Journal of Systematic Palaeontology 5 (2): 163–192 Issued 25 May 2007

doi:10.1017/S1477201906002021 Printed in the United Kingdom

C The Natural History Museum

Hemipatagus, a misinterpreted
Loveniid (Echinodermata:
Echinoidea)
Andreas Kroh∗
Naturhistorisches Museum Wien, Burgring 7, A-1010 Wien, Austria

SYNOPSIS The echinoid genus Hemipatagus is a poorly understood fossil spatangoid taxon that
is now usually treated as a subjective synonym of the extant genus Maretia, but was originally
subject to considerable dispute within the scientific community. Restudy of the species attributed
to Hemipatagus and a range of presumably related spatangoids including Lovenia and Maretia has
been carried out to solve the problem of its relationships. Cladistic analysis shows that Hemipatagus
is not close to Maretia, but is closely related to Lovenia and should be placed in the Loveniidae,
which is here confirmed as a monophyletic group. Characters of the adapical tuberculation suggest
that in Hemipatagus an internal fasciole is present in early ontogeny, but lost in the adults. A clade
comprising the genera Eurypatagus, Paramaretia and Platybrissus, and here named Eurypataginae,
appears in all resulting trees as sister group to Maretia.

KEY WORDS Echinoidea, Hemipatagus, Loveniidae, Cenozoic, Eurypataginae nov.

Contents
Introduction 163
History of nomenclature 164
Material and methods 164
Abbreviations and repositories 167
Results 167
Status of Hemipatagus 167
Phylogenetic affinities of Hemipatagus 169
Cladistic analysis 171
Spatial and temporal distribution of Hemipatagus and Maretia 172
Systematic Palaeontology 173
Order Spatangoida Claus, 1876 173
Family Loveniidae Lambert, 1905 173
Genus Hemipatagus Desor, 1858 173
Family Spatangidae Gray, 1825 174
Subfamily Eurypataginae nov. 174
Acknowledgements 174
References 174
Appendix 1 Characters scored in the cladistic analysis 178
Appendix 2 Data matrix 179
Appendix 3 Annotated list of the nominal species of Hemipatagus and Maretia 180

Introduction exploiting organic detritus as their prime food resource.

Spatangoids are echinoids characterised by their bilaterally
symmetrical corona and highly modified ambulacra form-
ing a complex respiratory apparatus. They are well adapted
to a life as burrowers and ploughers in mobile sediments,
* Corresponding author. E-mail address: andreas.kroh@nhm.wien.ac.at
Due to their life habit and evolutionary success they are
the most diverse of all extant sea urchin groups and one of
the echinoid clades with the best fossil record (Kier 1977).
Their highly adapted test morphology provides many fea-
tures that can be employed in classification. Consequently
spatangoid taxonomy is, in general, rather straightforward.
164 A. Kroh
Particular features, however, have recently been shown
to display high levels of convergence and homoplasy
(Markov & Solovjev 1995; Neraudeau et al. 1998; Villier
et al. 2004; Smith & Stockley 2005). Indeed, spatangoid
phylogeny and higher-rank taxonomy has been heavily re-
liant on fascioles. These are narrow bands of small cili-
ate spines secreting mucus that consolidates the walls of
their burrows and protects the respiratory tube feet from fine
mud (see Lawrence 1987). Unfortunately, fascioles were de-
veloped several times during spatangoid evolution and can
be secondarily lost (Villier et al. 2004; Smith & Stockley
2005) and the emphasis placed on these structures has led to
considerable confusion and problems in spatangoid system-
atics. The problem investigated here represents a case where
too much emphasis has been placed on presence and absence
of fascioles.
Hemipatagus (Pl. 1) is a genus widely distributed in the
Eocene to Miocene of Europe, occurring abundantly in the
famous Doberg section in northern Germany (Oligocene)
and in Early Miocene strata of the Rhône Basin (Philippe
1998), the Aquitaine Basin and the central Paratethys (Kroh
2005). It is a spatangoid characterised by its heart-shaped
outline and heterogeneous aboral tuberculation. The phylo-
genetic and systematic relationship of Hemipatagus within
the spatangoids is uncertain, although it has commonly been
synonymised with Maretia (Pl. 2, figs 1 & 2), a Pliocene to
extant inhabitant of the Indo-West Pacific. The complicated
nomenclatorial history of both genera is outlined below.
In this paper, the relationship between Hemipatagus,
Lovenia and Maretia is investigated using a cladistic ap-
proach based on morphological analysis. This generates a
phylogeny and revised taxonomy for Hemipatagus and its
relatives.
History of nomenclature
The genus Hemipatagus was established in 1858 by Desor
for the species Spatangus hoffmanni Goldfuss, 1829 (Fig. 1).
Subsequently, Agassiz (1873: 568) placed Hemipatagus in
synonymy with Maretia. In the following years this place-
ment was subject to some debate. Some echinologists suppor-
ted Agassiz’s view (Cotteau 1885: 24–25; Fourtau 1920: 83),
while others disagreed (Duncan 1877: 56–58, 1889: 222, 252;
Lambert 1909: 107, 1915: 188, 1927a: 87). In his ‘A Mono-
graph of the Echinoidea’ Mortensen (1951: 23–27) discussed
this matter at length and finally considered Hemipatagus
Desor, 1858 a junior synonym of Maretia Gray, 1855. His
opinion was based on the observation that only one of the
three features discussed in the literature seemed important
to him (the presence of internal ampullae associated with
spine tubercles in Hemipatagus). He argued that one feature
alone was not enough to justify separation of the two gen-
era. Moreover, he had also observed internal swellings that
he assumed were ampullae in juveniles of Maretia planu-
lata (Lamarck, 1816) and saw them as proof for his point of
view. In addition, Mortensen noted that the presence of am-
pullae was not verified in the type species of Hemipatagus.
The other two features listed by him were: (1) a more heart-
shaped outline with deeper frontal notch (he disregarded this
feature, due to the fact that large differences in the depth of the
frontal notch are also present in other genera, e.g. in Lovenia);
(2) the broader, bilobed subanal fasciole in Hemipatagus (he
also disregarded this feature, although noted that it is im-
portant for generic classification of some brissids). In the
‘Treatise on Invertebrate Paleontology’ Fischer (1966: U609)
adopted Mortensen’s view and thus it was soon widely accep-
ted in the scientific community, although some authors (e.g.
Philippe 1998: 222; Kroh & Harzhauser 1999: 163) still con-
tinued to express their doubts. Smith (2004: ‘The Echinoid
Directory’) noted the high similarity between Hemipatagus
and Lovenia and proposed its inclusion in the Loveniidae.
Part of the confusion surrounding Hemipatagus and
its relationship to Maretia and Lovenia resulted from
Desor’s (1858) statement that no fascioles are present in
Hemipatagus. Laube (1869) and Etheridge (1875), however,
showed that a ‘spectacle-shaped’ (bilobed) subanal fasciole
is present (see Fig. 2). Another major reason for the differing
opinions resulted from the habit of basing conclusions on fea-
tures observed in various non-type species attributed to either
genus (e.g. Duncan 1877). The type species of Hemipatagus
and Maretia, in contrast, were rarely investigated with regard
to this question. Furthermore, most authors placed a heavy
emphasis on the presence or absence of an internal fasciole,
a feature which is often not readily observed in fossil speci-
mens. Moreover, it is known now that fascioles may be lost
during ontogeny, being present in juveniles and absent in the
adults of many spatangoids (e.g. Meoma (partial loss of the
subanal fasciole: Kier & Grant 1965; Chesher 1969), Para-
maretia, Platybrissus and Eurypatagus (complete loss of the
subanal fasciole: Mortensen 1950, 1951); see also Néraudeau
et al. 1998; Smith & Stockley 2005).
Although discussed at length it is not really clear why
Mortensen (1951: 23–27) finally chose to synonymise Hemi-
patagus with Maretia. It would have been understandable if
he had synonymised Hemipatagus with Lovenia, as he stated
himself on p. 27 that ‘On the whole, the distinction between
Maretia [here actually referring to Hemipatagus] and Love-
nia is, in case of the fossils, very difficult, the presence or
absence of an inner fasciole – the only fully reliable distinc-
tion between these two genera – being, rather impossible to
ascertain beyond doubt, if the preservation is not very fine.’ It
is difficult to understand why he rejected the shape of the sub-
anal fasciole as a diagnostic feature in Hemipatagus, while
widely using it in brissid genera, particularly, as this feature
allows a very clear separation, with no known intermediate
species. Mortensen (1951) also made a strong case against
the use of the outline and the depth of the frontal sinus,
arguing that he would be forced to split Lovenia into sev-
eral genera when using these features to distinguish between
Hemipatagus and Maretia. Yet, it is well known that features
that are useful to separate some taxa might be highly variable
in others, not allowing a clear separation.
Material and methods
Two specimens of the extant Maretia planulata (Lamarck,
1816) from the Gulf of Siam, Mauritius (NHMW, 2nd
Zoological Department, collection Invertebrata-Varia no.
12715), three specimens of Lovenia elongata (Gray, 1845)
from the Red Sea (NHMW, 2nd Zoological Department, col-
lection Invertebrata-Varia no. 12714) and 24 well preserved
specimens of Hemipatagus hoffmanni (Goldfuss, 1829)
from the Oligocene of the type locality Doberg near Bünde,
 Hemipatagus – a misinterpreted Loveniid 165

Plate 1 Oligocene and Miocene Hemipatagus. Figs 1–10, 16, Hemipatagus hoffmanni (Goldfuss, 1829), all Late Oligocene of Doberg, near
Bünde, Germany. 1–3, MB.E4843; 4–7, NHMW 1852.II.1500a; 8–9, MB.E4839; 10, 16, NHMW 1855.I.801a. Figs 11–14, Hemipatagus ocellatus
(Defrance, 1827), all Late Eggenburgian (late Early Burdigalian, Early Miocene) of Grübern, Austria. 11–12, NHMW 1914.VII.22b; 13–14,
1914.VII.22a. Figs 15, Hemipatagus martensii (Ebert, 1889), Late Oligocene of Doberg, near Bünde, Germany, MB.E4840.
166 A. Kroh

Plate 2 Comparison between similar-sized specimens of Hemipatagus, Lovenia and Maretia. Figs 1–2, Maretia planulata (Lamarck, 1816),
extant, Gulf of Siam, Mauritius, NHMW, 2nd Zoological Department, collection Invertebrata-Varia no. 12715. Figs 3–4, Lovenia elongata (Gray,
1845), extant, Red Sea, NHMW, 2nd Zoological Department, collection Invertebrata-Varia no. 12714. Figs 5–6, Hemipatagus martensii (Ebert,
1889), Late Oligocene of Doberg, near Bünde, Germany, MB.E4840.
 Hemipatagus – a misinterpreted Loveniid 167

Figure 1 Hemipatagus hoffmanni (Goldfuss, 1829), syntype and figured specimen of Goldfuss (1829); specimen PIUB 357a. A, aboral view;
B, right lateral view; C, oral view.

Germany (NHMW, Geological-Palaeontological Depart-
ment 1852.II.1500, 1852.II,1501, 1855.I.800, 1855.I.801,
1862.XLIII.43, 1874.XXIX.1178 and MB, Palaeontology
Department MB.4839–4843, plus 6 unregistered speci-
mens) were examined. In addition, the type-material of
Hemipatagus hoffmanni was located (PIUB 357a-h) and
a photograph of Goldfuss’s original specimen (PIUB
357a) is presented for the first time (Fig. 1). Furthermore,
the type material of Hemipatagus perornatus (Schaffer
1912:2 syntypes – KM F/0127 and F/0128) was located and
reinvestigated, along with some new specimens (NHMW
Geology-Palaeontology Department 1914.VII.22a-c,
1999z0049/0003, 2002z0159/0001; see also Kroh 2005).
Cladistic analysis was carried out using the programme
PARS from the Phylogeny Inference Package PHYLIP
(Felsenstein 2002), using the ‘Randomize input order of
species option’ (1000 replicates). The program CONSENS
from the same package was used to obtain majority rule
consensus trees. The analysis involved 30 species from the
Spatangidae and Loveniidae together with Eupatagus valen-
ciennesi Agassiz, in Agassiz & Desor, 1847. A total of 50
characters were scored based on test shape, apical disc struc-
ture, plating pattern, tuberculation, sphaeridia, pore and peri-
proct structure. A number of additional, phylogenetically
uninformative characters (basal synapomorphies) as well as
some highly variable characters were excluded. The data
matrix and description of the remaining 42 characters are
found in Appendices 1 and 2. Due to the high degree of
morphological differentiation amongst the taxa investigated
many characters were multi-state. Characters have neither
been polarised nor differentially weighted.
Abbreviations and repositories
GSI, Geological Survey of India; KM, Krahuletz Museum,
Eggenburg, Austria; NHMW, Naturhistorisches Museum
Wien, Austria; MB, Naturkunde Museum Berlin, Germany;
PIUB, Institute of Palaeontology, University of Bonn,
Germany (Goldfuss Museum).
Results
Status of Hemipatagus
Material from the type species of Hemipatagus, Lovenia and
Maretia was investigated in detail, focusing on tubercula-
tion types and arrangement, gross morphology, pore morpho-
logy, fascioles (Fig. 2) and, most importantly, plating patterns
(Figs 3 & 4, Table 1).
When comparing H. hoffmanni with M. planulata, the
following differences were observed in M. planulata: (1) the
frontal sinus is shallower and often indistinct; (2) the lab-
rum is much narrower and more elongated; (3) the posterior

Figure 2 Oblique view of the subanal fasciole. A, Hemipatagus hoffmanni (Goldfuss, 1829), NHMW 1855.I.801a, Late Oligocene, Doberg,
Germany; B, Hemipatagus ocellatus (Defrance, 1827) NHMW 1914.VII.22b, Early Miocene, Grübern, Austria.
168 A. Kroh

Figure 3 Camera lucida drawings of oral plating. A, Hemipatagus hoffmanni (Goldfuss, 1829) NHMW 1855.I.801a, Late Oligocene, Doberg,
Germany; B, Hemipatagus ocellatus (Defrance, 1827) NHMW 1914.VII.22b, Early Miocene, Grübern, Austria. Interambulacra shaded, fasciole
finely stippled, small stippled circles are sphaeridial pits.

end is bluntly pointed not transversely truncated; (4) even in
large specimens the primary tubercles do not develop true
ampullae or strongly sunken areoles, especially not on the
oral surface; (5) the plastron is much more strongly con-
stricted between the first and second pair of episternal plates
(Fig. 4A); (6) the outline of the test is oval, antero-posteriorly
elongated, not cordiform; (7) the subanal fasciole is oval and
rather high instead of bilobed, very wide and low (Table
1); (8) the labrum and the primordial plates of the interam-
bulacra extend much further (Fig. 4A); (9) there are more
phyllodal pores (14–15) in ambulacra II and IV (only 6–7
in H. hoffmanni); (10) ambulacra II and IV are only slightly
constricted ambitally, whereas they are strongly constricted
in H. hoffmanni (compare Figs 3 and 4A); (11) a conspicuous
field of coarse tubercles adjacent to adapical ambulacrum III
is well developed in H. hoffmanni but lacking in M. planu-
lata (Pl. 3, figs 1–5). These differences provide ample means
to separate those two genera. Lovenia and Hemipatagus are
more difficult to separate. Hemipatagus differs mainly by the
absence of the internal fasciole and its different petal struc-
ture (never forming continuous arcs as in most species of
Lovenia).
When other species (e.g. H. ocellatus, H. grignonensis,
H. girundicus, M. planulata pliocenica, M. ? cordata: see Ap-
pendix 3) attributed to the three genera in question are taken
into account, it becomes apparent that most of the features

Figure 4 Camera lucida drawings of the oral plating. A, Maretia planulata (Lamarck, 1816) NHMW-EV 12715, Recent, Gulf of Siam, Mauritius;
B, Lovenia elongata (Gray, 1845) NHMW-EV 12714, Recent, Red Sea. Interambulacra shaded, fasciole finely stippled, small stippled circles are
sphaeridial pits.
 Hemipatagus – a misinterpreted Loveniid
Table 1 Summary of the differential diagnostic features of Hemipatagus and related respectively morphologically similar genera.
Amb, ambulacrum.
outlined above allow a clear separation of the two groups.
Concerning the structure of the primary tubercles, however,
Mortensen (1951) was quite correct to question their value
as a diagnostic feature. While some species of Hemipatagus
have truly camellate tubercles (i.e. tubercles with associated
pouch-like ampullae forming a bulge on the interior of
the test) on the oral and aboral surface (e.g. H. ocellatus,
H. perornatus, H. girundicus), others lack them either over
the oral surface (e.g. H. hoffmanni) or completely (H. carolin-
ensis, H. argutus). In those species where internal ampullae
are lacking, the areoles are, nevertheless, strongly sunken.
However, the other features, especially the distribution of the
aboral primary tubercles, number of phyllodal pores and all
features related to the plating pattern, are quite stable and
allow a consistent separation (see ‘Emended diagnosis’ and
‘Remarks’, below, for details).
Phylogenetic affinities of Hemipatagus
The systematic placement of Hemipatagus remains difficult.
Most spatangoid families are based on the type of primary
tuberculation, differentiation of fascioles, structure of the ap-
ical disc and plastron structure (see Fischer 1966). As pointed
169
out by various authors recently (e.g. Jeffery 1998; Néraudeau
et al. 1998; Smith & Jeffery 2000), in the past too much em-
phasis was placed on the presence/absence of certain types
of fascioles in the adult animal in developing a supra-generic
classification of the spatangoids. It has been clearly estab-
lished that fascioles may be lost during ontogeny, a fact that
has to be taken into account when trying to resolve this
question (e.g. Mortensen 1951; Kier & Grant 1965; Chesher
1969).
Here the conspicuous field of tubercles in adapical in-
terambulacral columns 2b and 3a (Pl. 3, figs 2–5) comes
into play. In all genera (except Hemipatagus) where this fea-
ture is present it is invariably associated with an internal
fasciole (Breynia, Echinocardium, Lovenia, Pseudolovenia
and Vasconaster). It is, therefore, possible to assume that
an internal fasciole was present during the early ontogeny
of Hemipatagus. The fact that this arrangement is not well
developed in some species of Hemipatagus does not neces-
sarily mean that these species never possessed an internal
fasciole. There are some species of Lovenia in which this
special arrangement is not very conspicuous (e.g. L. sub-
carinata). By contrast, such an arrangement has never been
observed in any member of the family Spatangidae (including
Maretia).
170 A. Kroh

Plate 3 Adapical tuberculation and phyllodes of Hemipatagus, Lovenia and Maretia. Figs 1 & 9, Maretia planulata (Lamarck, 1816), extant,
Gulf of Siam, Mauritius, NHMW, 2nd Zoological Department, collection Invertebrata-Varia no. 12715. Figs 2 & 8, Lovenia elongata (Gray, 1845),
extant, Red Sea, NHMW, 2nd Zoological Department, collection Invertebrata-Varia no. 12714. Figs 3–4, 6, Hemipatagus hoffmanni (Goldfuss,
1829), all Late Oligocene of Doberg, near Bünde, Germany. 3, MB.E4839; 4, MB.E4843; 6, NHMW 1855.I.801a. Figs 5 & 7, Hemipatagus ocellatus
(Defrance, 1827), all Late Eggenburgian (late Early Burdigalian, Early Miocene) of Grübern, Austria. 5, 1914.VII.22a; 7, NHMW 1914.VII.22b.
 Hemipatagus – a misinterpreted Loveniid 171

Figure 5 Result of the cladistic analysis based on the full dataset (A) and the reduced dataset (B). Italic numbers beside the nodes indicate
bootstrap values obtained (1000 replicates). Numbers in normal font beside the nodes in the 50% majority consensus tree (A) indicate the
percentage of times that this node occurred in the full set of most parsimonious trees.

On the basis of its strong similarity to Lovenia and the
inferred loss of the internal fasciole during early ontogeny,
Hemipatagus is here placed in the family Loveniidae.
Cladistic analysis
To test the inferences based on the subjective evaluation
of the morphological features of Hemipatagus, Lovenia and
Maretia, a cladistic analysis was carried out. In all analyses
Hemipatagus hoffmanni and H. ocellatus form a sister group
taxon to the extant species of Lovenia. Exclusion of the spe-
cies with a high number of unknown characters (Maretia
carinata, Hemimaretia elevata, Hemipatagus? subrostratus)
results in a very stable tree topology with a deep split
between a clade containing Eurypatagus, Maretia, Para-
maretia, Platybrissus and Spatangus (clade A) and a second
clade containing the classical Loveniidae (Breynia, Ech-
inocardium, Homolampas, Lovenia, Pseudolovenia) plus
Hemipatagus and Pseudomaretia (the latter traditionally
placed in the Spatangidae: see Fischer 1966) (clade B). Due to
the presence of several missing characters in Maretia cordata
and two old species tentatively placed in Hemipatagus (H.?
argutus and H.? carolinus, from the Middle Eocene and the
Oligocene, respectively) 30 equally parsimonious trees were
found. The majority consensus tree of these trees is shown
in Fig. 5A. The placement of M. cordata switches between
just two positions (at the base of clade A versus as sister-
group taxon to the two spatangids). Hemipatagus? argu-
tus and H.? carolinus, in contrast, are resolved either as
172 A. Kroh
Figure 6 Spatial distribution of the genera Hemipatagus and Maretia. Open symbols show taxa of doubtful generic affinity. Distribution of
extant and fossil M. planulata is based on data in: Stockley 1927; Brighton 1931; Mortensen 1951; Richer de Forges & Menou 1988; Baker &
Rowe 1990; Kerr et al. 1992; Liao & Clark 1995; Rowe & Gates 1995, USSMDC 1999; Lindley 2003.
stem-group loveniids (in the majority of the trees) or as
crown-group loveniids forming a polyphyletic clade with
Pseudolovenia, H. hoffmanni, H. ocellatus and the extant
species Lovenia. In just one of the 30 trees H.? argutus and
H.? carolinus are resolved as sister group to Echinocardium.
Exclusion of M.? cordata, H.? argutus and H.? carolinus
from the analysis results in a single most parsimonious tree
(Fig. 5B) very similar to the majority consensus tree of the
former analysis but with better resolution within clade B. It
is apparent that the loveniids as a whole are a very robust
group with well-defined monophyletic genera. The only ex-
ception is Lovenia, which is monophyletic, but paraphyletic
if Pseudolovenia is excluded. The present analysis suggests
that Pseudolovenia should either be synonymised with Love-
nia or that Lovenia should be split into two different genera.
The topology of the Lovenia clade resulting from the present
analysis is very similar to that published by Madon (1994)
using a different dataset. Another new result is the placement
of the Pseudomaretia-clade in the loveniids. Earlier Pseudo-
maretia (and the junior synonym Goniomaretia) were con-
sidered to be spatangids (Fischer 1966) or maretiids. The
genera Eurypatagus, Platybrissus and Paramaretia form a
distinct clade in the present analysis and are a sister group to
Maretia planulata in most trees. This clade appears also in
the strict consensus tree published by Stockley et al. (2005:
fig. 2), where it also forms the sister group of Maretia. The
Eurypatagus–Platybrissus–Paramaretia-clade is here named
Eurypataginae (see ‘Systematic palaeontology’ below, for a
formal definition).
Inclusion of a more distant outgroup (Brissus unicolor)
results in a distinctly higher number of most parsimonious
trees (42), the majority consensus trees of which show a very
similar topology to the trees discussed above.
The present analysis clearly shows that Hemipatagus
is a loveniid that evolved during the Middle or Late Eocene
from a common ancestor with Lovenia. Despite the superfi-
cial similarities with Maretia it is clearly not closely related
to that genus. This is also reflected by their temporal and
spatial distributions, which are disjunct with respect to each
other (see below). Earlier statements that Maretia and Hemi-
patagus are synonymous (e.g. Mortensen 1951) were based
on misinterpretation and erroneous weighting of a small set
of characters.
In a recent paper on spatangoid phylogeny, Stockley
et al. (2005) present the results of a cladistic analysis in-
volving several of the genera discussed here. In the strict
consensus tree resulting from their morphological analysis
(Stockley et al. 2005: fig. 2) the Loveniidae are a poly-
phyletic clade. According to co-author A.B. Smith (pers.
comm., 01.11.2005), however, their analysis produced a very
similar topology to the one presented here when restricted
to the micrasterine clade alone. Smith relates the differences
between their fig. 2 and the present tree (Fig. 5) to the global
parsimony analysis employed in Stockley et al. (2005), which
fails to group the loveniid taxa into a single clade.
Spatial and temporal distribution of Hemipatagus
and Maretia
Hemipatagus and Maretia are well separated in time and
space. The former occurs most commonly in the Eocene to
 Hemipatagus – a misinterpreted Loveniid
Middle Miocene of the Mediterranean region. In addition,
there are a few species on the east coast of North America
(see Fig. 6). Maretia, by contrast, occurs first in the Plio-
cene and is restricted to the Indian Ocean, the Indo–Malayan
Archipelago and the West Pacific Islands (farthest east
occurrences are on Hawaii). The only potential record of
Hemipatagus from the Pliocene and from the Indo–West Pa-
cific region is H.? bandaensis. The data available for this
taxon are not sufficient to firmly establish its generic affinity.
It might also be a misidentified Lovenia. A re-description and
new illustrations are urgently needed.
Another issue apparent in Fig. 6 is the disjunct distri-
butional pattern of the extant Maretia planulata. Apparently
there are no records from Western India, Pakistan, the Persian
Gulf or the Indian Ocean coast of the Arabian Peninsula. In
view of this the colouration differences noted by Mortensen
(1951: 36) may be more meaningful than he considered they
were. He described a whitish to yellow colouration in Indian
Ocean populations, versus Malayan specimens, which are
usually dark. As Mortensen (1951: 36) could find no apparent
morphological differences, he considered this phenomenon
to be local variation. Further investigations are needed to
judge whether there might actually be two species. A sim-
ilar situation has recently been discovered in Asthenosoma
by Weinberg & de Ridder (1998), who discovered that the
echinoids previously classified as A. varium belong to two
distinct species, which are geographically separated.
Systematic Palaeontology
Order SPATANGOIDA Claus, 1876
Family LOVENIIDAE Lambert, 1905
Genus HEMIPATAGUS Desor, 1858
TYPE SPECIES. Spatangus hoffmanni Goldfuss, 1829; by ori-
ginal designation (Desor 1858: 416). (nomen correctum pro
Spatangus hofmanni Goldfuss, 1829 – Grateloup (1836: 175)
(presumably intentionally) changed the name to S. hoffmanni.
According to the ICZN rules this is an unjustified emenda-
tion. However, as the name S. hoffmanni has been in prevail-
ing usage since 1836 and is attributed to the original author
and date of the original spelling (e.g. des Moulins 1837:
244; Agassiz & Desor 1847: 7), it has to be deemed to be a
justified emendation according to the ICZN (1999: 4th edn
Article 33.2.3.1.)).
RANGE. Eocene to Middle Miocene (Serravallian).
EMENDED DIAGNOSIS. Cordiform spatangoid differing from
other loveniids by its (1) heterogeneous aboral tubercula-
tion with large crenulate or non-crenulate primary tubercles
(Table 1); (2) conspicuous aboral primary tubercles in inter-
ambulacra 1–4, but missing in interambulacrum 5 and the
posterior part of interambulacral columns 1a and 4b (see
Table 1); (3) strongly sunken areoles or even the presence of
camellae in some species; (4) presence of a subanal fasciole
in adult specimens, which is strongly bilobed (Figs 2A, B);
(5) presence of a conspicuous field of coarse tubercles adja-
cent to adapical ambulacrum III in interambulacral columns
2b and 3a (Pl. 3, figs 3–5); (6) strong constriction of ambu-
lacra II and IV near the margin on the oral side in adult speci-
173
mens (Figs 3A, B); (7) moderately elongated labral plate not
extending beyond the second ambulacral plates (Figs 3A, B);
(8) prominent oral tubercles arranged in distinct rows, in
some species with deeply sunken areoles and helically spir-
alled parapet (e.g. Pl. 3, fig. 7); (9) three to four ambulacral
plates extending into the subanal fasciole in ambulacrum I
and V, respectively.
INCLUDED SPECIES. Apart from the type species H. hoff-
manni (Goldfuss, 1829) the following species can be con-
fidently assigned to Hemipatagus based on the published
information: H. calvimontanus (Cotteau, 1885); H. car-
olinensis (Kier, 1997); H. desmoulinsi (Cotteau, 1863); H.
girundicus Lambert, 1915; H. guebhardi (Lambert & Savin,
1902); H. hungarica (Vadsz, 1915); H. marianii (Airaghi,
1903); H. martensii (Ebert, 1889); H. ocellatus (Defrance,
1827) (=H. nicoleti (Agassiz, 1839) = H. perornata (Schaf-
fer, 1912)); H. pellati Cotteau 1863; H. saccoi (Airaghi,
1905); H. soubellensis (Peron & Gauthier, in Cotteau et al.,
1891); H. tenuis (Peron & Gauthier, in Cotteau et al., 1891).
In addition, there are several species that might also belong
to Hemipatagus, but which are too poorly known at present
to establish their generic affinities with certainty: H.? ar-
gutus Clark, in Clark & Twitchell, 1915; H.? bandaensis
Martin, in Jeannet & Martin, 1937; H.? carolinae (Gagel,
1905); H.? gauthieri (Lambert, in Lambert & Thiéry, 1924);
H.? gottschei (Gagel, 1905); H.? grignonensis (Cotteau,
1880); H.? hispanica (Cotteau, 1890); H.? mauritanica
(Pomel, 1885); H.? H.? regiomontanus (Mayer, 1860); H.?
sambiensis (Beyrich, 1848); H.? subrostratus Clark, in
Clark & Twitchell, 1915; H.? tuberculatus (Peron &
Gauthier, in Cotteau et al., 1885); H.? zeisei (Gagel, 1905).
See Appendix 3 for original generic assignments and more
data on the individual species listed here.
REMARKS. There are a number of genera that show superfi-
cial similarities to Hemipatagus. All of them, however, can
be clearly separated on the basis of structural features (see
Table 1). Lovenia differs by the presence of an internal fas-
ciole, high number of ambulacral plates extending into the
subanal fasciole (up to 11, although in some species there
are only three or four as in Hemipatagus) and less lanceol-
ate petals; in most species of Lovenia ambulacral columns
Ib–IIa and IVb–Va form continuous arcs (not observed in
any known Hemipatagus species); in some species of Hemi-
patagus the aboral and/or oral tubercles are not camellate,
while this is always the case in Lovenia (see also Fig. 4B; Pl.
2, figs 3–4; Pl. 3, fig. 8).
Maretia differs by possessing an extremely shallow
frontal notch, the distribution of the aboral primary tubercles
(primary tubercle reach the tips of the posterior paired petals),
the lack of strongly sunken areoles or camellate aboral and
oral primary tubercles, the strongly crenulate nature of the
aboral primary tubercles, the lack of a conspicuous field
of coarse tubercles in adapical interambulacral columns 2b
and 3a, its oval to shield-shaped subanal fasciole, the longer
primordial plates of the paired interambulacra (plates 1a1,
2a1, 3a1 and 4a1: which extend to the third or fourth am-
bulacral plate of the adjacent ambulacra in Maretia, while
they do not extend beyond the second ambulacral plate in
Hemipatagus or Lovenia), the labrum which extends to the
third or fourth ambulacral plate (not extending beyond the
second ambulacral plate in Hemipatagus and Lovenia) and
174 A. Kroh
the less constricted adoral ambulacra 2 and 3 (Fig. 4A; Pl. 2,
figs 1–2).
Breynia differs by its fully tuberculate sternum, pres-
ence of an internal and a peripetalous fasciole, shield-shaped
form of the subanal fasciole, shape of the petals (with am-
bulacral columns Ib–IIa and IVb–Va forming a continuous
arc in some species), by the labrum and primordial plates of
the paired interambulacra, which extend to the third ambu-
lacral plate, the presence of primary tubercles on the primor-
dial plates of the paired interambulacra; by the smaller oral
primary tubercles, which are never sunken or camellate; and
the high number of ambulacral plates extending into the sub-
anal fasciole (usually six to eight), by the distribution of the
aboral primary tubercles (confined to the area within the peri-
petalous fasciole, but reaching the tips of the posterior paired
petals) and the near-straight contact between the sternal and
episternal plates.
Eupatagus differs by its fully tuberculate sternum, pres-
ence of a peripetalous fasciole, oval shape of the subanal
fasciole, the different nature of the aboral and oral primary
tubercles (never camellate or strongly sunken), different dis-
tribution of the aboral primary tubercles (see Table 1), by the
labrum and primordial plates of the paired interambulacra
which extend to the third ambulacral plate and by the lack of
any frontal notch.
Pseudolovenia differs by its narrow, poorly-defined
petals with widely spaced, less well developed pores which
continue to the ambitus, restriction of the aboral primary
tubercles to the interambulacral columns adjacent to ambu-
lacra II and IV, its labrum which extends to the third am-
bulacral plates, the presence of an internal fasciole and the
higher number of plates extending into the subanal fasciole
(five).
Pseudomaretia and Goniomaretia differ by possessing
an ethmolytic apical disc with only three gonopores (no gono-
pore in genital plate 2), lack of a frontal notch, more reduced
nature of the anterior poriferous zones of the anterior paired
petals (well developed pores only in the most distal plates),
oval to shield-shaped subanal fasciole and the absence of
most or all aboral primary tubercles.
Paramaretia differs by the more reduced nature of the
anterior poriferous zones of the anterior paired petals (no
large, well developed pores present at all), its labrum which
extends to the fourth ambulacral plates, the primordial plates
extend to the fourth to sixth ambulacral plates, the loss of the
subanal fasciole in adults, having a plastron which is only
weakly indented behind the first pair of episternal plates, its
naked sternal plates and, most obviously, by the distribu-
tion of the aboral primary tubercles which are present in all
interambulacra, without obvious pattern.
Family SPATANGIDAE Gray, 1825
Subfamily EURYPATAGINAE nov.
TYPE GENUS. Eurypatagus Mortensen, 1948: 133.
OTHER GENERA INCLUDED. Paramaretia Mortensen, 1950,
? Platybrissus Grube, 1866 (see Fig. 5).
RANGE. Miocene to Recent.
DIAGNOSIS. Differing from the Maretiinae by their (1) lack
of any fascioles in adult individuals, (2) naked sternal plates
(partially tuberculate in Maretiinae), (3) structure of the
petals (narrowed, distally open, extending almost to the am-
bitus) and (4) labrum extending to the 4th plates of ambulacra
I and V. Differing from the Spatanginae by their (1) strongly
elongated labrum, (2) naked sternal plates (fully tuberculate
in Spatanginae), (3) crowding of the adoral plates of ambu-
lacra II and IV and (4) the structure of the petals (narrowed,
distally open, extending almost to the ambitus).
REMARKS. Additional features that are characteristic, but
not exclusive, for this family are: (1) an oval outline with
only feeble frontal sinus if present at all; (2) heterogeneous
aboral tuberculation (enlarged primary tubercles developed
in most forms; where developed, they are arranged without
distinct pattern).
The genus Platybrissus is another potential member of
the Eurypataginae. In some analyses with a different out-
group (Brissus unicolor), however, this genus forms the sis-
ter group to Eurypatagus, Paramaretia and Maretia. There-
fore it is only tentatively included in the Eurypataginae at
present.
An independent cladistic analysis presented by
(Stockley et al. 2005: fig. 2) likewise shows a monophyletic
clade containing Eurypatagus, Paramaretia, and Platybris-
sus. As in most trees resulting from the present analysis,
Stockley et al. (2005) found a sister-group relationship
between this clade and Maretia.
Acknowledgements
This study was supported by the Austrian Science Fund
(FWF) via project no. P-13466-Bio to Werner E. Piller (Uni-
versity of Graz). The critical reviews and helpful comments
of Andrew B. Smith, Rich Mooi and another anonymous re-
viewer greatly improved the paper. Furthermore, I would like
to express my sincere thanks to the staff of the Geological De-
partment of the Natural History Museum Vienna for provid-
ing scientific facilities. Special thanks to Johannes Tuzar and
Andreas Thinschmidt (both Krahuletz Museum, Eggenburg)
for helping to locate the type-material of H. peronatus and to
Manuel Kunz, Martin Sanders and Georg Oleschinski (all at
the Goldfuss Museum, Bonn) for locating the type material
of H. hoffmanni and providing photographs of it. Thanks also
to Alice Schumacher (NHM Vienna) for taking the photo-
graphs.
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Appendix 1 Characters scored in the cladistic analysis

Shape
1. Test outline: cordate (0); oval (1).
2. Frontal sinus: absent (0); weakly developed (1); well developed (2).
Apical disc
3. Apical disc, position: anteriorly displaced (0); subcentral (1); posteriorly displaced (2).
4. Number of gonopores: four (0); three (1).
Ambulacra
5. Shape of the paired ambulacra: petaloid (0); subpetaloid (1); non-petaloid (2).
6. Anterior PZ of PPP and posterior PZ of APP: form an angle (0); form a crescentic line (1).
7. Petals: broad, i.e. IPZPZ (0); moderately wide IPZ∼PZ (1); narrow, i.e. IPZPZ (2).
8. Pores in the paired petals: elongate isopores (0); other (1).
9. Rudimentary pores in the anterior PZ of the APP: none (0); in the adapical plates, gradual change to normal-sized pores (1); in the adapical
plates, abrupt change (2); in all but the most distal plates (3).
10. Adoral ambulacra II and IV between phyllodes and margin: only slightly constricted, with roughly isometric plates (0); strongly constricted,
with sinuous, elongated plates (1).
11. Phyllodal plates: plates roughly isometric and few plates included in phyllodes II and IV (up to plates II.4a and IV.4a, respectively) (O); plates
elongated, crowded and numerous plates in phyllodes II and IV (usually extending beyond plates II.5a and IV. 5a, respectively) (1).
12. Number of pores within the subanal fasciole (adult condition): up to five (0); usually more than 5 (1).
Interambulacra
13. Labrum, posterior extension to the: 1st (1); 2nd (2); 3rd (3); 4th (4) ambulacral plates.
14. Labrum, shape: broad, short crescent, length<<width (0); roughly isometric (1); elongated, length>>width (2).
15. Contact between labrum and sternal plates: broad (0); narrow (1); disjunct (2).
16. Contact between sternal and episternal plates: straight (0); markedly V-shaped (1).
17. Primordial plates of interambulacral 1a and 4b, posterior extension to the: 1st (1); 2nd (2); 3rd (3); 4th (4); 5th (5); 6th (6) ambulacral plates.
18. Primordial plates of interambulacral 1b and 4a, posterior extension to the: 1st (1); 2nd (2); 3rd (3); 4th (4); 5th (5); 6th (6) ambulacral plates.
19. Primordial plates of interambulacral 2a and 3b, posterior extension to the: 1st (1); 2nd (2); 3rd (3); 4th (4); 5th (5); 6th (6) ambulacral plates.
20. Primordial plates of interambulacral 2b and 3a, posterior extension to the: 1st (1); 2nd (2); 3rd (3); 4th (4); 5th (5); 6th (6) ambulacral plates.
21. Constriction between the 1st and 2nd pair of episternal plates: absent or very weak (0); strong (1).
22. Periproct, structure: normal (0); invaginated (1).
Tuberculation and Sphaeridia
23. Labrum: fully tuberculate (0); partially tuberculate (1); naked (2).
24. Sternal plates: fully tuberculate (0); partially tuberculate (1); naked (2).
25. Primordial plates of interambulacral 1 and 4: fully tuberculate (0); partially tuberculate (1); naked (2).
26. Primordial plates of interambulacral 2 and 3: fully tuberculate (0); partially tuberculate (1); naked (2).
27. Conspicuous tuberculation pattern in adapical interambulacral 2b and 3a: absent (0); present (1).
 Hemipatagus – a misinterpreted Loveniid 179

Appendix 1 Continued
28. Aboral tuberculation: homogenous (0); heterogeneous (1).
29. Large aboral primary tubercles in the paired interambulacral: absent (0); present (1).
30. Large aboral primary tubercles in interambulacral 5: absent (0); present (1).
31. Aboral primary tubercles, structure: absent (0); crenulate (1); partially crenulate (2); non-crenulate (3).
32. Aboral primary tubercles: not markedly sunken (0); markedly sunken (1); camellate (2).
33. Aboral primary tubercles, arrangement: no marked pattern present (0); en chevron-arrangement, like Spatangus purpureus (1); arranged in
single rows on the interambulacral plates (2).
34. Aboral primary tubercles, location in interambulacral 1 and 5: all over the interambulacral (0); restricted to the anterior columns (1b and 4a)
and the interradial part of the posterior columns (1a and 4b) (1); restricted solely to the anterior columns (1b and 4a) (2).
35. Aboral primary tubercles, location in interambulacral 2 and 3: all over the interambulacral (0); restricted to the posterior columns (2a and
3b) and the interradial part of the anterior columns (2b and 3a) (1); restricted solely to the posterior columns (2a and 3b) (2).
36. Oral primary tubercles: not markedly sunken (0); markedly sunken (1); camellate (2).
37. Oral primary tubercles, parapet shape: ‘normal’ (0); helically spiralled (1).
38. Periplastronal areas: tuberculate (0); granular (1); naked (2).
39. Sphaeridia, structure: not in pits (0); in pits (1); in cysts (2).
Fascioles (adult condition)
40. Internal fasciole: absent (0); present (1).
41. Subanal fasciole: absent (0); bilobed (1); oval (2); shield-shaped (3).
42. Peripetalous fasciole: absent (0); present (1).

APP, anterior paired petals; IPZ, interporiferous zone: PPP, posterior paired petals: PZ, poriferous zone.

Appendix 2 Data matrix

1 1 1 1 1 1 1 1 1 1 2 22222 22223 33333 33334 44
Character 1 2345 67890 1 2345 67890 1 2345 67890 1 2345 67890 1 2
Brissus unicolor (Leske, 1778) 1 0000 0201 0 1 01 00 1 3733 1 0000 00000 00033 001 00 11
Eupatagus valenciennesi Agassiz, 1847 1 0000 0001 1 00321 03222 1 0201 1 01 1 0 1 0000 0021 0 21
Spatangus inermis Mortensen, 1913 02000 0001 0 001 00 03444 00001 1 01 01 1 01 33 001 00 10
Spatangus purpureus Müller, 1776 02000 0001 0 1 01 00 03444 00001 1 01 1 1 1 01 00 001 00 10
Eurypatagus grandiporus Mortensen, 1948 1 0000 01 01 0 1 0422 1 4544 1 01 21 1 01 1 1 1 0000 1 0200 00
Eurypatagus ovalis Mortensen, 1948 1 0000 01 01 0 1 0421 1 4554 1 01 21 1 01 1 1 1 0000 1 0200 00
Eurypatagus parvituberculatus (Clark, 1924) 1 0000 01 01 0 1 0422 1 4443 1 01 21 1 01 1 1 1 0000 1 0200 00
Platybrissus roemeri Grube, 1865 1 0000 0001 0 1 0421 1 4343 1 021 1 1 01 00 1 0000 00200 00
Maretia carinata Bolau, 1873 1 0000 0001 0 00321 0? ? ? ? 1 01 1 ? ? 01 1 1 1 1 000 0? 21 0 20
Maretia? cordata Mortensen, 1948 02000 0001 0 00321 0343? 1 021 ? 1 01 1 1 1 1 000 0? 21 0 20
Maretia planulata (Lamarck, 1816) 1 1 000 0001 0 1 0321 04433 1 01 1 1 1 01 1 0 1 1 200 1 1 21 0 20
Hemimaretia elevata (Döderlein, 1906) 1 1 01 1 01 1 1 ? 00221 1 2? ? ? 1 01 1 1 1 01 1 0 ? 2? ? ? 21 1 00 10
Goniomaretia tylota Clark, 1917 1 01 1 0 00021 00221 1 21 22 1 01 01 11110 32223 1 1 21 0 30
Pseudomaretia alta Agassiz, 1863 1 001 0 00021 00221 1 2222 1 01 01 11110 32222 01 1 1 0 30
Paramaretia multituberculata Mortensen, 1950 1 1 000 01 01 0 1 0421 1 3445 00221 1 01 1 1 32000 1 1 200 00
Breynia australasiae (Leach, 1815) 1 1 000 1 0020 01 221 1 3332 1 01 00 01 1 1 0 32201 0021 1 31
Breynia desorii Gray, 1851 1 1 000 00020 01 321 1 3332 1 01 00 01 1 1 0 32200 0021 1 31
Lovenia cordiformis Agassiz, 1872 02200 1 0021 01 321 1 3222 01 1 1 1 11110 3221 1 21 221 10
Lovenia elongata Gray, 1845 02000 1 0021 01 221 1 21 22 01 1 1 1 11110 3221 1 21 221 10
Lovenia gregalis Alcock, 1893 031 00 00021 00221 1 2222 001 1 1 11110 3221 2 21 21 1 10
Lovenia subcarinata Gray, 1845 02000 1 0021 00321 1 3222 001 1 1 11110 32222 21 221 10
Lovenia triforis Koehler, 1914 0201 0 01 021 00221 1 2332 001 1 1 11110 3201 2 21 21 1 10
Pseudolovenia hirsuta Ag. & Clark, 1907 0201 1 01 01 1 00321 1 3332 ? 01 1 1 11110 3221 2 21 2? 1 10
Echinocardium cordatum (Pennant, 1777) 021 00 1 1 021 00200 1 2222 1 01 01 1 1 000 00033 001 01 40
Echinocardium mortenseni Thiéry, 1909 1 1 000 01 021 0021 0 1 2222 1 01 01 1 1 000 00033 00201 10
Homolampas lovenioides Mortensen, 1948 1 1 01 2 021 1 ? 00321 1 223? ? 01 1 1 11110 32222 00200 10
Homolampas fragilis (Agassiz, 1869) 1 1 01 2 021 1 ? 00221 1 2222 ? 01 1 1 11110 32222 00200 10
Hemipatagus? argutus Clark, 1915 02000 0201 1 0? ? 21 1???? ? 01 1 1 1 01 1 0 1 1 01 1 002? 0 10
Hemipatagus? carolinensis (Kier, 1997) 02000 00021 0? 221 1???? ? 01 1 ? ?1111 ?1112 002? 0 10
Hemipatagus hoffmanni (Goldfuss, 1826) 02000 00021 00221 1 21 22 0021 1 11110 2221 2 1 1 21 0 10
Hemipatagus ocellata (Defrance, 1827) 02000 00021 00221 1 2223 0021 1 11110 3221 2 21 21 0 10
Hemipatagus? subrostratus Clark, 1915 020? 0 020? ? ? ? ? 21 ????? ? 0? 1 1 1?110 1 1 01 2 002? 0 ?0

Note: In characters 17–20 there may be some variation in character state between the paired interambulacra, here the most common state was used.
180 A. Kroh
Appendix 3 Annotated list of the
nominal species of Hemipatagus
and Maretia
Also included are the species subsequently placed in either
genus (based on Lambert & Thiéry 1909–1925 and Kier &
Lawson 1978). Systematic decisions were made on the basis
of published descriptions and illustrations or taken from the
literature where modern revisions were available.
affinis Duncan & Sladen, 1883
∗
1883 Euspatangus [sic] affinis, Duncan & Sladen;
Duncan & Sladen: 46–47, pl. 12, fig. 2.
1924 Hemipatagus affinis Duncan et Sladen (Euspatan-
gus [sic]); Lambert & Thiéry: 457.
2004 Eupatagus (Eupatagus) affinis (Duncan & Sladen)
n. comb; Srivastava: 145–146; pl. 6, figs 5–6.
HOLOTYPE. The specimen figured by Duncan & Sladen
(1883); Geological Survey of India no. C 043 B (GSI Type
No. 2867 (Srivastava 2004)).
TYPE LOCALITY. One mile east of Goir, near Narainsir,
Kachchh, Pakistan.
OCCURRENCE. Numulitic Series of Kachchh.
AGE. Middle Eocene (Srivastava 2004).
TAXONOMIC PLACEMENT. Eupatagus Duncan & Sladen
(1883); on the basis of the occurrence of a peripetalous and
a subanal fasciole.
affinis Herklots, 1854
∗
1854 Spatangus affinis, nouv. esp.; Herklots: 12; pl. 2,
figs 5, 5a.
1937 Eupatagus (s. Brissoides) affinis (Herklots);
Jeannet & Martin: 274–275, fig. 51.
LECTOTYPE. The specimen figured by Herklots (pl. 2, figs
5, 5a), no. 477 (the same number is mentioned under his S.
pulchellus); no. L4292 according to Jeannet & Martin (1937:
274), Rijksmuseum van Geologie en Mineralogie, Leiden.
TYPE LOCALITY. Tjidamar, Java.
AGE. Pliocene.
TAXONOMIC PLACEMENT. Eupatagus (Henderson 1975:
33). According to Jeannet & Martin (1937: 274) and
Henderson (1975: 33), Herklots’ two syntypes represent two
different genera/species. One specimen (no. L4292) belongs
to the genus Eupatagus. The other specimen (no. L4291) is
a misidentified Maretia planulata (Lamarck) (Martin 1880:
81; Gerth 1922: 512). From Herklots’ paper alone, however,
it is not obvious that there were two specimens and it is un-
clear which of the two specimens mentioned by Jeannet &
Martin (1937) is the figured specimen.
alta A. Agassiz, 1864
∗
1864 Maretia alta A. Ag.; Agassiz: 360.
1951 Pseudomaretia alta (A. Agassiz); Mortensen: 58–
62; pl. 3, figs 12–14, 16–20; pl. 46, figs 1–6, 8–11,
13, 19, 20.
TYPE MATERIAL. Museum of Comparative Zoological at
Harvard College, Cambridge.
TYPE LOCALITY. Kagoshima Bay, Japan, ∼ 9 m water depth.
AGE. Recent.
TAXONOMIC PLACEMENT. Type species of Pseudomaretia
Koehler, 1914.
anomala Duncan, 1877
∗
1877 Maretia anomala, sp. nov.; Duncan: 53–53; pl. 4,
figs 1–4.
1985 Eupatagus anomalus (Duncan, 1877); Kruse &
Philip: 173–174; figs. 9a–b; pl. 2, fig. 5; pl. 3,
figs 1–3; pl. 5, figs 4–5.
HOLOTYPE. Natural History Museum, London 14072,
figured by Duncan (1877: pl. 4, figs 1–4).
TYPE LOCALITY. Cliffs at the mouth of the Sherbrooke River,
Victoria, Australia.
TYPE OCCURRENCE. Ruthledge Creek Member.
AGE. Balcombian, Middle Miocene.
TAXONOMIC PLACEMENT. Eupatagus (Kruse & Philip
1985: 173); based, among other features, on the presence
of a peripetalous fasciole, which had been previously recog-
nised by Duncan (1877) and subsequent authors, but still the
species remained in the genus Maretia until the revision by
Kruse & Philip (1985).
antillarum Cotteau, 1875
∗
1875 Euspatangus [sic] Antillarum, Cotteau, 1875;
Cotteau: 43; pl. 7, figs 7–11.
SYNTYPES. Collection Cleve, Museum Stockholm, Museum
of Upsala.
TYPE LOCALITY. St. Barthélemy Island.
AGE. Eocene.
TAXONOMIC PLACEMENT. Eupatagus (Henderson 1975:
33), senior synonym of Maretia clarki Lambert and Maretia
twitchelli Lambert according to Henderson (1975: 33).
aragonensis Cotteau, 1887
∗
1887 Maretia aragonensis Cotteau, 1887; Cotteau: 93–
95; pl. 11, figs 9–12; No. 57.
LECTOTYPE. Specimen figured by Cotteau (1887: pl. 11,
figs 9–12), collection Maurice Gourdon.
PARALECTOTYPES. Not figured, collection Cotteau.
TYPE LOCALITY. Pobla de Roda, Aragon, Spain.
AGE. Eocene.
 Hemipatagus – a misinterpreted Loveniid
TAXONOMIC PLACEMENT. Leiopneustes (Lambert & Thiéry
1924: 448); periproct high on the posterior face, visible in
aboral view, sternal plates fully tuberculate, no naked peri-
plastronal areas (providing Cotteau’s illustrations are correct;
the plating of the oral side, shown in fig. 11, however, is ob-
viously wrong), petals are strongly divergent, not at all like
anything found in Lovenia, Hemipatagus or Maretia. On the
whole the illustrations resemble much more a brissid or some
hemiasterids than a spatangid or loveniid.
archiaci Agassiz, in Agassiz & Desor, 1847
∗ ∗
1847 Spatangus Archiaci Agass.; Agassiz & Desor: 8.
TYPE MATERIAL. Collection d’Archiac.
TYPE LOCALITY. Oulchy-le-Châteaux, France.
LITHOSTRATIGRAPHY. Calcaire grossier.
AGE. Eocene.
TAXONOMIC PLACEMENT. Doubtful; according to Cotteau
(1885) this species is synonymous with ‘Spatangus’ grignon-
ensis Cotteau, 1880.
argutus Clark, in Clark & Twitchell, 1915
∗
1915 Hemipatagus argutus Clark, n. sp.; Clark &
Twitchell: 150; pl. 49, figs 1a–d.
1959 Maretia arguata (Clark); Cooke: 81; pl. 34,
figs 1–4.
1993 Maretia arguata (Clark, 1915); Zachos: 148–150;
figs 1.1–3.
HOLOTYPE. USNM 141107; U.S. National Museum, Smith-
sonian Institution, Washington, USA (according to Kier
(1980) only a single specimen known, although Cooke (1959)
listed three localities).
TYPE LOCALITY. Chickasawhay River at Enterprise, Clarke
County, Mississippi, USA.
TYPE OCCURRENCE. Winona Formation, Clairborne Group.
AGE. Middle Eocene.
OCCURRENCE. Middle Eocene of Mississippi and Texas,
USA.
TAXONOMIC PLACEMENT. Hemipatagus? Although this
species (like the other New World species of Hemipatagus)
has no camellate tubercles, it is tentatively attributed to Hemi-
patagus based on the characteristic cordiform outline, the
presence of rudimentary pores in the anterior poriferous
zones of petals II and IV and the distribution of the aboral
primary tubercles; according to Cooke (1959: 81) and Kier
(1980: 50) this species might be a junior synonym of H.
subrostratus Clark.
bandaensis Martin, in Jeannet & Martin, 1937
∗
1937 ? Hemipatagus bandaensis nov. spec. (R. Martin).;
Jeannet & Martin: 276; fig. 53a–c.
HOLOTYPE. Specimen figured by Jeannet & Martin (1937:
fig. 53a–c); private coll. Mieg, Basel.
181
TYPE LOCALITY. Banda Islands.
AGE. ? Pliocene.
TAXONOMIC PLACEMENT. Hemipatagus?; all features
(cordiform outline, frontal sinus, lack of internal or peri-
petalous fascioles, shape of the petals, presence of a bilobed
subanal fasciole, distribution of the aboral primary tubercles
. . .) in the original publication support an attribution to Hemi-
patagus.
buendensis Ebert, 1889
1889 Spatangus (Maretia) bündensis Ebert n. sp.; Ebert:
69–70; pl. 8, fig. 1a–c.
SYNTYPES. Two specimens in the Museum of Göttingen.
TYPE LOCALITY. Doberg, Germany.
AGE. Late Oligocene.
TAXONOMIC PLACEMENT. Hemipatagus; very similar to the
juveniles of H. martensii, as observed previously by Ebert
(1889); Lambert & Thiéry (1924: 457) consider H. buenden-
sis to be a junior synonym of H. hoffmanni.
calvimontanus Cotteau, 1885
∗
1885 Maretia calvimontana, Cotteau, 1885; Cotteau: 36–
38; pl. 5, figs 1–3.
HOLOTYPE. Specimen figured by Cotteau (1885: pl. 5, figs
1–3), collection ‘de la Sorbonne’, Paris.
TYPE LOCALITY. Chaumont (Oise), France.
LITHOSTRATIGRAPHY. Calcaire grossier inférieur.
AGE. Middle Eocene.
TAXONOMIC PLACEMENT. Hemipatagus; very similar to H.
grignonensis, possibly a juvenile or subadult specimen of
that species; the cordiform outline with distinct frontal sinus,
the shape and structure of the petals and the distribution
of the primary tubercles support an attribution to Hemi-
patagus; the original description does not reveal details on
the nature and tuberculation of the plastron and the shape of
the subanal fasciole.
carinata Bolau, 1873
∗
1873 [Maretia] carinata n. sp. Bolau: 6.
1951 Maretia carinata Bolau; Mortensen: 39–41; pl. 1,
figs 5–7; pl. 46, figs 14–16.
SYNTYPES. Museum of Hamburg, Germany (2 specimens in
alcohol, Viti Islands); Museum Godeffroy (2 dry specimens,
Viti Islands; 1 dry specimen, Ellice Islands), Musuem of
Göttingen, Germany (1 specimen in alcohol, Bay of Bengal).
TAXONOMIC PLACEMENT. Maretia; the elongate outline,
lack of a frontal sinus, long and straight petals and the distri-
bution of the non-camellate aboral primary tubercles support
an attribution to the genus Maretia; the shape of the subanal
fasciole is unknown and the adapical pores in the anterior
poriferous zones of the anterior paired petals are reduced in
size.
182 A. Kroh
carolinae Gagel, 1905
∗
1905 Chuniola Carolinae gen. nov. sp. n. Gagel: 531–
537; fig. 1; pl. 24, figs 1a–c, 2a–c, 3, 4a–b, 5.
SYNTYPES. Six whole and several fragmentary specimens
in the collections of the Geological Survey of Preussen, the
Museum of Lübeck and the Museum of Hamburg.
TYPE LOCALITY. Zarrentin, south-east Holstein, Northern
Germany.
AGE. Erratic boulders of Cenozoic age, presumably Mio-
cene.
TAXONOMIC PLACEMENT. Type species of Chuniola Gagel,
1905; probably a subjective junior synonym of Hemipatagus
Desor, 1858 according to Smith (2004: ‘The Echinoid Direct-
ory’); differs from typical Hemipatagus by lacking enlarged
aboral primary tubercles in interambulacra 2 and 3 and by
the presence of such tubercles in adapical interambulacrum
5; due to the poor preservation of the specimens the pres-
ence/absence of internal, subanal and peripetalous fascioles
could not be ascertained (a peripetalous fasciole, however,
seems to be lacking if Gagel’s figures are correct); Gagel
(1905: pl. 24, fig. 5) illustrated a plastron with broad, short
labrum very unlike anything observed in Maretia, Hemi-
patagus, or Lovenia.
carolinensis Kier, 1997
∗
1997 Maretia carolinensis, new species, Kier: 11–13;
fig. 6; pl. 9, figs 8–9; pl. 10, figs 1–7; pl. 11, figs
1–2.
HOLOTYPE. USNM 398338, figured by Kier (1997: pl. 10,
figs 1–4), U.S. National Museum, Smithsonian Institution,
Washington, USA.
PARATYPES. USNM 398337, 398339–398341, 492101–
492124.
TYPE LOCALITY. Polocksville state quarry, North Carolina,
USA.
TYPE OCCURRENCE. Trent Formation.
AGE. Late Oligocene.
TAXONOMIC PLACEMENT. Hemipatagus; although this spe-
cies has no truly camellate tubercles (even though the areoles
are distinctly sunken), it clearly belongs to Hemipatagus
based on the characteristic cordiform outline, typical shape
of the petals with rudimentary pores in the anterior porifer-
ous zones of petal II and IV and the conspicuous field of
tubercles in adapical ambulacrum III.
cartagensis Sánchez Roig, 1949
∗ are confined to the area inside the petalodium and seem to
1949 Hemipatagus cartagensis Sánchez Roig: 216–217
[not seen, fide Kier & Lawson 1978: 121].
AGE. Late Oligocene (Brodermann 1949: 323) or Late Eo-
cene (Sánchez Roig 1949).
TAXONOMIC PLACEMENT. Doubtful; not mentioned in
Kier’s (1984) revision of the fossil spatangoids of Cuba.
clarki Lambert, in Sanchez-Roig, 1924
1924 Maretia clarki Lambert in Sánchez Roig; Lambert
& Thiéry: 458. [nom. nov. pro specimen B of Eu-
patagus floridanus Clark, 1915 (in Clark & Twitchell
1915: 176–177; pl. 83, figs 1a–c)].
HOLOTYPE. Specimen figured by Clark & Twitchell (1915:
pl. 83, figs 1a–c); Wagner Free Institute of Science.
TYPE LOCALITY. Levy County, Florida, USA.
AGE. Early Oligocene.
TAXONOMIC PLACEMENT. Eupatagus; based on the pres-
ence of a peripetalous fasciole; junior synonym of Eupatagus
antillarum (Cotteau) according to Henderson (1975: 33).
cordata Mortensen, 1948
∗
1948 Maretia cordata, new species; Mortensen: 132–
133.
1951 Maretia cordata Mrtsn.; Mortensen: 41–45; pl. 1,
figs 8, 11–16; pl. 44, figs 9–10, 20, 25, 28, 30–31.
HOLOTYPE. USNM E7158 (one adult from the ‘Albatross’-
Expedition Stat. 5192 (11◦ 09 N, 123◦ 50 E, 59 m)).
ADDITIONAL MATERIAL. Four specimens from the
‘Albatross’-expedition (one juvenile from Stat. 5104
(14◦ 45 48 N, 120◦ 12 E, 60 m), 1 adult from Stat. 5257,
Mindanao (7◦ 22 N, 124◦ 12 E, 51 m), two adults from Stat.
5431 (10◦ 38 45 N, 120◦ 12 45 E, 93m).
TYPE AREA. Philippines.
AGE. Recent.
OCCURRENCE. Philippines; Palawan Islands; Bali Sea; ?
‘Siboga’-expedition Stat. 207 (Flores Sea).
TAXONOMIC PLACEMENT. Maretia?; more pronounced
cordate outline than M. planulata with distinct frontal notch;
the subanal fasciole is oval in outline as in M. planulata.
depressus Dubois, 1843
∗
1843 Spatangus depressus; Dubois: pl. 1, fig. 16.
1847 Spatangus depressus Dub.; Agassiz & Desor: 8.
1858 Hemipatagus depressus; Desor: 417.
1880 Hemispatangus [sic] depressus (Dubois), Desor.;
de Loriol: 135–136; pl. 11, fig. 6, 6a–c.
HOLOTYPE. The specimen figured by Dubois (1843).
TYPE AREA. Crimea, Ukraine.
AGE. ? Eocene.
TAXONOMIC PLACEMENT. Eupatagus?; the illustration in
Dubois’ paper shows the aboral surface of a damaged speci-
men with heterogeneous aboral tuberculation. The tubercles
have only slightly sunken areoles. This and the shape of the
petals is strongly reminiscent of Eupatagus and the species
is tentatively referred to that genus. The illustrations in de
Loriol (1880) are rather poor and do not offer additional
insights. Unfortunately, the present author did not have the
opportunity to check the text of Dubois’ work for a descrip-
tion of the species.
 Hemipatagus – a misinterpreted Loveniid
desmoulinsi Cotteau, 1863
∗
1863 Eupatagus Des Moulinsi, Cotteau, 1863; Cotteau:
148–149; pl. 7, fig. 11.
1885 Maretia Des Moulinsi (1863), Cotteau, 1885;
Cotteau: 26–28; pl. 2, figs 1–6.
HOLOTYPE. The specimen figured by Cotteau (1863: pl. 7,
fig. 11); collection of Cotteau.
TYPE LOCALITY. La Goureppe, near Biarritz, Basses-
Pyrénées, France.
AGE. Late Eocene.
TAXONOMIC PLACEMENT. Hemipatagus; initially Cotteau
(1863: 148) assumed the presence of a peripetalous fasciole,
but stated that he could not observe it in the specimens.
In 1885 he renounced this assumption and transferred the
species to Maretia; based on the presence of a distinct frontal
sinus, the distribution of the camellate primary tubercles, the
structure of the petals, and the presence of a bilobed subanal
fasciole, the species is here attributed to Hemipatagus.
duncani Lambert, in Lambert & Thiéry, 1924
∗
1924 H. Duncani Lambert; Lambert in Lambert &
Thiéry: 457 [nom. nov. pro Eupatagus rostratus
Duncan & Sladen (1884b: 267, pl. 43, fig. 6), non
d’Archiac].
1975 Eupatagus rostratus rostratus d’Archiac;
Henderson: 44–45.
2004 Eupatagus (Eupatagus) rostratus (d’Archiac) n.
comb.; Srivastava: 146–147; pl. 7, fig. 8.
HOLOTYPE. The specimen figured by Duncan & Sladen
(1884b: 267, pl. 43, fig. 6) as Eupatagus rostratus d’Archiac;
Geological Survey of India no. G226/76.
TYPE LOCALITY. West side of Bagothoro Hill, south of
Sehwan, India.
TYPE OCCURRENCE. Nari Series.
AGE. Middle Oligocene (Srivastava 2004).
OCCURRENCE. ?Eocene–Oligocene and, possibly, Miocene
of India, Japan and Java.
TAXONOMIC PLACEMENT. Eupatagus; junior synonym of E.
rostratus rostratus d’Archiac, 1850 according to Henderson
(1975: 44).
elevata Döderlein, 1906
∗
1906 Maretia elevata nov. spec.; Döderlein: 263–265;
pl. 33, figs 5, 5a–b; pl. 48, figs 6a–h.
1951 Hemimaretia elevata (Döderlein); Mortensen: 53–
55; pl. 1, fig. 4; pl. 44, figs 5–7, 21.
HOLOTYPE. Badly broken specimen in the Museum of
Berlin (Mortensen 1951).
TYPE LOCALITY. Station 251 (Valdivia-Expedition), off Wit-
uland, Eastern Africa (693 m depth).
AGE. Recent.
183
TAXONOMIC PLACEMENT. Type-species of Hemimaretia
Mortensen, 1950; this genus has only 3 gonopores (pore
of genital plate 2 lacking) and is thus easily distinguished
from Hemipatagus, Lovenia and Maretia.
elliptica Bolau, 1874
∗
1874 Maretia elliptica n. sp.; Bolau: 175–177; pl. 6,
figs 1–2.
HOLOTYPE. Figured by Bolau (1874: pl. 6, figs 1–2), Mu-
seum of Hamburg.
TYPE LOCALITY. Malden Island, Pacific Ocean.
AGE. Recent.
TAXONOMIC PLACEMENT. Insufficiently known; based on
a single specimen; according to Mortensen (1950: 281, 1951:
26) this species probably belongs to either Platybrissus or,
less likely, to Eurypatagus.
estenozi Sánchez Roig, 1926
1984 ? Maretia estenozi Sánchez Roig; Kier: 106–107;
pl. 55, fig. 5, pl. 56, figs 1–2.
HOLOTYPE. SRC 5491, figured by Sánchez Roig (1926:
pl. 37, fig. 1); Sánchez Roig collection, Cuban Academy
of Science.
TYPE LOCALITY. ‘Cervantes’ farm, San José de las Lajas,
Haban Province, Cuba.
AGE. Early – Middle Miocene.
TAXONOMIC PLACEMENT. Eupatagus (Kier, 1984: 107),
possible synonym of E. cubensis (Cotteau, 1875) according
to Kier (1984: 103–107).
fasciata Lambert, in Lambert & Thiéry, 1924
∗
1924 M. fasciata Lambert; Lambert in Lambert &
Thiéry: 458.
HOLOTYPE. Unknown.
TYPE LOCALITY. Philippines.
AGE. Recent.
TAXONOMIC PLACEMENT. Maretia; juvenile of M. planulata
according to Mortensen (1951: 38).
ficheuri Pomel, 1887
∗
1887 Hemipatagus Ficheuri; Pomel: 26 [figured on the
unpublished plate A.23 according to Cotteau et al.
(1891: 83)].
TAXONOMIC PLACEMENT. Nomen dubium; Cotteau et al.
(1891: 83) state that this species is unknown to them; never
described or illustrated; on page 79 of the same paper they
tentatively refer to it in the synonymy of Maretia tenuis Peron
& Gauthier, in Cotteau et al., 1891.
forbesii Woods, 1862
∗
1862 Spatangus Forbesii; Woods: 75, 83, 105; fig. on
p. 75 and 83 [nom. nov. pro Spatangus Hoffmanni
Sturt 1833 non Goldfuss].
184 A. Kroh
1864 Hemipatagus Forbesi, Woods & Duncan; Duncan:
165, pl. 6, fig. 3.
1877 Lovenia Forbesi, Woods and Duncan; Duncan: 44,
56–61, pls 4, figs 5–8.
1994 Lovenia forbesii (Woods, 1862); Irwin & Archbold:
2–5, figs 2A-K [cum syn.].
LECTOTYPE. Specimen figured by Sturt (1833: pl. 3, fig. 10),
designated by Irwin & Archbold (1994: 2); current where-
abouts unknown.
TYPE LOCALITY. Murray cliffs at Mannum, South Australia.
TYPE OCCURRENCE. Mannum Formation.
AGE. ?Longfordian (Burdiagalian), Early Miocene.
TAXONOMIC PLACEMENT. Lovenia; some references
(Woods 1859, 1862, 1865, 1878; Etheridge 1878; McCoy
1879; Dennant 1890) referring to this species actually
represent another species (Lovenia bagheerae Irwin in Irwin
& Archbold, 1994) according to Irwin & Archbold (1994).
formosus Zittel, 1867
∗
1867 Hemipatagus formosus Zitt.; Zittel: 63; pl. 12,
figs 2a–c.
1894 Eupatagus formosus; Tate: 126 [not seen].
1975 Eupatagus rostratus zitteli nomen nov.; Henderson:
45–46; fig. 10; pl. 11, figs 5–6; pl. 12, figs 1–3.
LECTOTYPE. NHMW 1970/1336, designated by Henderson
(1975: 45); Natural History Museum Vienna, Austria.
PARALECTOTYPE. NHMW 1959/335/50, 1970/1337.
TYPE LOCALITY. Cape Farewell, west Nelson, New Zealand.
AGE. ? Abel Head Fm., Late Oligocene.
OCCURRENCE. Whaingaroan to Altonian (Oligocene to
Early Miocene) of New Zealand.
TAXONOMIC PLACEMENT. Eupatagus; E. formosus (Zittel,
1867) is a secondary homonym of E. formosus de Loriol,
1863 and was renamed E. rostratus zitteli by Henderson
(1975: 45–46).
fuchsi Oppenheim, in Blanckenhorn, 1901
∗
1901 Maretia Fuchsi Oppenheim; Oppenheim in Blanck-
enhorn: 114–117, fig. 9.
SYNTYPES. 1 specimen from Mirsa Badia, Cyrenaica, Libya
(coll. Schweinfurth; figured by Oppenheim in Blancken-
horn, 1901: fig. 9); 1 fragment from Gebel Geneffe, Eastern
Desert, Egypt (coll. Blanckenhorn); 11 fragments from Wadi
Etthal, Sinai, Egypt (coll. Rothpeltz); current whereabouts
unknown.
AGE. Presumably Early or Middle Miocene.
TAXONOMIC PLACEMENT. Spatangus?; classification un-
clear; Oppenheim reports that he could not find any traces
of an internal or peripetalous fasciole, but records the pres-
ence of a subanal fasciole and a fully tuberculate plastron;
these features (if correct) suggest that the species is related
to Spatangus.
gauthieri Lambert, in Lambert & Thiéry, 1924
1885 Tuberaster tuberculatus, Peron & Gauthier, 1885;
Cotteau et al.: 47–49; pl. 3, figs 1–4 [type-species
of Tuberaster Peron & Gauthier in Cotteau et al.,
1885].
1893 Tuberaster tuberculatus, Peron & Gauthier, 1885;
Cotteau: 660–662; pl. 360, figs 5–10.
∗
1924 H. Gauthieri Lambert; Lambert in Lambert &
Thiéry: 457 [nom. nov. pro Tuberaster tuberculatus
Péron & Gauthier, 1885 non Zittel, 1867].
HOLOTYPE. Specimen figured by Cotteau et al. (1885: pl. 3,
figs 1–4); collection Gauthier.
TYPE LOCALITY. Kef Iroud, Algeria.
AGE. Oligocene.
TAXONOMIC PLACEMENT. Hemipatagus?; apart from the
fully tuberculate sternum (if correctly figured in Cotteau
et al. 1885) this species is very similar to typical Hemi-
patagus species (like H. hoffmanni); although Cotteau et al.
(1885) debate much about the presence and absence of fasci-
oles, none are ascertained due to the heavy abrasion of their
material.
girundicus Lambert, 1915
∗
1915 H. girundicus; Lambert: 191.
1924 H. girundicus Lambert; Lambert in Lambert &
Thiéry: 457; pl. XIII, fig. 5.
1927b Hemipatagus girundicus Lambert, 1915; Lambert:
119–121.
HOLOTYPE. Whereabouts unknown, a cast of the specimen
is figured in Lambert & Thiéry (1924: pl. 13, fig. 5).
TYPE AREA. Saucats, Bordelaise, France.
AGE. Early Miocene.
TAXONOMIC PLACEMENT. Hemipatagus; very similar to H.
ocellatus from the Burdigalian of the Rhône Basin.
gottschei Gagel, 1905
∗
1905 Spatangus ? (Eupatagus ?) Gottschëi sp. n.; Gagel:
539–541; pl. 24, fig. 7; pl. 25, figs 1a–c, 2, 8.
SYNTYPES. 3 natural moulds and one cast in the collections
of the Geological Survey of Preussen, and the Museum of
Hamburg.
TYPE LOCALITY. Zarrentin, south-east Holstein, Northern
Germany.
AGE. Erratic boulders of Cenozoic age, presumably
Miocene.
TAXONOMIC PLACEMENT. Hemipatagus?; based on the out-
line and the distribution of the aboral primary tubercles; type
material very poor and in need of redescription.
grignonensis Cotteau, 1880
1837 Spatangus grignonensis Desmarest, ined. in litt.;
des Moulins: 236–237 [nomen nudum].
 Hemipatagus – a misinterpreted Loveniid
∗
1885 Maretia grignonensis (Desmarets) [sic], Cotteau,
1880; Cotteau: 30–36; pl. 3, figs 1–8; pl. 4, 1–4.
TYPE MATERIAL. Unknown.
TYPE AREA. Paris Basin (Grignon, Cahumont, Fours,
Chenet . . .); France.
LITHOSTRATIGRAPHY. Calcaire grossier.
AGE. Middle Eocene.
TAXONOMIC PLACEMENT. Hemipatagus; Cotteau’s descrip-
tion and illustrations partly contradict each other; assuming
the description to be correct this species has a partially tuber-
culate sternum, a bilobed subanal fasciole, a cordate outline
with distinct frontal notch but no rudimentary pores in adap-
ical petals II and IV; generic attribution thus is doubtful but
Hemipatagus seems to fit best; according to Cotteau (1885)
this species is synonymous with Spatangus omali Galeotti,
1837 and S. archiaci Agassiz & Desor, 1847.
REMARKS. Although frequently cited, Desmarest never pub-
lished this species; it is first mentioned in des Moulins (1837)
referring to an unpublished manuscript by Desmarest; as des
Moulins provides no description or illustration he cannot be
considered the valid author either.
guebhardi Lambert & Savin, 1902
∗
1902 Maretia Guebhardi Lambert et Savin nov. sp.;
Lambert & Savin: 882–889; pl. 59, figs 1–5.
1915 Hemipatagus Guebhardi Lambert & Savin (Mare-
tia), 1905; Lambert: 189.
HOLOTYPE. Specimen figured by Lambert & Savin (1902:
pl. 59, figs 1–5), Savin collection.
TYPE LOCALITY. Gattières, Alpes-Maritimes, France.
AGE. Burdigalian, Early Miocene.
TAXONOMIC PLACEMENT. Hemipatagus; clearly a member
of the genus Hemipatagus with all the typical features de-
veloped (outline, tuberculation . . .); very similar to subadult
H. ocellatus.
heberti Cotteau, 1885
∗
1885 Maretia Heberti, Cotteau, 1885; Cotteau: 38–40;
pl. 5, figs 4–8.
TYPE MATERIAL. Sorbonne collection.
TYPE LOCALITY. La Mortagne, Oise, France.
LITHOSTRATIGRAPHY. Calcaire grossier.
AGE. Middle Eocene.
TAXONOMIC PLACEMENT. Referred to Phymapatagus
Lambert, 1910 (type-species: S. britannus Tournouer) by
Lambert & Thiéry (1924: 463); this is supported by the fully
tuberculate plastron and lack of camellate primary tubercles.
hispanica Cotteau, 1890
∗
1890 Maretia hispanica, Cotteau, 1889; Cotteau: 5–6;
pl. 22 (1), figs 1–3.
185
HOLOTYPE. Specimen figured by Cotteau (1890); Cotteau
collection.
TYPE LOCALITY. Callosa, Alicante, Spain.
AGE. ? Eocene.
TAXONOMIC PLACEMENT. Hemipatagus?; the shape of the
test and the distribution and nature of the primary tubercles
would indicate an attribution to Hemipatagus, yet, the
plastron is shown as being fully tuberculate in pl. 22 (1), fig.
3; it is known, however, that Cotteau’s artist often provided
highly idealized figures, adding plate sutures or tubercles
where there were none originally. Unfortunately the descrip-
tion gives no further clues.
hoffmanni Goldfuss, 1829
∗
1829 Spatangus hofmanni [sic] nobis; Goldfuss: 152–
153; pl. 47, figs 3a–c.
TYPE MATERIAL. Figured specimen and seven additional
syntypes in the collection of the Goldfuss Museum (Institute
of Palaeontology, University of Bonn, Germany), number
357a–h.
TYPE LOCALITY. Doberg, near Bünde, Germany.
AGE. Chattian, Late Oligocene.
TAXONOMIC PLACEMENT. Type species of Hemipatagus.
hungarica Vadász, 1915
∗
1915 Maretia hungarica n.sp.; Vadász: 237; pl. 9 (3),
figs 17–18.
1952 Maretia hungarica Vadász; Szörényi: 302, figs a–b.
HOLOTYPE. Specimen MAFI Ech-225, figured by Vadász
(1915: pl. 9 (3), figs 17–18) and Szörényi (1952: figs a–b);
housed at the Museum of the Hungarian Geological Survey,
Budapest.
TYPE LOCALITY. Matraverebély, Nógrad, Hungary.
AGE. Late Badenian (Early Serravallian), Middle Miocene.
REMARKS. Specimen damaged since the photograph of
Szörényi (1952), petal V now missing.
TAXONOMIC PLACEMENT. Hemipatagus; Lambert & Thiéry
(1924: 458) tentatively referred this species to Atelospatan-
gus Koch, 1885 because no anterior poriferous zones of petals
II and IV are visible in Vadsz’s drawing; the correctness of
this action was questioned by Mortensen (1951: 79), arguing
that the drawing might be incorrect; Mortensen was indeed
correct, as shown by Szörényi (1952: 302), only the adapical
pores of the anterior poriferous zones of petals II and IV are
rudimentary; no internal fasciole could be observed in the
type.
integer Sorignet, 1850
∗
1850 Spatangus integer; Sorignet: 49 [not seen, fide
Desor, 1858: 417].
1858 Hemipatagus integer; Desor: 417.
1885 Maretia integer (Sorignet), Cotteau, 1885; Cotteau:
40–41.
186 A. Kroh
HOLOTYPE. Unique specimen in the collection Sorignet;
whereabouts unknown.
TYPE LOCALITY. Fours, Eure, France.
LITHOSTRATIGRAPHY. Calcaire grossier.
AGE. Middle Eocene.
TAXONOMIC PLACEMENT. Doubtful; published descriptions
are insufficient.
koeneni Ebert, 1889
∗
1889 Spatangus (Maretia) Koeneni Ebert, n.sp.; Ebert:
60–62: pl. 8, figs 2a–d.
TYPE MATERIAL. Figured specimen in the collection of the
University of Göttingen; others in the collections of the Uni-
versity of Munich, the Museum of Breslau, the Natural His-
tory Museum of Osnabrück and the Geological Survey in
Berlin.
TYPE LOCALITY. Doberg, near Bünde, Germany.
AGE. Late Oligocene.
TAXONOMIC PLACEMENT. Similar to H. hoffmanni, but with
fully tuberculate sternal plates and non-camellate primary
tubercles, probably fits best in Spatangus (Phymapatagus)
? (based on the description of Ebert; re-examination neces-
sary).
lorioli Cotteau, 1878
∗
1878 Lovenia Lorioli, Cotteau, 1878; Cotteau: 216–217;
pl. 30, figs 2–7; No. 120.
HOLOTYPE. Specimens figured by Cotteau (1878: pl. 30,
figs 2–7); Gauthier collection.
TYPE LOCALITY. Drôme, France.
AGE. Burdigalian, Early Miocene.
TAXONOMIC PLACEMENT. ? Lovenia; Cotteau’s fig. 7
provides a magnification of the aboral primary tubercles,
showing distinct crenulations; no extant species of Love-
nia with crenulated primary tubercles are known (Mortensen
1951); although Cotteau explicitly stated that an internal and
a subanal fasciole are present, Lambert & Savin (1902: 884)
and Lambert & Thiéry (1924: 467) state that this species
could be a Hemipatagus, an opinion not shared by any sub-
sequent author; lately Philippe (1998: 225) suggested that
the type specimen might be a juvenile Maretia ocellata (here
Hemipatagus); he argued that he could find no trace of an in-
ternal fasciole in the cast of the holotype studied by him and
that the general appearance was much more like H. ocellatus
than like a Lovenia.
madurae Böhm, 1882
∗
1882 Hemipatagus Madurae n. sp.; Böhm: 366–367;
pl. 2, fig. 3a–c.
HOLOTYPE. Specimen figured by Böhm (1882: pl. 2,
figs 3a–c); whereabouts unknown.
TYPE LOCALITY. Sepoloc, northern coast of Madura, N of
Java.
AGE. ? Neogene.
TAXONOMIC PLACEMENT. ? Lovenia (referred to the genus
Vasconaster by Lambert & Thiéry (1924: 466), today con-
sidered a junior synonym of Lovenia); generic attribution
unclear, type material needs to be re-examined to verify the
presence or absence of an internal fasciole and to study the
morphological details.
marianii Airaghi, 1903
∗
1903 Maretia Marianii n. sp.; Airaghi: 423–424; fig. 10.
HOLOTYPE. Specimen figured by Airaghi (1903: fig. 10),
Museo civico Milano, Italy.
TYPE LOCALITY. Tirolo, Veneto, Italy.
AGE. Eocene.
TAXONOMIC PLACEMENT. Hemipatagus; typical member of
the genus Hemipatagus with the characteristic cordiform out-
line with distinct frontal sinus, typical distribution of the
aboral primary tubercles (restricted to the anterior part near
the interradial suture in the interambulacra 1 and 4) and
typical shape of the petals, with rudimentary pores in the
adapical anterior poriferous zones of ambulacra II and IV.
martensii Ebert, 1889
∗
1889 Spatangus (Maretia) Martensii Ebert n.sp.; Ebert:
67–69; pl. 9, fig. 2a–d.
HOLOTYPE. Specimen figured by Ebert (1889), Museum of
Göttingen, Germany.
TYPE LOCALITY. Doberg, near Bünde, Germany.
AGE. Late Oligocene.
TAXONOMIC PLACEMENT. Hemipatagus; this species exhib-
its all the features typical for Hemipatagus and is actually
rather similar to H. hoffmanni; it differs from the latter spe-
cies by its lower profile, prominent keel in interambulac-
rum 5, more eccentric peristome and higher number of en-
larged primaries compared to H. hoffmanni of similar size;
Ebert (1889) also saw similarities with H. ocellatus (however,
part of the illustrations he refers to actually depict Lovenia
duncani (namely those of Wright 1864)).
mascareignarum Michelin, 1863
∗
1863 Hemipatagus Mascareignarum N.; Michelin in
Maillard: A 6; pl. 16, fig. 2a–i.
HOLOTYPE. Specimen figured by Michelin (1863: pl. 16,
figs 2a–d); whereabouts unknown.
TYPE LOCALITY. Réunion Island, Indian Ocean.
AGE. Recent.
TAXONOMIC PLACEMENT. Maretia; typical Maretia with
long, straight petals, characteristic distribution of the aboral
primary tubercles (also present in the posterior part of the
 Hemipatagus – a misinterpreted Loveniid
interambulacra 1a and 4b, along the adradial suture), oval,
outline with only faint frontal notch, low profile, oval sub-
anal fasciole, well developed phyllodes and so on. Obviously
Michelin was not aware of the genus Maretia, as he states
‘Le spatangus planulatus de Lamarck, qui se trouve dans
la mer Rouge, et subfossile sur ses bords, est une espèce
d’hemispatangus, vivant aujourd’hui’ (1853: A 6); according
to Mortensen (1951: 38–39) this species is a junior synonym
of M. planulata (Lamarck).
mauritanica Pomel, 1885
∗ Eupatagus here (contrary to Jeannet & Martin (1937: 277),
1885 Sarsella mauritanica; Pomel: 16, pl. 1, figs 6–7;
pl. 2, figs 5–6 [not seen].
1885 Sarsella mauritanica, Pomel, 1885; Cotteau et al.:
36–39; pl. 1, figs 4–8.
1893 Sarsella mauritanica, Pomel, 1885; Cotteau: 667–
670; pl. 362, figs 1–6.
TYPE LOCALITY. Kef Iroud, Dépt. d’Alger, Algeria.
AGE. Eocene or Oligocene.
TAXONOMIC PLACEMENT. Lovenia or Hemipatagus;
without re-examining the types or topotypic material it is
impossible to decide to which of the two genera the species
belongs; none of the authors that have studied the specimens
could confirm the presence or absence of a internal fasciole
(e.g. Cotteau et al. 1885: 37–38); Contrary to the figures in
Cotteau’s Paléontologie Française (1893; pl. 362, figs 1–6),
the plastron is not fully tuberculate (compare description and
illustration in Cotteau et al. 1885); Cotteau et al. (1885) state
explicitly that the aboral primary tubercles are crenulated in
this species (which can also be seen on their pl. 1, fig. 5), a
feature which does not occur in extant Lovenia according to
Mortensen (1951); however, it seems that many ambulacral
plates reach within the subanal fasciole (5 or 6, Cotteau et al.
1885: pl. 1, fig. 6), a common feature in Lovenia.
martini Gerth, 1922
∗
1922 Eupatagus martini; Gerth: 511–512; pl. 52, figs 1,
1a.
1975 Eupatagus rostratus rostratus d’Archiac;
Henderson: 44–45.
HOLOTYPE. Specimen St.4286 (figured by Gerth 1922); pa-
laeontological collection of the Rijksmuseum van Geologie
en Mineralogie, Leiden.
TYPE LOCALITY. Goenoeng Spolong, Java.
OCCURRENCE. ? Eocene, Oligocene to Miocene of India,
Japan and Java.
TAXONOMIC PLACEMENT. Eupatagus; junior synonym of E.
rostratus rostratus d’Archiac, 1850 according to Henderson
(1975: 44).
mojsvari Böhm, 1882
∗
1882 Brissomorpha mojsvari n. sp.; Böhm: 364–365;
pl. 2, figs 1a–d.
HOLOTYPE. Specimen figured by Böhm (1882: pl. 2,
figs 1a–d); whereabouts unknown.
187
TYPE LOCALITY. Sepocloc, northern coast of Madura, N of
Java.
AGE. ? Neogene.
TAXONOMIC PLACEMENT. ? Eupatagus; this species has
been referred to a variety of genera by different authors: Eu-
patagus by Gerth (1922: 512), Brissoides by Wanner (1931:
455) and Maretia by Jeannet & Martin (1937: 277); these dif-
ferent opinions obviously result from the poor illustration in
Böhm; as Böhm (1882: 365) explicitly mentions the presence
of a peripetalous fasciole this species is tentatively referred to
who assume that a peripetalous fasciole is lacking based on
the distribution of the aboral primary tubercles in Böhm’s
figure).
nicklesi Cotteau, 1890
∗
1890 Maretia Nicklesi, Cotteau, 1889; Cotteau: 6–7;
pl. 33 (12), figs 7–10.
HOLOTYPE. Specimen figured by Cotteau (1890); Nicklès
collection.
TYPE LOCALITY. Callosa, Alicante, Spain.
AGE. ? Eocene.
TAXONOMIC PLACEMENT. Eupatagus; based on labrum
shape, fully tuberculate sternum and poorly developed phyl-
lodes (Mortensen, 1951: 26).
nicoleti Agassiz, 1839
∗
1839 Spatangus Nicoleti Ag.; Agassiz: 33; pl. 4, figs 7–8.
HOLOTYPE. Specimen figured by Agassiz (1839).
TYPE LOCALITY. La Chaux-de Fonds, Jura, Switzerland.
AGE. Burdigalian.
TAXONOMIC PLACEMENT. Hemipatagus; a junior synonym
of H. ocellatus according to Agassiz & Desor (1847: 7) and
Philippe (1998: 222).
ocellatus Defrance, 1827
∗
1827 Spatangus ocellatus nov. sp.; Defrance in Blainville
et al.: vol. 50, 96–97 [not seen, fide Philippe, 1998:
220].
1998 Maretia ocellata (Defrance, 1827); Philippe: 220–
223; pl. 22, figs 6, 7, 8a–b, 9a–b, 10a–c.
2005 Hemipatagus ocellatus (Defrance, 1827); Kroh:
191–194, fig. 94, pl. 81, figs 1–4, pl. 82, figs 1–
2.
HOLOTYPE. Whereabouts unknown.
TYPE LOCALITY. Saint-Paul-Trois-Châteaux, Rhône Basin,
France.
AGE. Burdigalian.
TAXONOMIC PLACEMENT. Hemipatagus; this well-known
species shows all features characteristic of Hemipatagus: a
cordate outline, camellate primary tubercles on the aboral
188 A. Kroh
and oral side, a conspicuous field of tubercles in adapical in-
terambulacra 2 and 3, a bilobed subanal fasciole. . . ; Philippe
(1998) recognised the differences to ‘true’ representatives of
Maretia and expressed his reservations concerning that gen-
eric attribution.
omali Galeotti, 1837
TYPES AND PROVENANCE. Unknown.
TAXONOMIC PLACEMENT. Doubtful; according to Cotteau
(1885) this species is synonymous with ‘Spatangus’ grignon-
ensis Cotteau, 1880.
ovata Leske, 1778
∗
1778 Spatangus ovatus; Leske: 188; pl. 49, figs 12–13
[not seen; fide Mortensen, 1951].
TYPES AND PROVENANCE. Unknown.
TAXONOMIC PLACEMENT. Doubtful; according to
Mortensen (1951: 37) the identity of this species is
unclear; Leske’s illustrations were based on those from
Seba’s Thesaurus III (pl. 15, figs 27–28), which according
to Mortensen are rather poor and of unknown affinity; He
strongly argues against synonymising this species with
Maretia planulata as proclaimed by other authors (e.g. Clark
1925).
pareti Manzoni, 1879
∗ TYPE LOCALITY. 26 miles south-west of cape Everard,
1879 Maretia Pareti Manzoni; Manzoni: 158–159; pl. 1,
figs 1–2; pl. 2, fig. 18; pl. 4, figs 33–39.
TYPE MATERIAL. Specimens figured by Manzoni (1879).
TYPE AREA. ‘Colline de Bologna e di Aucona’, Italy.
AGE. ? Miocene.
TAXONOMIC PLACEMENT. Type species of Mazettia
Lambert & Thiéry in Lambert, 1915 (replacement for Manzo-
nia Pomel, 1883 preoccupied by Manzonia Brusina, 1870),
regarded as subgenus of Prospatangus Lambert, 1902 by
Lambert (1915); current status unclear, but certainly not
closely related to either Hemipatagus or Maretia (based on
the fully tuberculate sternal plates, the en-chevron arrange-
ment of the aboral primary tubercles, the presence of primary
tubercles in all aboral interambulacra and the different shape
(presence of a frontal sinus in combination with an oval,
elongated outline and long, straight petals)).
parvituberculata H. L. Clark, 1924
∗
1924 Maretia parvituberculata; Clark: 13–15; pl. 4,
figs 1–5 [not seen].
1925 Maretia parvituberculata; Clark: 227.
1950 Eurypatagus parvituberculatus (H. L. Clark);
Mortensen: 260–261: pl. 17, figs 1, 5–7; pl. 18,
figs 9, 11; pl. 19, figs 2, 11, 19.
TYPE LOCALITY. South Africa.
AGE. Recent.
TAXONOMIC PLACEMENT. Eurypatagus (Mortensen 1950:
260).
pellati Cotteau 1863
∗
1863 Hemipatagus Pellati, Cotteau, 1863; Cotteau: 150.
1885 Maretia Pellati (1863), Cotteau, 1885; Cotteau: 28–
30; pl. 2, figs 7–11.
HOLOTYPE. The specimen described by Cotteau (1863); col-
lection of Pellat.
TYPE LOCALITY. Biarritz, Basses-Pyrénées, France.
TYPE OCCURRENCE. Couche à Eupatagus ornatus.
AGE. Late Eocene.
TAXONOMIC PLACEMENT. Hemipatagus (this paper); based
on the cordate outline, the presence of a distinct frontal notch,
rudimentary pores in the adapical part of the anterior porifer-
ous zones of the anterior paired petals, and the presence and
distribution of the primary tubercles; single known specimen
is rather small and very similar to juvenile H. desmoulinsi
from the same area.
peloria Clark, 1916
∗
1916 Maretia peloria; Clark: 121; pl. 44, figs 1–3 [not
seen].
1917 Maretia peloria; Clark: 248–249; pl. 146, fig. 25
[illustration of pedicellaria].
1951 Paramaretia peloria (H. L. Clark); Mortensen: 51–
52.
Victoria, Australia; ∼ 165 m water depth.
AGE. Recent.
TAXONOMIC PLACEMENT. Paramaretia (Mortensen 1951).
pendulus Agassiz, in Agassiz & Desor, 1847
∗
1847 Spatangus pendulus Agass.; Agassiz & Desor: 8.
1880 Hemispatagus [sic] pendulus (Agassiz), Desor; de
Loriol: 133–135; pl. 11, figs 7, 7a–c.
TYPE MATERIAL. Two poorly preserved syntypes in the Mu-
seum of Paris (probably not conspecific according to de
Loriol 1880).
TYPE AREA. Sinai Peninsula, Egypt.
AGE. ? Eocene.
TAXONOMIC PLACEMENT. ? Eupatagus; the specimen
figured by de Loriol (1880) clearly seems to be a Eupatagus
based on the shape of the petals, the size and distribution of
the enlarged primary tubercles; the presence of a peripetal-
ous fasciole is not confirmed but seems highly probable; the
identity of the second syntype remains dubious.
perornata Schaffer, 1912
∗
1912 Spatangus (Maretia) perornatus Schff.; Schaffer:
190–191; pl. 59, fig. 4–6.
SYNTYPES. 2 syntypes (KM F/0127, F/0128), figured in
Schaffer (1912: pl. 59, figs 4–6); housed at the Krahuletz-
Museum, Eggenburg, Austria.
 Hemipatagus – a misinterpreted Loveniid 189

TYPE LOCALITY. Eggenburg (Kremserberg), lower Austria.
TYPE OCCURRENCE. Zogelsdorf Formation.
AGE. Late Eggenburgian (Early Burdigalian), Early Mio-
cene.
TAXONOMIC PLACEMENT. Hemipatagus; junior synonym of
H. ocellatus (Desmarest) (see Kroh 2005).
planulatus Lamarck, 1816
∗ b.
1816 Spatangus planulatus; Lamarck: 31.
1951 Maretia planulata (Lamarck); Mortensen: 27–39;
pl. 4, figs 1–5, 10–11; pl. 5, figs 3–17; pl. 44,
figs 1–4, 18, 26–27, 29, 32–33.
TYPE MATERIAL. Probably at the Museum Histoire Naturelle
Paris.
(1937: 277); Herklots (1854: 11) mentioned a second speci-
men (no. 452b; pl. 4, figs 5, 5a) which he tentatively referred
to his S. praelongus, but which, in fact is indeterminable.
pulchellus Herklots, 1854
∗
1854 Spatangus pulchellus, nouv. esp.; Herklots: 12;
pl. 4, figs 7, 7a–b.
1858 [Hemipatagus] pulchellus; Desor: 418.
1880 Maretia? pulchella Herkl. spec.; Martin: 81.
1922 Maretia? pulchella (Herkl.); Gerth: 512, figs 9, 9a–
1937 Eupatagus (s. Brissoides) pulchellus (Herklots);
Jeannet & Martin: 273–274, figs 50a–b.
HOLOTYPE. The specimen figured by Herklots (1854: pl. 4,
figs 7, 7a–b), no. 447 (the same number is listed under S.
affinis).

TYPE AREA. ‘les mers australes’. TYPE LOCALITY. Tjidamar, Java.

AGE. Recent. AGE. Pliocene.

OCCURRENCE. South-western Red Sea; Indian Ocean and TAXONOMIC PLACEMENT. Eupatagus; Lambert & Thiéry
western Pacific, from Eastern Africa to Fiji and Gilbert (1924: 451, 458) listed this species both under the pre-
Islands, and from Sagami Bay (Japan) to Port Jackson Linnaean genus Brissoides and under Maretia; the oval,
(Australia) anteroposteriorly elongated outline, lack of a frontal sinus,
shape of the petals, fully tuberculate plastron, restriction of
TAXONOMIC PLACEMENT. Type-species of Maretia; Indian the aboral primary tubercles to the area between the petals
Ocean populations are usually whitish to yellow, Malayan and the presence of a peripetalous fasciole indicate an attri-
specimens are dark coloured, but show no morphological bution to the genus Eupatagus.
differences according to Mortensen (1951: 36).
regiomontanus Mayer, 1860
planulata abbassi Ali, 1985 ∗
1860 Hemispatangus [sic] Regiomontanus Mayer;
∗
1985 Maretia planulata abbassi n. sp.; Ali: 294–295; Mayer: 11 [pagination of the reprint].
figs 12A–B.
TYPE MATERIAL. Current whereabouts unknown.
TYPE MATERIAL. USNM 339748.
TYPE LOCALITY. Kleinkuhren, north-west of Königsberg,
TYPE LOCALITY. Wadi Abu Abraiki, Western Egypt. Samland, Russia.
AGE. Early Pliocene. AGE. Late Eocene.
TAXONOMIC PLACEMENT. Maretia; very similar to extant TAXONOMIC PLACEMENT. Hemipatagus?; poorly known;
M. planulata. a junior synonym of H. sambiensis according to Cotteau
(1885).
praelongus Herklots, 1854
∗
1854 Spatangus praelongus, nouv. esp.; Herklots: 11–12;
rotundus Duncan & Sladen, 1884
pl. 2, fig. 6. ∗
1884a Macropneustes roundus, Duncan & Sladen;
1858 [Hemipatagus] praelongus; Desor: 418. Duncan & Sladen: 232–233; pl. 38, figs 6–7.
1880 Maretia planulata Gray; Martin: 81. 1924 Hemipatagus rotundus Duncan & Sladen (Mac-
1924 Atelospatangus praelongus; Lambert & Thiéry: ropneustes); Lambert & Thiéry: 457.
458. 2004 Macropneustes (Macropneustes) roundus (Duncan
1937 Maretia planulata (Lamarck); Jeannet & Martin: & Sladen) n. comb.; Srivastava: 147–148; pl. 7, figs
277–278. 12–13.
HOLOTYPE. The specimen figured by Herklots (pl. 2, fig. g), HOLOTYPE. Geological Survey of India no. G302/100 (GSI
no. 433. Type No. 2685 (Srivastava 2004)).
TYPE LOCALITY. Tjidamar, Java. TYPE LOCALITY. Teyón (or Tiyún), east of Chorla, Sind,
India.
AGE. Pliocene.
LITHOSTRATIGRAPHY. Kithar Series.
TAXONOMIC PLACEMENT. Maretia; junior synonym of the
extant Maretia planulata according to Jeannet & Martin AGE. Middle Eocene.
190 A. Kroh
TAXONOMIC PLACEMENT. Eupatagus or Macropneustes
(Macropneustes); Duncan & Sladen (1884a) expressed their
problems concerning the generic attribution of this species
and named both genera mentioned above as potential candi-
dates. They choose Macropneustes because of the numerous
and prominent development of the aboral primary tubercles
and the inferred absence of the subanal fasciole (specimen
damaged posteriorly). Neither Henderson (1975: 33) nor
Srivastava (2004: 147–148) state why they changed the gen-
eric attribution and, apparently, no additional specimens have
been found since the discovery of the types. Until more spe-
cimens are described the generic attribution of this species
remains tentative.
saccoi Airaghi, 1905
∗
1905 Maretia Saccoi n.sp.; Airaghi: 53–54; pl. figs 21–
22.
HOLOTYPE. The specimen figured by Airaghi (1905: pl.
fig. 21–22); Museo geologico Torino, Italy.
TYPE LOCALITY. C. Dogana, Umbria, Italy.
AGE. Miocene.
TAXONOMIC PLACEMENT. Hemipatagus; typical Hemi-
patagus with the characteristic arrangement of the aboral
primary tubercles (missing in posterior part of interambulac-
ral columns 1a and 4b), typical petals with rudimentary pores
in the anterior poriferous zones of the anterior paired petals,
cordiform outline and strong frontal sinus; in Airaghi’s de-
scription no fascioles are mentioned, should an internal fas-
ciole be discovered in the type material this species would
have to be transferred to Lovenia.
sambiensis Beyrich, 1848
∗
1848 Spatangus Sambiensis n. sp.; Beyrich: 100
[pagination of the reprint].
1883 Maretia Sambiensis Beyr. sp.; Noetling: 688.
1885 Maretia sambiensis (Beyrich), Noetling, 1883;
Cotteau: 43.
TYPE MATERIAL. Current whereabouts unknown.
TYPE LOCALITY. Between Warnicken and Groß-Kuhren,
north-west of Königsberg, Samland, Russia.
AGE. Late Eocene.
TAXONOMIC PLACEMENT. Hemipatagus ?; poorly known;
H. regiomontanus Mayer, 1860 is a junior synonym of this
species according to Cotteau (1885).
savini Cotteau, 1893
∗ ∗
1893 Maretia Savini, Cotteau, 1893; Cotteau: 637–639;
pl. 354, figs 1–4.
TYPE MATERIAL. Two poorly preserved specimens; collec-
tion Savin, current whereabouts unknown.
TYPE LOCALITY. Vallée du Trapel, Aude, France.
AGE. Middle Eocene.
TAXONOMIC PLACEMENT. Doubtful; referred to Leiop-
neustes by Lambert & Thiéry (1924: 448); type material
poorly preserved, large parts of the aboral and oral surface
are missing; referred to Maretia by Cotteau (1893) on the
basis of the missing peripetalous fasciole; the features that
can be observed, however, are in favour of Eupatagus (oval,
elongated outline; very shallow frontal sinus; non-camellate
primary tubercles; arrangement of primary tubercles).
soubellensis Peron & Gauthier, in Cotteau et al.,
1891
∗
1891 Maretia soubellensis, Peron & Gauthier; Cotteau
et al.: 81–83; pl. 1, fig. 3.
HOLOTYPE. Specimen figured by Cotteau et al. (1891: pl. 1,
fig. 3); collection Peron, current whereabouts unknown.
TYPE LOCALITY. Foum-Soubella, south of Sétif, Dépt. Con-
stantine, Algeria.
AGE. ? Middle Miocene (‘Langhien’ of Cotteau et al., 1891).
TAXONOMIC PLACEMENT. Hemipatagus; Cotteau et al.
(1891) provide no information on the presence or absence
of an internal fasciole, but mention in their discussion that
this species is similar to H. ocellatus and explicitly state that
the latter lacks an internal fasciole, even in perfectly pre-
served specimens; the cordate shape, the pronounced frontal
sinus, the rudimentary pores in the adapical poriferous zone
of ambulacra II and IV, as well as the distribution of the
primary tubercles suggest an attribution to Hemipatagus.
subrostratus Clark, in Clark & Twitchel, 1915
∗
1915 Hemipatagus subrostratus Clark, n. sp.; Clark &
Twitchel: 151; pl. 49, figs 2a–b.
1959 Maretia subrostrata (Clark); Cooke: 81; pl. 34, figs
5–6.
HOLOTYPE. USNM 164652; U.S. National Museum, Smith-
sonian Institution, Washington, USA.
TYPE LOCALITY. Wilmington, North Carolina, USA.
TYPE OCCURRENCE. Castle Hayne Limestone.
AGE. Late Middle Eocene.
TAXONOMIC PLACEMENT. Hemipatagus?; generic attribu-
tion uncertain but outline, distribution of primary tubercles
and shape of the petals suggest the attribution to Hemi-
patagus; according to Cooke (1959: 81) this species is very
similar to H. desmoulinsi Cotteau (1885: 26; pl. 2, figs 1–6).
tenuis Peron & Gauthier, in Cotteau et al., 1891
1891 Maretia tenuis, Peron & Gauthier; Cotteau et al.:
79–81; pl. 1, fig. 2.
HOLOTYPE. Specimen figured by Cotteau et al. (1891: pl. 1,
fig. 2); collection Peron, current whereabouts unknown.
TYPE LOCALITY. Oued Sebt, near Tizi-Ouzou, Algeria.
AGE. ? Middle Miocene (‘Langhien’ of Cotteau et al., 1891).
 Hemipatagus – a misinterpreted Loveniid
TAXONOMIC PLACEMENT. Hemipatagus; according to
Cotteau et al. (1891) no traces of an internal fasciole could
be found; the cordate shape, the pronounced frontal sinus,
the rudimentary pores in the adapical poriferous zone of am-
bulacra II and IV, as well as the distribution of the primary
tubercles suggest an attribution to Hemipatagus.
tuberculata Agassiz & Clark, 1907
∗ oles, none are ascertained due to the heavy abrasion of their
1907 Maretia tuberculata A. Ag. & Clark; Agassiz &
Clark: 134.
1917 Maretia tuberculata; Clark: 246; pl. 160, figs 5–7.
HOLOTYPE. Single immature specimen.
TYPE LOCALITY. Eastern channel, Korea Strait, west of Shi-
monoseki, Japan; 108 m water depth (59 fathoms).
AGE. Extant.
TAXONOMIC PLACEMENT. Doubtful (juvenile specimen);
Mortensen (1951: 45) refers this species to Maretia, but an
attribution to Lovenia is also not improbable (the holotype
has 7 ambulacral plates reaching into the subanal fasciole
(2×5 tube feet within the subanal fasciole) and no gono-
pores developed at a size of 26 mm; in Maretia planulata
the gonopores open at a size of 18 mm and it has 4 ambulac-
ral plates reaching into the subanal fasciole (2×2 tube feet);
in M. cordata the gonopores open at 13 mm test length; in
Lovenia scaber, by contrast, the gonopores open at 30 mm).
tuberculatus Zittel, 1867
∗ 1924
1867 Hemipatagus tuberculatus Zitt.; Zittel: 63–64; pl.
12, figs 1a–c.
1894 Lovenia tuberculata; Tate: 126 [not seen].
1975 Lovenia tuberculata (Zittel); Henderson: 31–32;
pl. 4, figs 5–6; pl. 5, figs 3–6.
LECTOTYPE. NHMW 1959/335/51, figured by Zittel (1867:
pl. 12, figs 1a–c), designated by Henderson (1975: 31); Nat-
ural History Museum Vienna, Austria.
PARALECTOTYPE. NHMW 1970/1338.
TYPE LOCALITY. Cape Farewell, west Nelson, New Zealand.
TYPE OCCURRENCE. ? Abel Head Formation.
AGE. Late Oligocene.
OCCURRENCE. Whaingaroan to Altonian (Oligocene to
Early Miocene) of New Zealand.
TAXONOMIC PLACEMENT. Lovenia.
tuberculatus Peron & Gauthier, in Cotteau et al.,
1885
∗ seum, London.
1885 Tuberaster tuberculatus, Peron & Gauthier, 1885;
Cotteau et al.: 47–49; pl. 3, figs 1–4 [= H. gauthieri
Lambert in Lambert & Thiéry, 1924: 457 (nom. nov.
pro Tuberaster tuberculatus Péron & Gauthier, in
Cotteau et al., 1885 non Zittel, 1867); see above].
HOLOTYPE. Specimen figured by Cotteau et al. (1885: pl. 3,
figs 1–4); collection Gauthier.
191
TYPE LOCALITY. Kef Iroud, Algeria.
AGE. Oligocene.
TAXONOMIC PLACEMENT. Hemipatagus?; apart from the
fully tuberculate sternum (if correctly figured in Cotteau
et al. (1885)) this species is very similar to typical Hemi-
patagus species (like H. hoffmanni); although Cotteau et al.
(1885) debate much about the presence and absence of fasci-
material.
tuberosus Fraas, 1867
∗
1867 Eupatagus tuberosus Frs.; Fraas: 279; pl. 6, fig. 8.
1880 Euspatangus [sic] tuberosus, Fraas; de Loriol: 141–
142; pl. 11, fig. 5.
HOLOTYPE. Fragmentary specimen figured by Fraas (1867);
Museum Stuttgart, Germany.
TYPE LOCALITY. Ouadi-el-Tih, near Caire, Egypt.
AGE. ? Eocene.
TAXONOMIC PLACEMENT. Doubtful; the holotype is a frag-
ment of the aboral side preserving a petal (I, or more likely
IV) and the adjoining interambulacra which bear camelate
tubercles; currently impossible to decide whether it is a Love-
nia, Hemipatagus or something else.
twitchelli Lambert, in Lambert & Thiéry, 1924
Maretia twitchelli Lambert in Sanchez Roig;
Lambert & Thiéry: 458 [nom. nov. pro specimen
C of Eupatagus floridanus Clark, 1915 (in Clark &
Twitchell 1915: 176–177; pl. 83, figs 2a–d)].
HOLOTYPE. Specimen figured by Clark & Twitchell (1915:
pl. 83, figs 2a–d); Museum of Comparative Zoology, Har-
vard, USA.
TYPE LOCALITY. Levy County, Florida, USA.
TYPE OCCURRENCE. Vicksburg Group.
AGE. Early Oligocene.
TAXONOMIC PLACEMENT. Eupatagus; junior synonym of
Eupatagus antillarum (Cotteau) according to Henderson
(1975: 33).
variegata Gray, 1866
∗
1866 Spatangus (Maretia) variegatus, n.sp.; Gray: 170–
171.
TYPE MATERIAL. Two specimens in the Natural History Mu-
TYPE LOCALITY. Pulo Taya, China Sea.
AGE. Recent.
TAXONOMIC PLACEMENT. Maretia; according to Mortensen
(1951: 36) M. variegata is a dark coloured variant of M.
planulata and synonymous with that species.
192 A. Kroh
vicentina Dames, 1878
∗ Victoria, Australia.
1878 Breynia vicentina nov. sp.; Dames: 75–76; pl. 7,
figs 7a–b.
HOLOTYPE. Specimen figured by Dames (1878: pl. 7, figs
7a–b); current whereabouts unknown.
TYPE LOCALITY. Longio, Vicentin, Northern Italy.
AGE. ? Eocene.
TAXONOMIC PLACEMENT. ? Lovenia; Dames could not ob-
serve any fascioles on the single specimen available, but
inferred the presence of an internal fasciole on the basis of
the absence of any pore pairs in the adapical ambulacra;
the figures provided by Dames do not fully fit his descrip-
tion (the whole adapical area seems to be missing according
to the figure, whereas he mentions details of the adapical area
in the description; likewise he reports a naked plastron, while
the figures show that the plastron is largely missing); it is un-
clear why Lambert & Thiéry (1924: 466) refer this species
to Hemipatagus; the available information rather suggests
an attribution to Lovenia, providing Dames’ description is
correct.
woodsii Etheridge, 1875
∗
1875 Hemipatagus Woodsii, sp. nov.; Etheridge: 445–
447, pl. 21, figs 1–7.
1877 Lovenia Forbesi, var. Woodsi, Etheridge; Duncan:
44, 56–61.
1994 Lovenia woodsii (Etheridge, 1875); Irwin & Arch-
bold: 6–10, figs 4A–C, 6A–D, 7A–M [cum syn].
LECTOTYPE. Specimen F17500, figured by Etheridge (1875:
pl. 21, figs 1–7), designated by Irwin & Archbold (1994: 6,
fig. 6A–D); Australian Musuem, Sydney.
TYPE LOCALITY. Beaumaris, east shore of Port Phillip Bay,
TYPE OCCURRENCE. Black Rock Sandstone, Brighton
Group.
AGE. Cheltenhamian (Messinian), Late Miocene.
OCCURRENCE. Early (?) Miocene – Pliocene (Zanclean) of
the Port Phillip Basin, Victoria, Australia; possibly also in
Tasmania.
TAXONOMIC PLACEMENT. Lovenia.
zeisei Gagel, 1905
∗
1905 Maretia Zeisëi spec. nov.; Gagel: 541–543; pl. 24,
fig. 6; pl. 25, figs 3a–c, 4, 5.
SYNTYPES. Four poorly preserved specimens in the collec-
tions of the Geological Survey of Preussen, and the Museum
of Lübeck, Germany.
TYPE LOCALITY. Zarrentin, south-east Holstein, Northern
Germany.
AGE. Erratic boulders of Cenozoic age, presumably Mio-
cene.
TAXONOMIC PLACEMENT. ? Hemipatagus; based on the
outline and the distribution of the aboral primary tubercles;
type material very poor and in need of re-description; junior
synonym of H. gottschei according to Lambert & Thiéry
(1924: 457); or juveniles of H. hoffmanni as discussed by
Gagel (1905: 543; he rejected this hypothesis because of the
naked plastron of H. zeisei, which could, however, also be
related to the poor preservation).