continuous row of
inferomarginals, partially overlying inner discontinuous row of superomarginals impressed into former substrate (e.g., Fig. 4A). C, Top view of depressed aboral side on upper
bedding plane, with sunken disc and continuous row of inferomarginals (e.g., Figs. 4BeC, 6AeB, 7C, 8e10), in part with discontinuous row of superomarginals preserved in
interbrachial arcs and/or arm tips (Figs. 4C1, 6A6, 8A1eA3, 9) and inner surfaces of abactinal plates (e.g., Figs. 4B2, 8A4) impressed into former substrate.

as bulbs in several specimens, in a central position of the disc (see
Figs. 2B1, 6B and 7B) and are visible, in different dimensions, in the
interbrachial arcs of Figs. 5A1 (lower left arc), 6B (lower, left and
upper left arc) and 7B (upper right arc). As shown in other aster-
oids, the cardiac stomach can be reduced alongside a conspicuous
development of the Tiedemann's pouches (Rowe et al., 1982) which
are seen with these sand bulbs. Tiedemann's organs are known to
be capable of pumping food into the digestive tract; they also act to
augment the efficiency of water circulation inside the digestive
tract (Jangoux, 1982b). Based on the digestive organisation, it may
be assumed that C. michaelisi nom. nov. was a microphagous
asteroid that fed on particulate matter, e.g., epibenthic films.
In the lectotype (MMG: SaK 12512), the lower right arm shows a
healed predation scar (Fig. 2B3); the arm is short and rounded with
large end plates and the inferomaginals are connected just from the
point where the animal was injured. The abactinals close to the
point of injury were reinforced during healing.
In order to withstand current traction, the animals sunk their
arm tips into the sandy substrate (area of downward bend in
Fig. 3B). As a consequence, arm tips often are either not preserved
on the aboral side or occur as a cavities due to the dissolution of
marginals (Figs. 4BeC, 6A1, 6A4e6, 6B, 7BeC, 8A1e2).
6.6. Preservation
All asteroids studied were embedded in life position with the
oral side below and the aboral side up (Niebuhr and Seibertz, 2016).
During life, the inferomarginals contacted the substrate directly so
that the continuous row of ossicles in their moulds represents the
exact horizontal outline of the living starfish. Superomarginals are
occasionally sunken and laterally displaced due to compaction of
the sand and appear in an outer discontinuous row of moulds in the
inner interbrachial arcs of some specimens. In most cases imprints
of superomarginals only appear in 1e3 arcs, indicating the current
direction during tempestite deposition. However, in the less com-
pacted specimen (Fig. 4A), supero- and inferomarginals are
opposed precisely as during life.
142 B. Niebuhr, E. Seibertz / Cretaceous Research 87 (2018) 126e144
All oral sides (Figs. 2B, 5, 7A) are located on upper bedding
planes and are represented by inner views of imprints into the
former substrate of the animals (compare Fig. 12A1), showing discs
with partially preserved ambulacral and adambulacral plates
covered by a continuous row of inferomarginals; actinal plates are
not visible e such would appear only in bottom-up views which are
not preserved. Inner views of the aboral sides are preserved by
imprints in lower bedding planes of the tempestitic layers
(compare Fig. 12A2), and represented by the more or less uncom-
pacted specimen (Fig. 11), in which the pillow-like, convex-up disc
is not preserved. The arms show a continuous row of super-
omarginals and portions of the underlying inferomarginals;
polygonal abactinals appear in a regular, tessellate arrangement,
their outer surfaces being flat (Fig. 11A3, A5eA7). In both inner
views of oral and aboral side parts of the opposite body disc can be
preserved, which is seen, e.g., in sunken abactinals of Fig. 2B. [The
scenario of Fig. 12A1 and A2 is based on Calliderma ottoi, the single
asteroid individual from the Quader sandstone facies known to be
preserved in both inner views of oral and aboral sides (see Niebuhr
and Seibertz, 2016, fig. 12a)].
The other twelve aboral sides appear to be top views on the
upper bedding planes of the former substrate. Top views of less
compacted aboral sides (compare Fig. 12B) are visible in Fig. 4A and,
in part, in Fig. 7B. The pillow-like, convex-up disc is surrounded by
an outer row of inferomarginals that partially overlies the sunken
superomarginals of the arms or precisely opposing in the inner
interbrachial arcs (Fig. 4A3). The specimen in Fig. 4A is closest to
what a living Comptoniaster michaelisi nom. nov. would have looked
like.
Specimens in Figs. 4BeC, 6, 7C and 8e10 are preserved as
demonstrated in Fig. 12C. Strong compression pushed the dorsal
surface of the discs downwards to the level of the continuous row of
inferomarginals, which is particularly well visible in Figs. 4BeC,
6AeB, 8 and 10. Parts of overlying superomarginals, which appear
in an outer discontinuous row, are best seen in Figs. 8A1eA3 and
10A1. The inner surfaces of abactinals that taper inwardly are well
visible in Figs. 4B2, 7A7 and 8A3. Downward-bending of arm tips
into the substrate for anchorage of the animals was carried out only
in depressed aboral sides, as demonstrated in Fig. 12C.
7. Conclusions
The Cretaceous goniasterid starfish species described under the
names of Pentagonaster semilunatus by Schulze (1760) and Asterias
schulzii Cotta by Roemer (1840, 1841) is restudied. Both Schulze's
and Roemer's names are shown to be invalid, and a replacement
name, michaelisi, is proposed. Schulze's (1760, pl. 2, fig. 6) illus-
tration allows recognition of a specimen now housed in the
palaeozoological collections of the Senckenberg Naturhistorische
Sammlungen Dresden, Museum für Mineralogie und Geologie
(MMG), and this specimen is here designated lectotype of the
species which also is reassigned to the genus Comptoniaster Breton,
1983. Occurrences of the species are documented from the middle
Turonianemiddle Coniacian of the Saxonian, North and Intra-
sudetic Cretaceous subbasins in the border triangle of Germany,
Poland, and the Czech Republic. Fourteen, out of the available 18,
well-preserved specimens are figured. These specimens allow a
precise reconstruction of morphological details; single skeletal
morphologies of the asteroid are described, such as infero- and
superomarginals, abactinals, madreporite, odontophore, ambula-
cral, adambulacral and mouth-angle plates. Accessory ossicles such
as pedicellariae, fascioles, spinelets and granules are not observed.
Comptoniaster michaelisi nom. nov. is a medium-sized species
with an outer radius up to 90 mm that is characterised by 45e50
paired marginals and an arm-to-disc ratio of 2.2e2.5. Preferred
habitats were shallow-marine environments above the fair
weather wave base with medium- to coarse-grained sandy firm-
grounds. According to its digestive organisation, C. michaelisi nom.
nov. was a microphagous asteroid feeding on deposit particulate
matter. Predation observed in the lectotype is described. The
taphonomy of the well-preserved, articulated specimens has been
reconstructed, with both death and rapid burial most likely caused
by tempestites. Great morphological affinities exist to Comp-
toniaster comptoni (Forbes, 1848) that is interpreted to be the late
Albianeearly Cenomanian precursor of Comtoniaster michaelisi
nom. nov.
Acknowledgements
For the loan of specimens or supply of photographs of Creta-
ceous asteroids studied we thank Birgit Gaitzsch (TU Bergakade-
mie, Freiberg), Peter Suhr (formerly LfULG Freiberg, now MMG
Dresden), Christian Neumann (Museum für Naturkunde, Berlin),
Mathias Harzhauser (Naturhistorisches Museum Wien), Jan Sklena r
(National Museum, Prague) and Loïc Villier (Pierre et Marie Curie
Universite, Paris). Photographs of the MMG and Berlin specimens
were taken by Ronald Winkler (MMG). Many thanks go to Andy S.
Gale (Portsmouth) and Loïc Villier (Paris) for their very helpful
reviews, as well as Gerd Geyer (Würzburg) for help in taxonomic
issues. The editors of Cretaceous Research, in particular John W.M.
Jagt, are thanked for their friendly support.
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