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IMPLICATIONS FOR THE STUDY OF FOSSIL ASTEROIDEA
(ECHINODERMATA) OF NEW GENERA AND SPECIES FROM THE
EOCENE OF FLORIDA
DANIEL B. BLAKE
1
AND ROGER W. PORTELL
2
1
Department of Geology, University of Illinois, Urbana 61801, ,dblake@uiuc.edu.; and
2
Florida Museum of Natural History, University of Florida,
Gainesville 32611, ,portell@flmnh.ufl.edu.
ABSTRACT—Oyenaster oblidus, Ocalaster timucum, and Ocalaster seloyi are new genera and species of the family
Goniasteridae (Asteroidea) described from the Eocene Ocala Limestone of Florida. Although the fossil record of
asteroids is sketchy, goniasterids appear to have been important contributors to marine communities since at least
the Middle Jurassic. Similarities between living goniasterids and their fossil precursors indicate that plesiomorphy
and convergence have been important in family history, and as a result, taxonomic interpretation is challenging.
Even partial fossil goniasterids are rare, forcing systematists to rely heavily on isolated marginal ossicles, although
some authors have expressed the need for caution. Building around three new taxa, we suggest that broader
approaches can aid systematic interpretation of all crown-group asteroids. We also suggest that the inevitably
idiosyncratic interpretations of marginal-based systematics can be partially tested using blind evaluations.
INTRODUCTION
M
ESOZOIC AND Cenozoic asterozoans have an extended
history of study in Europe, but they are relatively
neglected in North America, although paleontologists long
have been aware of their presence (e.g., Howe, 1942).
Nevertheless, asteroids are important in many extant marine
communities, and even the poor fossil record offers important
data on phylogeny, paleobiogeography, and paleoecology.
Extant asteroid shallow shelf diversity of the Florida region
is limited (Clark and Downey, 1992). Arguments of Stanley
(1986) on molluscan extinction together with the widespread
occurrence of asteroid ossicles in Cenozoic sediments (Portell
and Oyen, 2001) and the few known more nearly complete
specimens (Jones and Portell, 1988; Ivany et al., 1990; Oyen
and Portell, 2001) now augmented by those described here
indicate more diverse faunas in the geologic past, and together
they indicate good opportunities for future research.
Asteroids are skeletally complex organisms, and therefore
many characters are available to systematists. Nevertheless,
expression of both primary ossicles and their accessories
demonstrate widespread character survival (plesiomorphy)
and convergence among modern species and their fossil
precursors. More or less intact fossil asteroids are rare in all
regions, imposing emphasis on isolated marginal ossicles, yet
focus on individual skeletal elements is problematic for any
taxon. Study of other multielemental groups, such as
conodonts, has benefited from use of all available portions
of the skeleton, and multielemental approaches and other
techniques can aid the study of asteroids. In addition,
comparatively simple tests are available to partially test the
reliability of systematics based on isolated ossicles.
Except as noted, specimens herein are catalogued into the
Invertebrate Paleontology Collection, Florida Museum of
Natural History, University of Florida, and are identified by
the prefix UF followed by the catalogue number. Fossil
localities referred to throughout the paper are denoted by the
prefix FLMNH IP followed by a letter and number
designation.
TERMINOLOGY
Terminological usage follows Spencer and Wright (1966)
and Blake and Hagdorn (2003). A usually double marginal
series (inferomarginals, IM, and superomarginals, SM) arise at
the unpaired terminal at the tip of the arm and separate
abactinal ossicles of the dorsal surface from the actinal ossicles
developed between the marginals and the adambulacrals. A
differentiated primary circlet of abactinals is developed near
the dorsal center of the disk of most asteroids, and a
differentiated carinal series extends along the dorsal arm
midline of many genera. The ambulacral and adambulacral
series also arise at the terminal and extend proximally to the
mouth frame. Accessory spines, spinelets, granules, and
pedicellariae are variously developed.
OCCURRENCES, STRATIGRAPHY, AND DEPOSITIONAL SETTINGS
A single fairly complete specimen and four associated
fragments of Oyenaster oblidus n. gen. and sp. were collected
in situ from near the top of an eight-meter quarried section of
Upper Ocala Limestone at FLMNH IP AL004, Alachua
County, Florida. Two incomplete external molds of two
goniasterids (Ocalaster timucum and O. seloyi n. gen. and spp.)
were recovered from a quarried dolomitized section of Lower
Ocala Limestone (just above contact with the middle Eocene
Avon Park Formation) at FLMNH IP CI009, Citrus County,
Florida. Before the present occurrences, no articulated body-
fossil goniasterid specimens had been recovered from the
Ocala Limestone.
Robust isolated marginal ossicles of goniasterid asteroids
are common in Florida exposures of the Eocene Ocala
Limestone. An especially large ossicular concentration
(.15,000 specimens) was sampled in a sinkhole exposed
during excavation of the Cross Florida Barge Canal (FLMNH
IP UF CI001, Citrus County). The solution feature was
infilled with approximately 300 m
3
of eroded Lower Ocala
Limestone, washed-in late Pliocene terrestrial sediments, and
Holocene sands (Klein, 1971; Meylan, 1980). On-site screen
washing of some of these sediments, primarily for vertebrate
remains, yielded 15,324 small echinoid tests (Zachos, 2005;
personal commun., 2008) along with the abundant isolated
goniasterid ossicles. Geographic data are summarized in
Figure 1.
The Ocala limestone was named by W. H. Dall (in Dall and
Harris, 1892) for all Jacksonian limestones found in central
Florida. Cooke (1915) acknowledged the formation and
J. Paleont., 83(4), 2009, pp. 562–574
Copyright ’ 2009, The Paleontological Society
0022-3360/09/0083–0562$03.00
562
recognized the unit as late Eocene. Applin and Applin (1944)
divided the Ocala Limestone into two members (Upper and
Lower) based on microfossil and lithologic evidence. Vernon
(1951) and Puri (1957) further subdivided the unit, recognizing
the Ocala Group that consisted in sequence of the Inglis,
Williston, and Crystal River Formations. This terminology
remained in use until the Florida Geological Survey (Scott,
1991; personal commun., 2008) returned to the broader
stratigraphic terminology of Applin and Applin (1944) in order
to adhere to the North American Stratigraphic Code (North
American Commission on Stratigraphic Nomenclature, 1983).
Scott’s decision was based solely on lithologic differences, the
Upper Member consisting of a soft, friable, variably muddy,
granular limestone and the Lower Member being a more
granular limestone that in some localities is partially to
completely dolomitized. Nevertheless, the Inglis, Williston,
and Crystal River terminology is thoroughly entrenched in the
literature. Stratigraphic data are summarized in Figure 2.
The Ocala Limestone is exposed in northwestern peninsular
Florida and a small portion of the Florida panhandle along
the border with Georgia and Alabama. The thickness of the
Ocala Limestone ranges from less than 30 m in the northern
panhandle and central peninsular region to more than 120 m
in southern Florida and the Gulf Coast region of the
panhandle (Chen, 1965).
The environment of deposition suggested by Cheetham
(1963) and Chen (1965) for the Ocala Limestone across the
northwestern portion of the Florida peninsula is that of a
carbonate buildup in an open, warm, shallow sea in 45 m of
water depth or less. Based on endolithic, nestling, and
burrowing bivalves, Krumm and Jones (1993, p. 949) further
determined that a ‘‘complex network of hardgrounds, firm
muds, and small coral patches, along with unconsolidated
sediment’’ were present at the time of deposition. Fenk (1979)
reported the lithology of the Inglis Formation (Lower Ocala
Limestone) to be primarily a clean packstone and grainstone
that represents a higher energy, subtidal environment, whereas
the Crystal River Formation (Upper Ocala Limestone) fauna
and muddy packstone and wackestone lithologies represent a
lower energy (below wave base), deeper subtidal deposit.
SYSTEMATICS OF FOSSIL CROWN-GROUP ASTEROIDS
General background.—Many fossil asteroids are similar to
others alive today, and as a result paleontologists long have
assigned ancient species to modern genera (e.g., Forbes, 1848).
During the 19th century, however, efforts to compare fossil
and living taxa were hampered by the still-emerging under-
standing of the systematics of the extant fauna. The basic
familial concepts and assignments in use today, developed
through the efforts of many researchers, have been in place at
least since the monographs of Fisher (1911, 1919, 1928, 1930).
Clark and Downey (1992), Clark (1989, 1993, 1996), Clark
and Mah (2001), and Mah (2009) surveyed living representa-
tives, and knowledge continues to expand with revisions,
exploration of still poorly known regions, and the discovery of
new species (e.g., Mah, 2007).
The focal area for the study of fossil post-Paleozoic
asteroids has been northwest Europe, and many of the
important references are cited in discussions below. Significant
papers continue to appear, including Breton (1988, 1992, 1995,
1997a, b), Breton and Vizcaı
¨
no (1997), Gale (1986, 1987,
1989), Jagt (2000), and Villier et al. (2004b). The valuable
relatively comprehensive comparative generic surveys of
Spencer and Wright (1966) and especially Breton (1992) are
particularly useful. Work has tended to emphasize Mesozoic
faunas, but important papers on Cenozoic fossils are
available, for example Nielsen (1943), Rasmussen (1950,
1972), and Jagt (2000).
Difficulties of study.—Many difficulties burden the system-
atics of fossil crown-group asteroids. The fossil record is poor,
although more or less complete specimens such as those
described here are widely illustrated, they generally are very
rare. An articulated skeleton of comparatively small elements
FIGURE 1—Map of Florida showing type localities of Oyenaster oblidus,
Ocalaster timucum, and O. seloyi n. gen. and spp., FLMNH IP AL004,
Alachua County, Newberry Quadrangle, USGS 7.59, 1988; T9S, R17E;
and CI009, Citrus County, Yankeetown Quadrangle, USGS 7.59, 1988;
T17S, R16E. Also shown is FLMNH IP CI001 (Citrus County, Yankee-
town Quadrangle, USGS 7.59, 1988, T17S, R16E) site of large
ossicular concentration.
FIGURE 2—Historical summary of stratigraphic usage of Ocala
Limestone in Florida (modified from Oyen and Portell, 2001).
BLAKE AND PORTELL—FLORIDA EOCENE ASTEROIDS 563
and the presence of an enlarged coelom with voluminous soft
organs render asteroids readily disrupted and destroyed by
both physical and biological processes. Further, skeletal
element types appear susceptible to selective destruction
because published specimen listings indicate that marginals
predominate, although ossicles of a number of other series
(e.g., ambulacrals, adambulacrals) are at least as abundant in
the living individual.
Adding to difficulties of working with isolated skeletal
elements, ossicular types vary significantly with ontogeny and
position in the body, as well as among individuals. Grada-
tional changes in morphology (as along an arm) are difficult to
categorize for cladistic purposes or in descriptions.
Expression of accessory ossicles (spines, spinelets, granules,
pedicellariae) also has been important to the taxonomy of
living asteroids. Nevertheless, taxonomically widely distribut-
ed similarities of accessory expression indicate evolutionary
convergence. Adding to difficulties, accessories can be largely
lost from otherwise skeletally complete fossils; where pre-
served, these generally crowded small elements tend to hide
underlying primary ossicles. During preservation, life expres-
sion is obscured by fusing, displacement, overlap, and
recrystallization.
As a result of the poor fossil record, many taxa have been
based on isolated ossicles. As a result, taxonomic interpreta-
tion is difficult and diverse research approaches are needed.
Background to the Goniasteridae.—The Goniasteridae has
perhaps the best fossil record among crown-group asteroid
families. Goniasterids are both numerically abundant and
diverse today, and although the fossil record of all asteroids is
limited, an extended literature (e.g., Hess, 1972; Breton, 1992)
indicates that familial importance has been ongoing at least
since the Middle Jurassic.
The configuration of the new taxa, including large disks
with curved interbrachial margins, more or less extended arms,
tabular interbrachial marginals wider than long that grade
distally into those of more equidimensional proportions, and
rather small, similar abactinals, are widely distributed among
valvatidan and paxillosidan asteroids throughout their histo-
ry. For example, Hess (1972) viewed Calliderma Gray, 1847;
Comptonia Gray, 1840; and Tylasteria Valette, 1929, as
representing a widely distributed Jurassic and Cretaceous
asteroid type. Breton (1984, 1992) in effect agreed with Hess’s
overview with his recognition of Comptoniaster Breton, 1984,
for a relatively large group of similar species including one
assigned to Tylasteria by Hess (1972). Subsequently, Villier et
al. (2004b) argued that Comptoniaster is based on an
association of primitive character states, that the genus might
be paraphyletic, and that it might provide sister taxa for
multiple stellate Mesozoic goniasterid lineages. Villier et al.
(2004b) thus in effect recognized strong similarities among
ancient species, and monographs of living faunas (e.g., Fisher,
1919) demonstrate that similarities recognized by these
authors are still to be found.
Shallower shelf settings with particulate substrates that are
conducive to goniasterid occurrence have endured through
geologic time, promoting survival and plesiomorphy and
encouraging convergence and homoplasy; convergence is
illustrated by a superficially goniasterid-like Early Ordovician
asteroid (Blake et al., 2007) that is only remotely related.
Because of convergence, even comparatively complete fossils
can be taxonomically difficult.
Difficulties of fossil asteroid systematics.—Body form where
preserved is important to the taxonomic interpretation of both
living and fossil asteroids (e.g., Spencer and Wright, 1966).
Nevertheless, overall proportions can change significantly
during ontogeny (Breton, 1997b), indicating a need for
caution.
Unlike still-articulated specimens, isolated marginal ossicles
locally are comparatively common and therefore have received
much attention. Nielsen (1943) structured his introductory
material around individual ossicular types; although treatment
was expanded later in this text, his characterizations of
marginals in effect recognized form genera. Some researchers
have reconstructed the complete individual from isolated
ossicles (e.g., Villier, 1999; Zitt, 2005), and such approaches
have been tested (Breton, 1995). Marginal form has been
emphasized in inferences on phylogeny (Spencer, 1913;
Breton, 1997a, 1997b), biogeography (Gale, 1989), paleoecol-
ogy and biostratigraphy (e.g., Villier et al., 1997; Jagt, 1999,
2000; Villier et al., 2004b), and taphonomy (Zaton, et al.,
2007).
Along with progress in the interpretation of the significance
of marginal morphology, authors have expressed concerns.
Rasmussen (1950) noted that the species diversity within the
Danish chalks recognized by Nielsen (1943) was ‘‘suspicious’’
(Rasmussen, 1950, p. 15) given apparent equivalent modern
faunal occurrences; the latter author thus tested Nielsen based
on extant faunas. Villier et al. (2004b) cogently defended the
taxonomy of isolated ossicles, but these authors also observed
that comparisons are more difficult without extant or well-
defined fossil controls. In proposing a method of reconstruct-
ing skeletal architecture and body shape from isolated ossicles,
Villier (1999, p. 354) demanded ‘‘no ambiguity concerning the
identification of the species and the deposits must contain a
few dozen marginal plates.’’ In discussing his collection of
marginals of the type species of the goniasterid Ophyraster
Spencer, 1913, Nielsen (1943, p. 61) in a sense presaged the
concerns of Villier: ‘‘If such plates are found isolated, it is
impossible to imagine that marginalia from the disc and the
arm belong to the same species, even to the same genus.’’
Many of Nielsen’s species are no longer recognized, logically
because he followed his own form-taxon approach and split
more finely than recognized by subsequent workers, but also
perhaps because he did not sufficiently heed his own warning.
Regardless of why Nielsen’s species concepts have been
challenged, this author deemed it worthwhile to forcefully
call for caution in the interpretation of isolated ossicles.
Future researchers seem likely to continue to evolve their own
approaches to marginal analysis and taxon recognition,
perhaps even resurrecting some concepts of Nielsen. Breton
(1992) illustrated considerable variation among marginal
ossicles for Crateraster quinqueloba (Goldfuss, 1831), a species
recognized from comparatively abundant material. Weather-
ing prior to final burial can cause significant damage (e.g.,
Villier, 2008), a difficulty also encountered with the isolated
Ocala marginals. These conclusions of capable practitioners
must raise concerns about taxa based on small or scattered
ossicular samples.
Available approaches to asteroid systematics.—Species and
generic concepts in paleontology for multielemental organism
groups such as the Asteroidea almost inevitably are based on
incomplete specimens. Perhaps no example of the significance
of this limitation is more clear than that of conodonts, in
which studies have progressed from elemental form taxonomy
to a still-developing multielemental approach (Sweet and
Donoghue, 2001). In parallel with work on conodont
assemblages, studies of those fossil asteroids that are still
partially articulated need to fully exploit all available ossicular
types. However, the fossils here also illustrate inevitable
564 JOURNAL OF PALEONTOLOGY, V. 83, NO. 4, 2009
difficulties in that important ossicular types are unexposed,
incompletely exposed, and poorly preserved.
Paleontologists have interpreted fossils based on compari-
sons with extant species since some of the earliest studies;
however, comprehensive efforts to link systematics of fossils to
living forms are comparatively few. Taxonomic keys provide
another useful tool in that they enable at least approximate
identification of crown-group fossil asteroids. The mono-
graphs of Fisher (1911, 1928, 1930) provided comprehensive
keys to extant taxa, and more detailed treatments (e.g., Clark
and Downey, 1992; Mah, 2005, 2007) are becoming available.
A key to fossil marginal ossicles was provided by Spencer
(1907), and Breton (1992) included keys to genera as well as to
species within genera.
Developing cladistic treatments offer insights not available
to earlier workers. Mah (2007), for example, was able to
clarify the systematic position of Buterminaster Blake (in Blake
and Zinsmeister, 1988) in a study that included a number of
extant genera. Villier et al. (2004a) used cladistic analysis in
description of a new genus and reevaluation of the small
extinct family Stauranderasteridae. The fragmentary nature of
fossil asteroids is a limitation for cladistic study, but many
parallel examples from fossil vertebrates suggest at least
partially surmountable difficulties.
Molecular studies on asteroids to date have yielded
inconsistent results. Such work in the future, however, can
be expected to broaden the understanding of both systematics
and phylogeny, and these results provide useful perspectives
for paleontology.
There has been little attempt to evaluate ranges of variation
within genera, species, populations, and individuals of either
fossil or extant asteroids. Computer-imaging accompanied by
statistical analysis is becoming more common for other phyla,
and such work appears to promise important advances for the
evaluation of ossicular morphology. The complexly articulat-
ed ambulacral column and mouth frame offer opportunities
for understanding of not only systematics and phylogeny but
also functional morphology [e.g., Eylers (1976); O’Neil (1989,
1990)]. Logically, imaging work begins with populations of
extant species so that closely related taxa can be compared.
Ecologic and biogeographic variation need to be treated.
Study of living representatives can provide insight into ranges
of variation to be expected among ancient species as well as
the reproducibility of species concepts.
Testing the reliability of marginal taxonomy.—The literature
of fossil crown-group Asteroidea includes many examples of
assignment of isolated marginal ossicles to genera and species,
including taxa based on ossicles alone. This literature has been
developed by careful and thoughtful paleontologists, yet
reassignments are not uncommon; we argue that the example
of conodont assemblages (and examples from other groups
could be selected) indicate the need to test the reliability of
isolated elemental taxonomy. At least two test approaches are
available. First, a paleontologist asks an assistant to isolate
perhaps 100 ossicles taken from modern individuals repre-
senting different genera and species. Both inferomarginal and
superomarginal ossicles are included, and marginals are taken
from specimens of different sizes and positions on the disk and
arms; numerical balance among sampled species is avoided (or
not). The paleontologist has only isolated ossicles and must
decide values for many parameters for the suite. The study
program can be expanded with multiple tests of differing levels
of difficulty by varying test taxon similarities, ontogenetic
stages, source positions on parent specimens, and numbers of
ossicles representing individual species. In a second experi-
mental type, the assistant enables blind re-evaluation of large
and diverse fossil samples some months after the original
treatment by the paleontologist to determine how well later
interpretations support earlier assignments.
Study of the new specimens.—New material documented
here illustrates many of the study difficulties touched upon
above. Cladistic treatment and statistical analyses entail
larger-scale projects than the present study, and a more
limited approach is appropriate. We follow a ‘‘multielemen-
tal’’ methodology in documenting as possible all available
ossicular types. Assessments then began with the comprehen-
sive treatment of living taxa provided by Mah (2005) before
turning to the literature on fossils. Interpretations stressed
available more-objective criteria such as spines and pedicellar-
iae, although workers with modern species recognize impor-
tant variability in these features. Although hundreds of
isolated marginal ossicles were available and many are
tentatively separated and available in Florida Museum of
Natural History collections, isolates did not provide a basis
for any of the new species and they are not a part of any of the
type suites.
SYSTEMATIC PALEONTOLOGY
Class ASTEROIDEA de Blainville, 1830
Order V
ALVATIDA Perrier, 1884
Family G
ONIASTERIDAE Forbes, 1841
Discussion.—The families Goniasteridae and Oreasteridae
as traditionally formulated (e.g., Spencer and Wright, 1966)
encompass similar genera; in different studies, some genera
have been assigned to both families. Larger oreasterids
typically have relatively small and more or less rounded
marginals, unlike the robust, tabular, closely abutted ossicles
of the Goniasteridae, including Oyenaster and Ocalaster. Most
oreasterids have a well-developed granular-dermal layer that
extends into depressions between the marginals; any indication
of such a layer is lacking from the new genera. Oreasterids
have comparatively tall adambulacral ossicles that contribute
to a protective armor, whereas adambulacrals of Oyenaster are
low (those of the Ocalaster specimens are not exposed). The
rather uniform, tabular abactinals with more or less promi-
nent basal flanges (which provide for discrete papulae rather
than papulary fields), the absence of strongly differentiated
and enlarged carinal series and primary circlet, and the
absence of robust, conical nodes or spines all indicate that the
new genera are goniasterids in the traditional sense.
Genus O
YENASTER new genus
Type species.—O. oblidus, new species; the genus is
monospecific.
Diagnosis.—As for the species.
Description.—As for the species.
Etymology.—Named in honor of Dr. Craig Wendell Oyen
(1963–2005), who discovered the fossil and contributed
significantly to the study of Florida fossil echinoderms.
Occurrence.—As for the type species.
O
YENASTER OBLIDUS new species
Figures 3.1–3.6, 4
Diagnosis.—Overall form and ossicular form robust; arms
broad, short, triangular; abactinals probably reaching termi-
nal. Spines, enlarged accessory bases, pedicellariae absent
from all ossicular systems. Marginals robust; interbrachial
ossicles wide, outer face transverse profile weakly positively
arched medially, curving abruptly at abradial margin to form
a shallow grove or furrow zone along marginal series contact.
BLAKE AND PORTELL—FLORIDA EOCENE ASTEROIDS 565
FIGURE 3—1–6. Oyenaster oblidus n. gen. and sp., holotype, Florida Museum of Natural History UF 50000a, Upper Ocala Limestone, Florida. Scale
bars 1, 2, 5–6, 15 mm; 3, 4, 5 mm. 1, 2, dorsal and ventral views of entire specimen; 3, 4, ossicular arrangement of ventral surface, arrows identify
adambulacral series positioned at the straight furrow spine attachment line; two series of actinals adjacent to adambulacrals are robust and uniform; 5, 6,
oblique and lateral views of arm margin showing ossicular form and overhang of marginals to form a shallow furrow between the marginal series. The
tiny distal adambulacrals are upturned so that podia were directed distally; 7, 8, Calliderma spectabilis Fisher, 1906, Bishop P. Museum W3732. Scale
bars 10 mm. 7, dorsal lateral view of portions of disk and arm, small spine bases occur on marginals (arrow); 8, ventral view of disk with interbrachium
partially cleared.
566 JOURNAL OF PALEONTOLOGY, V. 83, NO. 4, 2009
Marginal outer surfaces covered by uniform, densely spaced
small accessory depressions. First actinal series ossicles
adjacent to adambulacrals somewhat irregular in outline.
Adambulacrals slightly wider than long. Furrow margin
spines probably aligned in a straight, longitudinal series (i.e.,
not an angular series on each ossicle).
Description.—Five-armed asteroid; outline subpentagonal,
interbrachia broadly rounded and grading evenly into short,
triangular arms; abactinal field narrows sharply but super-
omarginals not appearing to abut at arm midline, i.e., few
abactinal series appearing to reach terminal area; arm tips
broadly rounded. Arrangement of preserved abactinal ossicles
suggest body profile low in life (e.g., as in Mediaster Stimpson,
1857, but not as in Oreaster Mu¨ller and Troschel, 1842).
Spines lacking. Arm radius of holotype 115 mm, interbrachial
radius 80 mm.
Abactinals appearing small for holotype size; ossicles low,
tabulate; ossicles polygonal, flanged for papulae; extended
papulary areas intercalated among ossicular series not recog-
nized; dorsal ossicular surfaces flat. Carinal series, primary
circlet ossicles weakly enlarged. No enlarged accessories
developed, smaller accessories appear to have been closely
fitted, uniform, granular or perhaps weakly elongate, spine-like.
Marginals in double, nearly paired series, about 12 from
interbrachial sinus to arm tip of holotype. Marginals simple in
form; robust; interbrachial ossicles tabular, broad relative to
FIGURE 4—Oyenaster oblidus n. gen. and sp., holotype, Upper Ocala Limestone, Florida. Views of marginal series and adjacent series, oriented with
IM below. Scale bars, 5 mm. 1–4, UF 50000e; 1, inclined ventral view; 2, lateral view; 3, IM and edge of SM series, abradial up; 4, abradial view. 5–9,UF
50000c, 5, 6, inclined dorsal views, right SM as seen in 6 partially replaced by small ossicles; 7, lateral view; 8, IM series, abradial up; 9, abradial view. 10–
12, UF 50000d; 10, inclined dorsal view; 11, IM and edge of SM series, abradial up; 12, abradial view.
BLAKE AND PORTELL—FLORIDA EOCENE ASTEROIDS 567
height, short; ossicular plan-view outlines becoming more
nearly equidimensional on arms. Marginals of two series
closely abutted, side faces closely abutted; adradial edges of
both series generally straight, can be angular if abutting
multiple adjacent abactinals or actinals. Groove between
successive marginals narrow but sharply defined. Transverse
ossicular profiles convex, can be arched such that most dorsal
point of SM (most ventral of IM) about at mid-ossicle.
Ossicles of both series curving abruptly at abradial margin,
abutting to form a shallow, peripheral furrow zone. At mid-
ossicle, longitudinal ossicular profiles weakly arched, that of
SM perhaps more positive than that of IM; profile saddle-
shaped abradially. Enlarged accessory bases not recognized;
ossicular surfaces covered by uniform, closely spaced small
depressions; depressions appearing uniform to ossicular edges,
i.e., no differentiated border zone nor reduction of depressions
near adradial and intermarginal edges; no bare or raised areas
developed. Side faces bordered by low rim, tissue surfaces flat,
closely appressed. Adradial faces bear subdued articular
facets. Skeletal debris suggests granular accessories. Distal-
most inferomarginals deflected dorsally but arm tip as a whole
at most only weakly upturned; inferomarginals near tip might
protrude laterally slightly beyond superomarginals.
Terminal small, triangular, appearing flush with arm tip.
Actinals tabular, closely fitted, forming robust ventral
pavement. Longitudinal actinal series immediately adjacent
to adambulacrals enlarged, ossicles rectangular; second
longitudinal series similar to first but ossicles smaller, nearly
square in outline; second series ossicles nearly paired with first
series. Two or perhaps three additional series of smaller,
robust, rectangular ossicles present in each interbrachial arc.
Actinal ossicular arrangement appearing locally irregular.
Outer surfaces flat, bearing small spinelets.
Adambulacral ossicles tabular, upright, weakly overlapping;
articular flanges contacting ambulacrals positioned near
adradial margin; outer face rectangular, long axis transverse,
surface bearing small granules. Adradial face edge notched for
enlarged furrow spines, furrow spines arranged in linear (not
curved or arched) longitudinal series (numbers in series
uncertain). Adradial surfaces flattened (allowing tight furrow
closure). Ambulacral ossicles of typical goniasterid form:
robust, adradial head strongly overlapping.
Neither pedicellariae valves nor pedicellariae alveolae
recognized on any ossicular series. (Linear markings on
certain ossicles visible on ventral ossicles of Fig. 3.3, 3.4 are
preservational artifacts and debris.) No enlarged accessories
recognized.
Mouth frame ossicles not exposed.
Etymology.—oblido, L., squeeze together, for the distinc-
tive, tightly appressed marginals.
Material and occurrence.—Holotype UF 50000a, consisting
of the central portion of the disk, which is largely obscured by
sediment and skeletal debris, one complete arm, and portions
of two more arms. In addition, four associated fragments (UF
50000b–e) were available.
Dickerson Limerock Mines (Haile Compex) (FLMNH IP
AL004), Alachua County, Florida. Newberry Quadrangle,
USGS 7.59, 1988; T9S, R17E. Upper Ocala Limestone, upper
Eocene.
More than 600 isolated ossicles (UF 38231, UF 38266, UF
107216, UF 116669–116671) were examined; all are abraded
and have suffered some leaching. Marginal ossicles with
proportions such as those found in the Ocala Limestone are
widely encountered in the Goniasteridae, and surface features
of Ocala ossicles are comparatively simple and provide few
characters. Ossicles are abraded, and wear could have led to
convergence of apparent morphologies from different taxa.
Presence of a few astropectinid marginals in the collection
indicates a greater diversity than that documented by partial
specimens, and perhaps also including further goniasterid
species. In accord both with the implications of weathering in
the Ocala environments and also with perspectives expressed
here, we have not based interpretations of any of the three new
species on isolated ossicles, and we do not include isolated
ossicles as a part of any of the type suites.
Discussion.—Asteroid morphology can change significantly
with growth and therefore character values change as well.
Only a single articulated specimen of Oyenaster is available; it
is of moderate or moderately large size for its family, and
descriptions above and interpretations below assume approx-
imate final morphology for its species. Many fossil goniasterid
genera are not treated in the survey below because their form
and specifics of ossicular morphology are subjectively
considered distinct from those of Oyenaster. Names treated
as synonyms by Breton (1992) are not included.
Taxonomic assessment began here with the cladistic analysis
and comprehensive diagnostic key of Mah (2005), which
focused search on Calliderma (Fig. 3.7, 3.8) and Nymphaster,
two genera also recognized in the fossil record. Calliderma is
rarely encountered today, whereas Nymphaster is quite
common; of the two, Calliderma emerged as the more
promising.
Two extant species have been recognized for Calliderma, C.
emma Gray, 1847, and C. spectabilis Fisher, 1906. Fisher
(1906) found C. spectabilis to be very close to C. emma,
differing reliably only in development of abactinal spines. Liao
and Clark (1995, p. 93) suggested that Fisher did not realize
‘‘that the apparent absence of such spines in Gray’s figure of
the holotype of C. emma is due to damage in preservation’’
(even the systematics of Recent species can be affected by
small sample sizes and ‘‘taphonomic’’ alteration). Liao and
Clark (1995), however, did not quite take the step of
synonymizing the two species.
Neither Mah (2005) nor Fisher (1911) provided a diagnosis
for Calliderma, whereas the description of Fisher (1906) was
detailed but directed toward C. spectabilis. A number of
Cretaceous species have been assigned to Calliderma, and
diagnoses focused on fossils have been provided by Schulz and
Weitschat (1971, 1975), Gale (1987), and Breton (1992). In
contrast, comparisons of the Oyenaster material with Calli-
derma focused on the extant type species. Marginals in
Oyenaster and most fossils assigned to Calliderma are quite
simple in form, and simple marginals in both provide
similarities and unfortunately also limits available characters.
Secondary accessory depressions are subtle in both genera.
Oyenaster and extant Calliderma share a straight series of
furrow spines, abruptly rounded marginal series that together
yield an abradial furrow zone between marginal series, and
enlarged interbrachial actinal regions. These character expres-
sions occur widely among goniasterids.
Oyenaster and extant Calliderma differ in that Oyenaster
lacks spines, and marginals do not abut at the midline of the
comparatively short (R:r of Oyenaster ,1.5:1; of C. spectabilis
2.37:1.) arms. Both of these characters can vary among
recognized living genera, although not in extant Calliderma.
Other differences include adambulacrals that are wider than
long in Oyenaster but longer than wide in Calliderma.
Abactinals, poorly exposed in available Oyenaster, do not
appear to be as low (i.e., the vertical dimension) as in
Calliderma. The actinal series are more uniform in arrange-
568 JOURNAL OF PALEONTOLOGY, V. 83, NO. 4, 2009
ment in Calliderma. Pedicellariae have not been recognized in
either Oyenaster nor C. emma, whereas both Schulz and
Weitschat (1975) and Breton (1992) stressed the significance of
pedicellariae in fossils assigned to Calliderma. Oyenaster and
C. emma therefore are similar but judged generically distinct.
Fossils assigned to Calliderma are similar to Oyenaster in
overall form and presence of tabular marginals (see especially
drawings of Sladen, 1891), but such marginal form likely is
plesiomorphic. More comprehensive treatment of fossil species
assigned to Calliderma is needed, but the fossils are judged
different enough from the type species as well as Oyenaster
that they are here considered a separate issue.
Tylasteria Valette, 1929, (as Tylaster Spencer, 1913) was
proposed as a subgenus of Calliderma. Breton (1992) found
the differences between Tylasteria and Comptoniaster to be
subtle, and Villier et al. (2004b) suggested that Comptoniaster
is based on association of primitive character states and that it
might be paraphyletic, providing sister taxa for multiple
stellate Mesozoic goniasterid lineages. Species assigned to
Tylasteria share generalized form with Oyenaster, although
the arms are longer, marginals of both are tabulate, becoming
more strongly equidimensional on the arms. Transverse profile
is not as angular in Tylasteria, and accessories appear more
robust. Tomidiaster Sladen, 1891 was synonymized with
Calliderma by Gale (1987) but retained by Breton (1992). It
has a large disk but also long arms and apparently deep
interbrachial sinuses; pedicellariae are present.
Nymphaster is the sister genus to Calliderma sensu Mah
(2005). It includes many species, all with superambulacrals
abutted across the midline beginning near the base of the arm.
Abutted marginals appear to be a synapomorphy of the
Calliderma-Nymphaster pair, a character excluding Oyenaster.
In Nymphaster, the furrow margin spines are arranged in an
angular row, whereas they lie in a straight series in C. emma,
and although revealed only by the linear furrow ridge on
denuded adambulacrals of Oyenaster (Fig. 3.3, 3.4), the
arrangement appears linear here as well. Number of furrow
spines is unknown in Oyenaster.
Other surviving genera broadly similar to Oyenaster are
those belonging to what Mah (2005) referred to as the
Ceramaster/Peltaster/Sphaeriodiscus Complex; this author
found significant convergence or plesiomorphy and paraphyly
among the species of the group. Marginal form appears to
provide useful discriminating criteria for these genera in that
the marginal accessories appear more complexly differentiated
in all three than they are in Oyenaster; further, raised, bare
areas are typical. Variation among extant species assigned to
Ceramaster suggested a need for re-study to that author;
nevertheless, marginals in this genus appear comparatively
elongate, whereas those of Sphaeriodiscus Fisher, 1910, and
Peltaster Verrill, 1899, are robust and nearly equidimensional.
In Sphaeriodiscus, distal arm marginals are abutted, the
penultimate enlarged in some species. Abactinals of Cera-
master are more nearly paxilliform and more regular in form
and arrangement than those of Oyenaster, whereas those of
Peltaster are flat-topped. Although differences are of degree,
actinal ossicles in the Complex sensu Mah (2005) lack the
differentiation found in Oyenaster.
Among extinct genera, Crateraster Spencer, 1913 shares a
broad disk and short arms with Oyenaster; marginals are short
and the marginal profile abruptly curved, and the intermar-
ginal contact can be depressed (Gale, 1987; Breton, 1992). In
Crateraster, however, there are only 5 to 10 marginals in each
half radius as opposed to about 12 in Oyenaster. In some
species of Crateraster, distal superomarginals abut at the arm
midline (Spencer and Wright, 1966; Gale, 1987; Manso, 2006).
Marginals in assigned species are shorter than those of
Oyenaster and can be approximately equidimensional, unlike
the tabular ossicles of Oyenaster. Certain species now assigned
to Crateraster lack the crater-like accessory bases but
accessories are nevertheless more complexly differentiated
than in Oyenaster. Breton (personal commun. in Blake and
Reid, 1998) reasonably suggested assignment of Formalhautia
Blake and Reid (1998) to Crateraster, although these authors
decided inclusion would unduly expand the generic concept.
Illustrations of specimens of Crateraster suggest absence of the
actinal surface ossicular arrangement of Oyenaster. Nielsen
(1943) also stressed presence of fairly large circular spine
depressions.
Like Oyenaster, Recurvaster Nielsen, 1943, has short arms,
but these are distally upturned, and borders of the marginals
are differentiated. Differentiation of accessories appears to
occur in at least most species, although the character is not
included in all diagnoses of the genus.
Comptoniaster, which has been recognized only from
Mesozoic rocks, is broadly similar to Oyenaster. The
interpretation of Villier et al. (2004b) of Comptoniaster as a
relatively plesiomorphic genus is noted above under Tylas-
teria. Arms are relatively long in Comptoniaster, and the
interbrachial sinus is not as broadly curved as in Oyenaster.
Interbrachial marginals are broad, although those of the arms
more nearly equidimensional. Illustrations (Breton, 1992)
suggest marginal profile and accessory base development can
resemble those of Oyenaster, but ventral interbrachial ossicles
are uniform in size in Comptoniaster and unlike those of
Oyenaster. Marginals in Cretaceous Comptoniaster are most
distinctive in that they have large, oval pedicellariae alveolae
(Villier et al., 2004b).
Genus O
CALASTER new genus
Type species.—O. timucum, new species
Diagnosis.—Disk large, interbrachial sinuses broadly
rounded, arms probably elongate. Carinal series distinctly
enlarged relative to adjacent ossicles; carinals wider than long,
with prominent basal facets. Other abactinals approximately
equidimensional, faceting distinct only on abactinals near
carinals (i.e., disk papulae confined to area above ambulacral
columns). Abactinal spines, small pedicellariae alveolae can
occur. Interbrachial marginals rectangular, convex, some
superomarginals in O. timucum with one or two small spines,
remainder of surfaces with subdued, uniform accessory
depressions. Marginal side face tissue depressions deep, outer
faces bordered by prominent articular facets. Marginal
adradial faces faceted for intermarginal, adradial ossicular
contacts.
Description.—Disk large, low; interbrachial sinuses broadly
rounded; arms distinct in both known species, length
uncertain, but marginals appearing to converge toward
midline.
Abactinals small, uniform low tabular, closely fitted,
polygonal; dorsal surfaces nearly flat to weakly domed.
Carinals larger than adjacent abactinals, wider than long,
basal facets prominent; ossicles of adjacent series more nearly
equidimensional, rows adjacent to carinals with prominent
facets, faceting on remaining abactinals subdued. Small medial
spines with circular bases present on some abactinals,
remainder of surface granulate; small pedicellariae can occur.
Interbrachial marginals tabular, sides weakly convergent
abradially. SM transverse profile convex, curving more
abruptly near abradial margin but abradial profile not
BLAKE AND PORTELL—FLORIDA EOCENE ASTEROIDS 569
FIGURE 5—Ocalaster timucum n. gen. and sp., holotype, Florida Museum of Natural History UF 20565, Lower Ocala Limestone, Florida. Scale bars
all 5 mm. 1, Dorsal view of entire specimen, grooved ridge-like feature in upper center is a sand dollar fragment; 2, lower interbrachium of 5.1; three SM
flanked by IM showing dorsal aspect; abactinals with accessories, a few with basal facets; 3, convergence of marginals suggests a narrow, elongate arm;
to right is marginal series of 5.2; 4, three each SM and IM in abradial view, series of 5.2; 5, SM series, abactinals, some with basal facets, pedicellariae
alveolae; marginal series to left of 5.3; 6, abactinals, basal facets, pedicellariae alveolae, circular spine bases (arrow); 7, lateral view of SM-IM pair,
abradial right; 8, dorsal view of two SM, that to left with two circular bases toward abradial (upward) edge, fine accessory bases over remainder of
surface; 9, slightly oblique view of SM to left, interior of IM to right; abradial to top of photograph; side faces are deep and adradial faces have
distinct facets.
570 JOURNAL OF PALEONTOLOGY, V. 83, NO. 4, 2009
angular; longitudinal profile convex, not saddle-shaped
abradially. Spines can be present but absent from most
marginals, otherwise outer face covered by subdued equidi-
mensional accessory depressions for spherical or tessellated
polygonal granules. SM side faces with deep tissue depressions
rimmed by well-developed ridges; adradial faces faceted for
abactinal, actinal contacts. Zone of juncture between SM and
IM series only weakly depressed or not depressed to form
marginal furrow. Ambulacrals of conventional goniasterid
form: Adradial flange triangular, overlapping proximally;
dorsal ridge narrow, rounded; ambulacral articular wings
prominent, symmetrical.
Etymology.—The fossil is derived from the Ocala Lime-
stone, which is named for Ocala, Florida. The word ‘‘Ocala,’’
variously rendered, is thought to be from the Timucua
language; the Timucua are an extinct Native American tribe.
The name is in honor of the tribe and its language, the town
and the region, and the Ocala Limestone.
FIGURE 6—Ocalaster seloyi n. gen. and sp., holotype, Florida Museum of Natural History UF 17244, Lower Ocala Limestone, Florida. Scale bars all
5 mm. 1, Dorsal view of entire specimen; 2–7, abradial toward bottom of photograph; 2, interior of SM series, with abactinals to top; 3–6, dorsal of SM,
abactinals with accessories, accessory depressions; 7, at top, MAO and ambulacral series above actinals; actinal interior surfaces are smooth; to left,
abradial of ambulacrals and carinals, a few abactinals with medial spines, accessories; 8, at top, carinal series with facets, spines; circumoral,
foreshortened ambulacrals to middle left; also interbrachial abactinals with fine accessories, only small facets.
BLAKE AND PORTELL—FLORIDA EOCENE ASTEROIDS 571
Discussion.—Two species of Ocalaster n. gen. are recog-
nized, both based on partial specimens, both with only a part
of the disk and arm bases preserved. The dorsal surfaces of
both specimens are exposed and as a result, the best available
information is for the superomarginals and abactinals with
only limited data on inferomarginals, ambulacrals, and
actinals; adambulacrals and the jaw frame are unavailable.
Primary ossicular form is well preserved, but data on
accessories are incomplete.
Overall form and marginal and abactinal morphology allow
assignment to the Goniasteridae. Abactinal form is very
similar between the two specimens. Abactinal expression has
not received detailed emphasis in the systematics of either
living or fossil goniasterids, and similar patterns are found in
many living genera. Marginals, especially inferomarginals, are
incompletely known; nevertheless, the two specimens are
considered similar enough to be assigned to a single genus but
different enough to indicate separate species, see below.
Ocalaster shares much with extant Nymphaster, including
both the type species, N. protetus Sladen, 1889 (SD, Fisher,
1917) and the living Florida species N. arenatus (Perrier, 1881);
similarities include presence of a large disk with broadly
curved interbrachial arcs, presence of low, tabulate abactinals,
the expression of the superomarginals, and presence of
granular accessories. Diagnoses of Nymphaster (Fisher, 1919;
Spencer and Wright, 1966; Halpern, 1970; Gale, 1987; Breton,
1992; Clark and Downey, 1992) all cite presence of contiguous
superomarginals on the arms, and most also cite presence of
an angular furrow margin on the adambulacral ossicles;
neither feature is exposed in the available material, although
convergence of the marginal series at the margins of the disk
suggest slender, extended arms and perhaps abutted marginals.
Both Breton (1992) and especially Gale (1987, fig. 3) recognized
a broad range of variation of marginal ossicles within fossil
species assigned to Nymphaster, again in effect challenging the
significance of isolated ossicles, and for Nymphaster,emphasiz-
ing the importance of arm and adambulacral form as well as the
need for a multielemental taxonomy. Ocalaster is based on
abactinal expression together with aspects of marginal and
accessory form; however, generic assessment will need to be
reconsidered with discovery of new material and further
documentation of recognized fossil species.
Separation of Ocalaster n. gen. from Oyenaster n. gen. is
based on overall form, differences of superomarginal form,
and accessory development. Abactinals are poorly known in
Oyenaster but appear higher, and enlarged carinals with
prominent basal facets have not been recognized. Among
other similar taxa, marginals of broadly similar form but
distinctive accessories, as further noted under Oyenaster
discussion, are found in Crateraster, Tomidiaster and Tylas-
teria. The possibly plesiomorphic Comptoniaster is similar, but
differs in accessory style and absence of marginal adradial
flanges. Caletaster also is similar in many of the same features,
but accessory development differs and marginal form is
‘‘twisted’’ (Breton, 1992).
O
CALASTER TIMUCUM, n. sp.
Figure 5
Diagnosis.—Ocalaster with small pedicellariae alveolae on
some abactinals, possibly concentrated near ambulacra;
abactinal spines occur. Superomarginal profile low, adradial
margin rounded; adradial terminus appears to extend beyond
that of IM. Shallow peripheral furrow zone probably present
between marginal series. One or two transversely aligned
circular spine bases on some SM, most bases near transverse
ossicular midline and near abradial edge but bases can be
more adradial, offset from midline. IM side face outline nearly
rectangular. Side face-adradial face edge with distinct facet
dorsally.
Discussion.—Arm bases and marginal form suggest elongate
arms. Abactinal series adjacent to carinals might be slightly
enlarged relative to more lateral series, wider than long.
Etymology.—Named in honor of the Timucua Native
American tribe.
Material and occurrence.—Holotype UF 20565, one silicone
rubber peel made from external mold. Peel later cut into two
unequal pieces to allow for easier examination. Largest piece
consisting of most of the disk and bases of two arms and a
portion of a third arm, exposed in dorsal view; interbrachial
radius 45 mm–50 mm; superomarginals, a small portion of a
few inferomarginals, and abactinals are exposed. Smaller piece
includes only a few poorly preserved marginals.
Dolime Quarry 01 (FLMNH IP CI009), Citrus County,
Florida. Yankeetown Quadrangle, USGS 7.59, 1988; T17S,
R16E. Bed 2, Periarchus Zone, Lower Ocala Limestone, upper
Eocene.
O
CALASTER SELOYI, n. sp.
Figure 6
Diagnosis.—Ocalaster with small medial spines present on
numerous abactinals; abactinal pedicellariae alveolae possibly
present but not identified with certainty. SM profile quite
high, adradial margin straight to weakly curved; spine bases
unknown. SM not appearing wider than IM; adradial face
appearing bordered by ridge rather than simple facet; (only
adradial faces of IM available). Adradial face facets well
defined. Intermarginal contact unknown. Actinals similar in
size and proportion to abactinals (only interiors exposed,
overlain by abactinals). Circumorals of conventional gonias-
terid form; robust, cross furrow articular depression strong,
ossicle constricted immediately abradial to articular surfaces.
Etymology.—Named for Seloy, a Timucua chief who on
September 8, 1565, watched the Spanish fleet anchor offshore.
Material and occurrence.—Holotype UF 17244, one silicone
peel made from an external mold consisting of a portion of the
disk, superomarginals on one interbrachial arc leading toward
two arms, but the arms are not preserved. Also, abactinals,
some ambulacrals, and one circumoral are exposed, all in
dorsal view. Occurrence as for O. timucum.
ACKNOWLEDGMENTS
The authors are indebted to C. W. Oyen (1963–2005), who
found the holotype of Oyenaster and brought it to the
attention of RWP at the Florida Museum of Natural History.
Field assistance at the Dolime Quarry 01 (FLMNH IP CI009)
was provided by J. Bryan and D. Kendrick; J. T. Milanich
advised on the Timucua. Figures 1 and 2 were drafted by S.
Roberts. A preliminary review was provided by J. Jagt, and C.
Mah, and L. Villier both read early drafts and provided
reviews for the Journal; C. Sumrall served as Associate Editor.
All are thanked for their helpful contributions. This is
University of Florida Contribution to Paleobiology 592.
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