Hemipatagus, amisinterpreted Loveniid(Echinodermata: … · Hemipatagus, amisinterpreted...

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 thatis now usually treated as a subjective synonym of the extant genus Maretia, but was originallysubject to considerable dispute within the scientific community. Restudy of the species attributedto Hemipatagus and a range of presumably related spatangoids including Lovenia and Maretia hasbeen carried out to solve the problem of its relationships. Cladistic analysis shows that Hemipatagusis 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 suggestthat in Hemipatagus an internal fasciole is present in early ontogeny, but lost in the adults. A cladecomprising 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.

* Corresponding author. E-mail address: [email protected]

Contents

Introduction 163

History of nomenclature 164

Material and methods 164Abbreviations and repositories 167

Results 167Status of Hemipatagus 167Phylogenetic affinities of Hemipatagus 169Cladistic analysis 171Spatial and temporal distribution of Hemipatagus and Maretia 172

Systematic Palaeontology 173Order Spatangoida Claus, 1876 173

Family Loveniidae Lambert, 1905 173Genus Hemipatagus Desor, 1858 173

Family Spatangidae Gray, 1825 174Subfamily 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

Spatangoids are echinoids characterised by their bilaterallysymmetrical corona and highly modified ambulacra form-ing a complex respiratory apparatus. They are well adaptedto a life as burrowers and ploughers in mobile sediments,

exploiting organic detritus as their prime food resource.Due to their life habit and evolutionary success they arethe most diverse of all extant sea urchin groups and one ofthe echinoid clades with the best fossil record (Kier 1977).Their highly adapted test morphology provides many fea-tures that can be employed in classification. Consequentlyspatangoid taxonomy is, in general, rather straightforward.

164 A. Kroh

Particular features, however, have recently been shownto display high levels of convergence and homoplasy(Markov & Solovjev 1995; Neraudeau et al. 1998; Villieret al. 2004; Smith & Stockley 2005). Indeed, spatangoidphylogeny 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 oftheir burrows and protects the respiratory tube feet from finemud (see Lawrence 1987). Unfortunately, fascioles were de-veloped several times during spatangoid evolution and canbe secondarily lost (Villier et al. 2004; Smith & Stockley2005) and the emphasis placed on these structures has led toconsiderable confusion and problems in spatangoid system-atics. The problem investigated here represents a case wheretoo much emphasis has been placed on presence and absenceof fascioles.

Hemipatagus (Pl. 1) is a genus widely distributed in theEocene to Miocene of Europe, occurring abundantly in thefamous Doberg section in northern Germany (Oligocene)and in Early Miocene strata of the Rhone Basin (Philippe1998), the Aquitaine Basin and the central Paratethys (Kroh2005). It is a spatangoid characterised by its heart-shapedoutline and heterogeneous aboral tuberculation. The phylo-genetic and systematic relationship of Hemipatagus withinthe spatangoids is uncertain, although it has commonly beensynonymised with Maretia (Pl. 2, figs 1 & 2), a Pliocene toextant inhabitant of the Indo-West Pacific. The complicatednomenclatorial 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 aphylogeny and revised taxonomy for Hemipatagus and itsrelatives.

History of nomenclature

The genus Hemipatagus was established in 1858 by Desorfor the species Spatangus hoffmanni Goldfuss, 1829 (Fig. 1).Subsequently, Agassiz (1873: 568) placed Hemipatagus insynonymy 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) discussedthis matter at length and finally considered HemipatagusDesor, 1858 a junior synonym of Maretia Gray, 1855. Hisopinion was based on the observation that only one of thethree features discussed in the literature seemed importantto him (the presence of internal ampullae associated withspine tubercles in Hemipatagus). He argued that one featurealone was not enough to justify separation of the two gen-era. Moreover, he had also observed internal swellings thathe assumed were ampullae in juveniles of Maretia planu-lata (Lamarck, 1816) and saw them as proof for his point ofview. 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 thisfeature, due to the fact that large differences in the depth of thefrontal notch are also present in other genera, e.g. in Lovenia);

(2) the broader, bilobed subanal fasciole in Hemipatagus (healso 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 EchinoidDirectory’) noted the high similarity between Hemipatagusand Lovenia and proposed its inclusion in the Loveniidae.

Part of the confusion surrounding Hemipatagus andits relationship to Maretia and Lovenia resulted fromDesor’s (1858) statement that no fascioles are present inHemipatagus. Laube (1869) and Etheridge (1875), however,showed that a ‘spectacle-shaped’ (bilobed) subanal fascioleis present (see Fig. 2). Another major reason for the differingopinions resulted from the habit of basing conclusions on fea-tures observed in various non-type species attributed to eithergenus (e.g. Duncan 1877). The type species of Hemipatagusand Maretia, in contrast, were rarely investigated with regardto this question. Furthermore, most authors placed a heavyemphasis 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 lostduring ontogeny, being present in juveniles and absent in theadults of many spatangoids (e.g. Meoma (partial loss of thesubanal fasciole: Kier & Grant 1965; Chesher 1969), Para-maretia, Platybrissus and Eurypatagus (complete loss of thesubanal fasciole: Mortensen 1950, 1951); see also Neraudeauet al. 1998; Smith & Stockley 2005).

Although discussed at length it is not really clear whyMortensen (1951: 23–27) finally chose to synonymise Hemi-patagus with Maretia. It would have been understandable ifhe had synonymised Hemipatagus with Lovenia, as he statedhimself on p. 27 that ‘On the whole, the distinction betweenMaretia [here actually referring to Hemipatagus] and Love-nia is, in case of the fossils, very difficult, the presence orabsence of an inner fasciole – the only fully reliable distinc-tion between these two genera – being, rather impossible toascertain beyond doubt, if the preservation is not very fine.’ Itis difficult to understand why he rejected the shape of the sub-anal fasciole as a diagnostic feature in Hemipatagus, whilewidely using it in brissid genera, particularly, as this featureallows a very clear separation, with no known intermediatespecies. Mortensen (1951) also made a strong case againstthe 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 betweenHemipatagus and Maretia. Yet, it is well known that featuresthat are useful to separate some taxa might be highly variablein 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, 2ndZoological 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 preservedspecimens of Hemipatagus hoffmanni (Goldfuss, 1829)from the Oligocene of the type locality Doberg near Bunde,

Hemipatagus – a misinterpreted Loveniid 165

Plate 1 Oligocene and Miocene Hemipatagus. Figs 1–10, 16, Hemipatagus hoffmanni (Goldfuss, 1829), all Late Oligocene of Doberg, nearBunde, 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 Grubern, Austria. 11–12, NHMW 1914.VII.22b; 13–14,1914.VII.22a. Figs 15, Hemipatagus martensii (Ebert, 1889), Late Oligocene of Doberg, near Bunde, 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 Bunde, Germany, MB.E4840.


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, PalaeontologyDepartment MB.4839–4843, plus 6 unregistered speci-mens) were examined. In addition, the type-material ofHemipatagus hoffmanni was located (PIUB 357a-h) anda photograph of Goldfuss’s original specimen (PIUB357a) is presented for the first time (Fig. 1). Furthermore,the type material of Hemipatagus perornatus (Schaffer1912:2 syntypes – KM F/0127 and F/0128) was located andreinvestigated, along with some new specimens (NHMWGeology-Palaeontology Department 1914.VII.22a-c,1999z0049/0003, 2002z0159/0001; see also Kroh 2005).

Cladistic analysis was carried out using the programmePARS from the Phylogeny Inference Package PHYLIP(Felsenstein 2002), using the ‘Randomize input order ofspecies option’ (1000 replicates). The program CONSENSfrom the same package was used to obtain majority ruleconsensus trees. The analysis involved 30 species from theSpatangidae and Loveniidae together with Eupatagus valen-ciennesi Agassiz, in Agassiz & Desor, 1847. A total of 50characters were scored based on test shape, apical disc struc-ture, plating pattern, tuberculation, sphaeridia, pore and peri-proct structure. A number of additional, phylogeneticallyuninformative characters (basal synapomorphies) as well assome highly variable characters were excluded. The datamatrix and description of the remaining 42 characters arefound in Appendices 1 and 2. Due to the high degree of

morphological differentiation amongst the taxa investigatedmany characters were multi-state. Characters have neitherbeen polarised nor differentially weighted.

Abbreviations and repositories

GSI, Geological Survey of India; KM, Krahuletz Museum,Eggenburg, Austria; NHMW, Naturhistorisches MuseumWien, 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 andMaretia 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, thefollowing differences were observed in M. planulata: (1) thefrontal 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, Grubern, 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, Grubern, Austria. Interambulacra shaded, fasciolefinely stippled, small stippled circles are sphaeridial pits.

end is bluntly pointed not transversely truncated; (4) even inlarge specimens the primary tubercles do not develop trueampullae or strongly sunken areoles, especially not on theoral 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-posteriorlyelongated, not cordiform; (7) the subanal fasciole is oval andrather high instead of bilobed, very wide and low (Table1); (8) the labrum and the primordial plates of the interam-bulacra extend much further (Fig. 4A); (9) there are morephyllodal pores (14–15) in ambulacra II and IV (only 6–7in H. hoffmanni); (10) ambulacra II and IV are only slightlyconstricted ambitally, whereas they are strongly constricted

in H. hoffmanni (compare Figs 3 and 4A); (11) a conspicuousfield of coarse tubercles adjacent to adapical ambulacrum IIIis well developed in H. hoffmanni but lacking in M. planu-lata (Pl. 3, figs 1–5). These differences provide ample meansto separate those two genera. Lovenia and Hemipatagus aremore difficult to separate. Hemipatagus differs mainly by theabsence of the internal fasciole and its different petal struc-ture (never forming continuous arcs as in most species ofLovenia).

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 takeninto 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 aresphaeridial pits.


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 valueas a diagnostic feature. While some species of Hemipatagushave truly camellate tubercles (i.e. tubercles with associatedpouch-like ampullae forming a bulge on the interior ofthe test) on the oral and aboral surface (e.g. H. ocellatus,H. perornatus, H. girundicus), others lack them either overthe oral surface (e.g. H. hoffmanni) or completely (H. carolin-ensis, H. argutus). In those species where internal ampullaeare lacking, the areoles are, nevertheless, strongly sunken.However, the other features, especially the distribution of theaboral primary tubercles, number of phyllodal pores and allfeatures related to the plating pattern, are quite stable andallow 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 primarytuberculation, differentiation of fascioles, structure of the ap-ical disc and plastron structure (see Fischer 1966). As pointed

out by various authors recently (e.g. Jeffery 1998; Neraudeauet al. 1998; Smith & Jeffery 2000), in the past too much em-phasis was placed on the presence/absence of certain typesof fascioles in the adult animal in developing a supra-genericclassification of the spatangoids. It has been clearly estab-lished that fascioles may be lost during ontogeny, a fact thathas to be taken into account when trying to resolve thisquestion (e.g. Mortensen 1951; Kier & Grant 1965; Chesher1969).

Here the conspicuous field of tubercles in adapical in-terambulacral columns 2b and 3a (Pl. 3, figs 2–5) comesinto play. In all genera (except Hemipatagus) where this fea-ture is present it is invariably associated with an internalfasciole (Breynia, Echinocardium, Lovenia, Pseudoloveniaand Vasconaster). It is, therefore, possible to assume thatan internal fasciole was present during the early ontogenyof Hemipatagus. The fact that this arrangement is not welldeveloped in some species of Hemipatagus does not neces-sarily mean that these species never possessed an internalfasciole. There are some species of Lovenia in which thisspecial arrangement is not very conspicuous (e.g. L. sub-carinata). By contrast, such an arrangement has never beenobserved in any member of the family Spatangidae (includingMaretia).

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 Bunde, 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 Grubern, Austria. 5, 1914.VII.22a; 7, NHMW 1914.VII.22b.


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

On the basis of its strong similarity to Lovenia and theinferred 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 evaluationof the morphological features of Hemipatagus, Lovenia andMaretia, a cladistic analysis was carried out. In all analysesHemipatagus hoffmanni and H. ocellatus form a sister grouptaxon to the extant species of Lovenia. Exclusion of the spe-cies with a high number of unknown characters (Maretiacarinata, 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 secondclade containing the classical Loveniidae (Breynia, Ech-inocardium, Homolampas, Lovenia, Pseudolovenia) plusHemipatagus and Pseudomaretia (the latter traditionallyplaced in the Spatangidae: see Fischer 1966) (clade B). Due tothe presence of several missing characters in Maretia cordataand two old species tentatively placed in Hemipatagus (H.?argutus and H.? carolinus, from the Middle Eocene and theOligocene, respectively) 30 equally parsimonious trees werefound. The majority consensus tree of these trees is shownin Fig. 5A. The placement of M. cordata switches betweenjust 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 ofextant 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 ascrown-group loveniids forming a polyphyletic clade withPseudolovenia, H. hoffmanni, H. ocellatus and the extantspecies Lovenia. In just one of the 30 trees H.? argutus andH.? carolinus are resolved as sister group to Echinocardium.

Exclusion of M.? cordata, H.? argutus and H.? carolinusfrom the analysis results in a single most parsimonious tree(Fig. 5B) very similar to the majority consensus tree of theformer analysis but with better resolution within clade B. Itis apparent that the loveniids as a whole are a very robustgroup with well-defined monophyletic genera. The only ex-ception is Lovenia, which is monophyletic, but paraphyleticif Pseudolovenia is excluded. The present analysis suggeststhat 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 presentanalysis is very similar to that published by Madon (1994)using a different dataset. Another new result is the placementof the Pseudomaretia-clade in the loveniids. Earlier Pseudo-maretia (and the junior synonym Goniomaretia) were con-sidered to be spatangids (Fischer 1966) or maretiids. Thegenera Eurypatagus, Platybrissus and Paramaretia form adistinct clade in the present analysis and are a sister group toMaretia planulata in most trees. This clade appears also inthe strict consensus tree published by Stockley et al. (2005:fig. 2), where it also forms the sister group of Maretia. TheEurypatagus–Platybrissus–Paramaretia-clade is here namedEurypataginae (see ‘Systematic palaeontology’ below, for aformal 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 verysimilar topology to the trees discussed above.

The present analysis clearly shows that Hemipatagusis a loveniid that evolved during the Middle or Late Eocenefrom a common ancestor with Lovenia. Despite the superfi-cial similarities with Maretia it is clearly not closely relatedto that genus. This is also reflected by their temporal andspatial distributions, which are disjunct with respect to eachother (see below). Earlier statements that Maretia and Hemi-patagus are synonymous (e.g. Mortensen 1951) were basedon misinterpretation and erroneous weighting of a small setof characters.

In a recent paper on spatangoid phylogeny, Stockleyet al. (2005) present the results of a cladistic analysis in-volving several of the genera discussed here. In the strictconsensus 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 verysimilar topology to the one presented here when restrictedto the micrasterine clade alone. Smith relates the differencesbetween their fig. 2 and the present tree (Fig. 5) to the globalparsimony analysis employed in Stockley et al. (2005), whichfails to group the loveniid taxa into a single clade.

Spatial and temporal distribution of Hemipatagusand Maretia

Hemipatagus and Maretia are well separated in time andspace. The former occurs most commonly in the Eocene to


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–MalayanArchipelago and the West Pacific Islands (farthest eastoccurrences are on Hawaii). The only potential record ofHemipatagus from the Pliocene and from the Indo–West Pa-cific region is H.? bandaensis. The data available for thistaxon are not sufficient to firmly establish its generic affinity.It might also be a misidentified Lovenia. A re-description andnew illustrations are urgently needed.

Another issue apparent in Fig. 6 is the disjunct distri-butional pattern of the extant Maretia planulata. Apparentlythere are no records from Western India, Pakistan, the PersianGulf or the Indian Ocean coast of the Arabian Peninsula. Inview of this the colouration differences noted by Mortensen(1951: 36) may be more meaningful than he considered theywere. He described a whitish to yellow colouration in IndianOcean populations, versus Malayan specimens, which areusually dark. As Mortensen (1951: 36) could find no apparentmorphological differences, he considered this phenomenonto be local variation. Further investigations are needed tojudge whether there might actually be two species. A sim-ilar situation has recently been discovered in Asthenosomaby Weinberg & de Ridder (1998), who discovered that theechinoids previously classified as A. varium belong to twodistinct 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 proSpatangus 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 authorand date of the original spelling (e.g. des Moulins 1837:244; Agassiz & Desor 1847: 7), it has to be deemed to be ajustified emendation according to the ICZN (1999: 4th ednArticle 33.2.3.1.)).

RANGE. Eocene to Middle Miocene (Serravallian).

EMENDED DIAGNOSIS. Cordiform spatangoid differing fromother 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 theposterior part of interambulacral columns 1a and 4b (seeTable 1); (3) strongly sunken areoles or even the presence ofcamellae in some species; (4) presence of a subanal fasciolein 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 columns2b 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-

mens (Figs 3A, B); (7) moderately elongated labral plate notextending beyond the second ambulacral plates (Figs 3A, B);(8) prominent oral tubercles arranged in distinct rows, insome species with deeply sunken areoles and helically spir-alled parapet (e.g. Pl. 3, fig. 7); (9) three to four ambulacralplates extending into the subanal fasciole in ambulacrum Iand 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 publishedinformation: 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 belongto Hemipatagus, but which are too poorly known at presentto establish their generic affinities with certainty: H.? ar-gutus Clark, in Clark & Twitchell, 1915; H.? bandaensisMartin, in Jeannet & Martin, 1937; H.? carolinae (Gagel,1905); H.? gauthieri (Lambert, in Lambert & Thiery, 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, inClark & Twitchell, 1915; H.? tuberculatus (Peron &Gauthier, in Cotteau et al., 1885); H.? zeisei (Gagel, 1905).See Appendix 3 for original generic assignments and moredata on the individual species listed here.

REMARKS. There are a number of genera that show superfi-cial similarities to Hemipatagus. All of them, however, canbe clearly separated on the basis of structural features (seeTable 1). Lovenia differs by the presence of an internal fas-ciole, high number of ambulacral plates extending into thesubanal fasciole (up to 11, although in some species thereare only three or four as in Hemipatagus) and less lanceol-ate petals; in most species of Lovenia ambulacral columnsIb–IIa and IVb–Va form continuous arcs (not observed inany 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 shallowfrontal 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 andoral primary tubercles, the strongly crenulate nature of theaboral primary tubercles, the lack of a conspicuous fieldof coarse tubercles in adapical interambulacral columns 2band 3a, its oval to shield-shaped subanal fasciole, the longerprimordial 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, whilethey do not extend beyond the second ambulacral plate inHemipatagus or Lovenia), the labrum which extends to thethird or fourth ambulacral plate (not extending beyond thesecond 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-shapedform of the subanal fasciole, shape of the petals (with am-bulacral columns Ib–IIa and IVb–Va forming a continuousarc in some species), by the labrum and primordial plates ofthe 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 oralprimary tubercles, which are never sunken or camellate; andthe high number of ambulacral plates extending into the sub-anal fasciole (usually six to eight), by the distribution of theaboral primary tubercles (confined to the area within the peri-petalous fasciole, but reaching the tips of the posterior pairedpetals) and the near-straight contact between the sternal andepisternal plates.

Eupatagus differs by its fully tuberculate sternum, pres-ence of a peripetalous fasciole, oval shape of the subanalfasciole, the different nature of the aboral and oral primarytubercles (never camellate or strongly sunken), different dis-tribution of the aboral primary tubercles (see Table 1), by thelabrum and primordial plates of the paired interambulacrawhich extend to the third ambulacral plate and by the lack ofany frontal notch.

Pseudolovenia differs by its narrow, poorly-definedpetals with widely spaced, less well developed pores whichcontinue to the ambitus, restriction of the aboral primarytubercles 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 thehigher number of plates extending into the subanal fasciole(five).

Pseudomaretia and Goniomaretia differ by possessingan ethmolytic apical disc with only three gonopores (no gono-pore in genital plate 2), lack of a frontal notch, more reducednature of the anterior poriferous zones of the anterior pairedpetals (well developed pores only in the most distal plates),oval to shield-shaped subanal fasciole and the absence ofmost or all aboral primary tubercles.

Paramaretia differs by the more reduced nature of theanterior poriferous zones of the anterior paired petals (nolarge, well developed pores present at all), its labrum whichextends to the fourth ambulacral plates, the primordial platesextend to the fourth to sixth ambulacral plates, the loss of thesubanal fasciole in adults, having a plastron which is onlyweakly indented behind the first pair of episternal plates, itsnaked sternal plates and, most obviously, by the distribu-tion of the aboral primary tubercles which are present in allinterambulacra, 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) lackof any fascioles in adult individuals, (2) naked sternal plates

(partially tuberculate in Maretiinae), (3) structure of thepetals (narrowed, distally open, extending almost to the am-bitus) and (4) labrum extending to the 4th plates of ambulacraI and V. Differing from the Spatanginae by their (1) stronglyelongated labrum, (2) naked sternal plates (fully tuberculatein 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, butnot exclusive, for this family are: (1) an oval outline withonly feeble frontal sinus if present at all; (2) heterogeneousaboral tuberculation (enlarged primary tubercles developedin most forms; where developed, they are arranged withoutdistinct pattern).

The genus Platybrissus is another potential member ofthe 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 atpresent.

An independent cladistic analysis presented by(Stockley et al. 2005: fig. 2) likewise shows a monophyleticclade containing Eurypatagus, Paramaretia, and Platybris-sus. As in most trees resulting from the present analysis,Stockley et al. (2005) found a sister-group relationshipbetween 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 commentsof Andrew B. Smith, Rich Mooi and another anonymous re-viewer greatly improved the paper. Furthermore, I would liketo 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 andAndreas Thinschmidt (both Krahuletz Museum, Eggenburg)for helping to locate the type-material of H. peronatus and toManuel Kunz, Martin Sanders and Georg Oleschinski (all atthe Goldfuss Museum, Bonn) for locating the type materialof H. hoffmanni and providing photographs of it. Thanks alsoto Alice Schumacher (NHM Vienna) for taking the photo-graphs.

References

Agassiz, A. 1864. Synopsis of the echinoids collected by Dr. W. Stimpsonon the North Pacific Exploring Expedition under the command of Cap-tains Ringgold and Rodgers. Proceedings of the Academy of NaturalSciences of Philadelphia 15 (1863): 352–361.

—— 1873. Revision of the Echini. Memoirs of the Museum of Compar-ative Zoology at Harvard College III: pt. 3: 379–628 +1, pls 50–77.

—— & Clark, H. L. 1907. Preliminary report on the Echini collected1906, from May to December, among the Aleutian Islands, in Ber-ing Sea, and along the coasts of Kamchatka, Saghalin, Korea, andJapan, by U.S. Fish Commission steamer “Albatross”, in charge ofLieut.-Commander L. M. Garrett, U.S.N., commanding. Bulletin of theMuseum of Comparative Zoology at Harvard College 51(3): 107–139.


Agassiz, L. 1839. Description des Echinodermes fossiles de la Suisse;premiere partie, Spatangoides et Clypeasteroides. Memoires de laSociete helvetique des Sciences naturelles 3: i–viii, 1–101, 14 pls.

—— & Desor, P. J. E. 1847. Catalogue raisonne des especes, des genres,et des familles d’echinides. Annales des Sciences Naturelles, TroisiemeSerie, Zoologie 8: 5–35, 355–380.

Airaghi, C. 1903. Alcuni Echinidi del Terziario Veneto. Atti della SocietaItaliana di Scienze naturali e del Museo Civico di Storia naturale inMilano 41: 415–425, 1 pl.

—— 1905. Echinodermi miocenici dei dintorni di S. Maria Tiberina (Um-bria). Atti della Reale Accademia della Scienze di Torino 40: 43–54,1 pl.

Ali, M. S. M. 1985. On some Pliocene echinoids from the area north ofMersa Alam, Red Sea coast, Egypt. Palaontologische Zeitschrift 59:277–300.

Baker, A. N. & Rowe, F. W. E. 1990. Atelostomatid sea urchins fromAustralian and New Zealand waters (Echinoidea: Cassiduloida, Hol-asteroida, Spatangoida, Neoplampadoida). Invertebrate Taxonomy 4:281–316.

Beyrich, E. 1848. Zur Kenntniss des tertiaren Bodens der Mark Branden-burg. Karsten’s und v. Dechen’s Archiv fur Mineralogie etc. 22: 1–102.

Blanckenhorn, M. 1901. Neues zur Geologie und Palaeontologie Ae-gyptens. III: Das Miozan. Zeitschrift der Deutschen GeologischenGesellschaft 53: 52–132.

Bohm, A. von 1882. Uber einige tertiare Fossilien von der Insel Madura,nordlich von Java. Denkschriften der kaiserlichen Akademie der Wis-senschaften, mathematisch-naturwissenschaftliche Classe 45: 359–372, 4 pls.

Bolau, C. C. H. 1873. Die Spatangiden des Hamburger Museums Natur-wissenschaftlicher Verein Hamburg, Abhandlungen aus dem Gebieteder Naturwissenschaften 5(4): 1–23, 1 pl.

—— 1874. Neue Spatangiden des Hamburger Museums. Archiv fur Nat-urgeschichte 40(1): 175–178, pl. 6.

Brighton, A. G. 1931. The Geology of the Farsan Islands, Gizan andKamaran Islands, Red Sea. Part 3. Echinoidea. Geological Magazine68: 323–333.

Brodermann, J. 1949. Signification Estratigrafica de los EquinodermosFosiles Cubanos. Pp. 305–330 in Sanchez Roig, M. (ed.) Los equino-dermos fosile de Cuba. Paleontologia Cubana 1: 305–330.

Brusina, S. 1870. Ipsa Chiereghinii conchylia, ovvero contribuzionepella malacologia adriatica desunta dal manoscritto dall’abate S.Chiereghini. Bibliotheca Malacologia 2: 1–280.

Chesher, R. H. 1969. Contributions to the biology of Meoma ventricosa(Echinoidea: Spatangoida). Bulletin of Marine Science 19: 72–110.

Clark, H. L. 1916. Report on the sea lilies, starfishes, brittle stars, and sea-urchins obtained by the F.I.S. “Endeavour” on the coast of Queensland,New South Wales, Tasmania, Victoria, South Australia, and WesternAustralia. Biological Results “Endeavour”, 1909–1914 4: 1–23, 44pls.

—— 1917. Hawaiian and other Pacific Echini. The Echinoneidae, Nuc-leolitidae, Urechinida, Echinocorythidae, Calymnidae, Pourtalesiidae,Palaeostomatidae, Aeropside, Palaeopneustidae, Hemiasteridae, andSpatangidae. Memoirs of the Museum of Comparative Zoology at Har-vard College 46: 81–283, 18 pls.

—— 1924. Echinoderms from the South African Fisheries and MarineBiological Survey, Part I: Sea-Urchins (Echinoidea). Fisheries andMarine Biological Survey Report 4: 1–16, 4 pls.

—— 1925. A Catalogue of the Recent Sea-Urchins (Echinoidea) in theCollection of the British Museum (Natural History). Oxford UniversityPress, London, 250 pp.

Clark, W. B. & Twitchell, M. W. 1915. The Mesozoic and CenozoicEchinodermata of the United States. Monographs of the United StatesGeological Survey 54: 1–341, 108 pl.

Claus, C. F. W. 1876. Grundzuge der Zoologie. 3rd edn. N. G. Elwert’scheUniversitatsbuchhandlung, Marburg & Leipzig, XII+1254 pp.

Cooke, C. W. 1959. Cenozoic Echinoids of Eastern United States.United States Geological Survey Professional Papers 321: III+1–106,43 pls.

Cotteau, G. 1863. Echinides fossiles de Pyrenees. F. Savy, Paris, 160 pp.—— 1875. Description des Echinides Tertiaires des ıles St-Barthelemy

et Anguilla. Kongelige Svenska Vetenskaps-Akademiens Handlingar13(6): 1–48, 8 pls.

—— 1878. Echinides nouveaux ou peu connus. 1re serie. (Extraits de laRevue et Magasin de Zoologie). J.-B. Bailliere, Paris, 1878: 202–217,pls 29–30 (No. 111–120).

—— 1880. Description des echinides tertiaires de la Belgique. Memoirescouronnes et Memoires des Savants etrangers de l’Academie royaledes Sciences, des Lettres et des Beaux-Arts de la Belgique 43: 1–90.

—— 1885. Terrain Tertiaire, Tome I. Echinides Eocenes. Familles desSpatangidees, des Brissidees, des Echinoneidees et des Cassidulidees(pars). Paleontologie Francaise ou description des fossiles de la Francecontinuee par une reunion de paleontologistes sous la direction d’uncomite special, 1re Serie. – Animaux Invertebres G. Masson, Paris,1–48, pls 1–12 (Nov. 1885).

—— 1887. Echinides nouveaux ou peu connus. 2e serie, article 6. (Extraitsdu Bulletin de la Societe Zoologique de France). Paris, 91–103, pls.11–12 (No. 56–62).

—— 1889. Echinides rescueillis dans la province d’Aragon (Espagne)par M. Maurice Gourdon. Annales des Sciences Naturelles 7e Serie,Zoologie et Paleontologie 8: 1–60, pls. i–iv.

—— 1890. Echinides eocenes de la province d’Alicante. Memoires de laSociete Geologique de France, Serie 3 5 (Memire No. 2): 1–107, pls22–37.

—— 1893. Terrain Tertiaire, Tome II. Echinides Eocenes. Familles desCassidulidees (pars), Conoclypeidees, Clypeasteroidees, Scutellidees,Scutellinidees, Fibularidees et les Echinides reguliers. PaleontologieFrancaise ou description des fossiles de la France continuee par unereunion de paleontologistes sous la direction d’un comite special, 1re

Serie. – Animaux Invertebres G. Masson, Paris, 625–672, pls 353–364(Dec. 1893).

——, Peron, A. & Gauthier, V. 1885. Echinides fossiles de l’Algerie, 9e

fasc.; Etages eocene. G. Masson, Paris, 89 pp.——, —— & —— 1891. Echinides fossiles de l’Algerie, 10e fasc.; Etages

miocene et pliocene. G. Masson, Paris, 273 pp.d’Archiac, E. J. A. D. d. St. S. 1850. Description des fossiles du groupe

nummulitique recueillis par M. S.-P. Pratt et M. J. Delbos aux environsde Bayonne et de Dax. Memoires de la Societe Geologique de France,Deuxieme Serie 3(1 (Mem. n◦ 6)): 397–456, 6 pls.

Dames, W. 1878. Die Echiniden der vicentinischen und veronesischenTertiarablagerungen. Palaeontographica 25: 1–100, 11 pls.

Defrance, M. J. L. 1827. Spatangus Ocellatus. Pp. 96–97 in Blainvilleet al. (eds.) Dictionnaire des Sciences Naturelles, Tome 50. F. G.Levrault, Strasbourg & Paris.

de Loriol, P. 1863. Description de deux echinides nouveaux de l’etagenummulitiques de l’Egypte. Memoires de la Societe de Physique etd’Histoire Naturelle de Geneve 17: 103–107, 1 pl.

—— 1880. Monographie des echinides contenus dans les couches nummu-litiques de l’Egypte. Memoires de la Societe de Physique et d’HistoireNaturelle de Geneve 27: 1–92, 11 pls.

Dennant, J. 1890. Observations on the Tertiary and post-Tertiary geo-logy of south-western Victoria. Pp. 441–452 in W. Baldwin Spencer(ed.) Report of the second meeting of the Australian Association forthe Advancement of Science (Melbourne, January 1890.). AustralianAssociation for the Advancement of Science, Melbourne.

des Moulins, C. 1837. Troisieme Memoire sur les Echinides – Synonymiegenerale. Actes de la Societe Linneenne de Bordeaux 9(6): 45–364.

Desor, E. 1855–1858. Synopsis des echinides fossiles. Reinwald, Paris,lxviii+490 pp.

Doderlein, L. 1906. Die Echinoiden der Deutschen Tiefsee-Expedition.Pp. 61–290 in C. Chun (ed.) Wissenschaftliche Ergebnisse derDeutschen Tiefsee-Expedition auf dem Dampfer “Valdivia” 1898–1899. Gustav Fischer, Jena.

Dubois de Montpereux, F. 1843. Voyage au Caucase, chez les Tch-erkesses et les Abkhases, en Colchide, en Georgie, en Armenie et enCrimee. Atlas Serie de Geologie ou Ve. Serie. Dubois & Gide Libraire,Neuchatel & Paris, 1–4 pp.

176 A. Kroh

Duncan, P. M. 1864. A description of some fossil corals and echinodermsfrom the South-Australian Tertiaries. The Annals and Magazine ofNatural History, 3rd Series 14(81): 161–168, pls 5–6.

—— 1877. On the Echinodermata of the Australian Cainozoic (Tertiary)deposits. Quarterly Journal of the Geological Society of London 33:42–73, pls 3–4.

—— 1889. A revision of the genera and great groups of the Echinoidea.Journal of the Linnean Society, Zoology 23: 1–311.

—— & Sladen, W. P. 1883. The fossil Echinoidea of Kachh and Kattywar.Palaeontologia Indica, Ser. XIV 1(4): 1–91, 13 pls.

—— & ——, 1884a. Fossil Echinoidea of western Sind and the Coast ofBaluchıstn and of the Persian Gulf, from the Tertiary Formations. Fasc.III. The Fossil Echinoidea from the Khirthar Series of NummuliticStrata in Western Sind. Palaeontologia Indica, Ser. XIV 1(3): 101–246, pls 21–38.

—— & ——, 1884b. Fossil Echinoidea of western Sind and the Coastof Baluchıstn and of the Persian Gulf, from the Tertiary Formations.Fasc. IV. The Fossil Echinoidea from the Nari Series, the OligoceneFormation of Western Sind. Palaeontologia Indica, Ser. XIV 1(3): 247–272, pls 39–43.

Ebert, T. 1889. Die Echiniden des nord- und mitteldeutschen Oligocans.Abhandlungen zur geologischen Specialkarte von Preussen und denThuringischen Staaten 9(1): v–vi, 1–111, 10 pls.

Etheridge, R. J. 1875. Description of a new species of the genus Hemi-patagus Desor, from the tertiary rocks of Victoria, Australia, withnotes on some previously described species from South Australia.Quarterly Journal of the Geological Society of London 31: 444–450,1 pl.

—— 1878. A catalogue of Australian fossils. Cambridge University Press,Cambridge, 232 pp.

Felsenstein, J. 2002. PHYLIP Version 3.6 (alpha3), July, 2002. Computerprogram distributed by the author.

Fischer, A. G. 1966. Spatangoids. Pp. U543–U628 in R. C. Moore (ed.)Treatise on Invertebrate Paleontology, U. Echinodermata. 3 (2). GSA& University of Kansas Press, Boulder, CO & Lawrence, KS.

Fourtau, R. 1920. Catalogue des Invertebres fossiles de l’Egypte. Ter-rains Tertiaires. 2de partie: Echinodermes Neogenes. PalaeontologicalSeries 4, Geological Survey of Egypt, Cairo, iv+100 pp.

Fraas, O. 1867. Aus dem Orient: Geologische Beobachtungen am Nil,auf der Sinai-Halbinsel, und in Syrien. Ebner & Seubert, Stuttgart,222 pp.

Gagel, C. 1905. Ueber einige neue Spatangiden aus dem Nord-deutschen Miocan. Jahrbuch der Koniglichen Preussischen Geologis-chen Landesanstalt und Bergakademie zu Berlin 23 (1902): 525–543,pls 24–25.

Galeotti, H. G. 1837. Memoire sur la constitution geognostique de laprovince de Brabant, en reponse a la question suivante: Decrire laconstitution geologique de la province Brabant, determiner avec soinles especes minerales et les fossiles que les divers terrains renfernantet indiquer la synonymy des auteurs qui en ont deja traite. Memoiresde la Academie Royale des Sciences, des Lettres et des Beaux-Arts deBelgique 12: 1–192, 4 pls.

Gerth, H. 1922. Echinoidea. Pp. 497–520 in Martin, K. (ed.) Die Fossilienvon Java. Samml. geol. R.-Mus. Leiden, N. F. 2(4): 497–520.

Goldfuss, A. 1829. Petrefacta Germaniæ tam ea, quae in Museo Uni-versitatis Regiae Borussicae Fridericiae Wilhelmiae Rhenanae servan-tur quam alia quaecunque in Museis Hoeninghusiano Muensterianoaliisque extant, Iconibus et Descriptionibus illustrata. Arnz & Comp.,Dusseldorf, 1(2): 77–164; pls 26–50.

Grateloup, J. P. S. d. 1836. Memoire de geo-zoologie sur les oursinsfossiles, (Echinides), qui se recontrent dans terrains calcaires des en-virons de Dax (Departement des des Landes). Actes de la SocieteLinneenne de Bordeaux 8(4): 103–191, 2 pls.

Gray, J. E. 1825. An attempt to divide the Echinida, or sea eggs, intonatural families. Annals of Philosophy, new series 10: 423–431.

—— 1845. Description of two new invertebrated animals from Australia.Pp. 435–436 in E. J. Eyre (ed.) Journals of Expeditions of Discoveryinto Central Australia and Overland from Adelaide to King Georg’sSound in 1840–41. Volume 1. T. & W. Boone, London.

—— 1855. Catalogue of the Recent Echinida, or Sea Eggs, in the Collec-tion of the British Museum. Part I. Echinida Irregularia. Woodfall &Kinder, London, 69 pp.

—— 1866. Notes on a new species of Spatangus (S. variegatus). Proceed-ings of the Zoological Society of London 1866: 170–171.

Grube, A. E. 1866. Platybrissus Roemeri. Jahresbericht und Abhandlun-gen der Schlesischen Gesellschaft fur vaterlandische Cultur 43 (1865):61–62.

Henderson, R. A. 1975. Cenozoic spatangoid echinoids from New Zea-land. Paleontological Bulletin, New Zealand Geological Survey 46:1–90, 18 pls.

Herklots, J. A. 1854. Fossiles de Java. Description des restes fossilesd’animaux des terrains Tertiaires de Iıle de Java, recueillis. . . par F.Junghuhn. Pt. IV. Echinodermes. E. J. Brill, Leiden, 24 pp.

ICZN (ed.) 1999. International Code of Zoological Nomenclature, 4thedn. International Trust for Zoological Nomenclature, London, xxix +306 pp.

Irwin, R. P. & Archbold, N. W. 1994. The spatangoid echinoid Loveniafrom the Tertiary of southeastern Australia. Proceedings of the RoyalSociety of Victoria 106: 1–15.

Jeannet, A. & Martin, R. 1937. Ueber Neozoische Echinoideaaus dem Niederlaendisch–Indischen Archipel. Leidsche GeologischeMededeelingen 8: 215–308.

Jeffery, C. H. 1998. Carrying on regardless: the echinoid genus Cyclasterat the Cretaceous–Tertiary boundary. Lethaia 31: 149–157.

Kerr, A. M., Norris, D. R., Schupp, P. J., Meyer, K. D., Pitlik, T. J.,Hopper, D. R., Chamberlain, J. A. & Meyer, L. S. 1992. Rangeextensions of echinoderms (Asteroidea, Echinoidea, Holothuroidea)to Guam, Mariana Islands. Micronesica 25: 201–216.

Kier, P. M. 1977. The poor fossil record of the regular echinoid. Paleobi-ology 3: 168–174.

—— 1980. The echinoids of the Middle Eocene Warley Hill Formation,Santee Limestone, and Castle Hayne Limestone of North and SouthCarolina. Smithsonian Contributions to Paleobiology 39: 1–102, 22pls.

—— 1984. Fossil spatangoid echinoids of Cuba. Smithsonian Contribu-tions to Paleobiology 55: 1–336, 90 pls.

—— 1997. Oligocene echinoids of North Carolina. Smithsonian Contri-butions to Paleobiology 83: 1–37, 11 pls.

—— & Grant, R. E. 1965. Echinoid distribution and habits, Key LargoCoral Reef Preserve, Florida. Smithsonian Miscellaneous Collection149/6: 1–68, 16 pls.

—— & Lawson, M. H. 1978. Index of living and fossil echin-oids 1924–1970. Smithsonian Contributions to Paleobiology 34: 1–182.

Koch, A. 1884–1987. Die Alttertiaren Echiniden Siebenburgens. Mit-teilungen aus dem Jahrbuch der koniglichen ungarischen geologischenAnstalt 7: 45–132, 4 pls.

Koehler, R. 1914. Echinides du Musee Indien a Calcutta. I. Spatangides.Echinoderma of the Indian Museum. Part 8, Echinoidea (1). ZoologicalSurvey of India. Calcutta, 258 pp.

Kroh, A. 2005. Catalogus Fossilium Austriae. Band 2. Echinoidea neo-genica. Osterreichische Akademie der Wissenschaften, Wien, lvi+210 pp, 82 pls.

—— & Harzhauser, M. 1999. An echinoderm fauna from the LowerMiocene of Austria: paleoecology and implications for Central Parat-ethys paleobiogeography. Annalen des Naturhistorischen Museums inWien 101A: 145–191, 9 pls.

Kruse, P. D. & Philip, G. M. 1985. Tertiary species of echinoid genusEupatangus from southern Australia. Special Publications, South Aus-tralian Department of Mines and Energy 5: 167–185, 5 pls.

Lamarck, J. B. P. M. d. 1816. Histoire Naturelle des Animaux sansVertebres, presentant les caracteres generaux et particuliers de cesanimaux, leur distribution, leur classes, leurs familles, leurs genres,et le citation des principales especes qui s’y rapportent; precedeed’une Introduction offrant la Determination des caracteres essentiellesde l’animal, sa distinction du vegetal et des autres corps naturels,enfin, l’Exposition des Principes fondamentaux de la Zoologie. TomeTroisieme. Verdiere, Paris, 586 pp.


Lambert, J. 1902. Description des echinides de la Province de Barcelone.Fasc. 1. Memoires de la Societe Geologique de France, Paleontologie9: 1–57, 4 pls.

—— 1905. Notes sur quelques Echinides eoceniques de l’Aude et del’Herault. Pp. 129–164, pl. 6 in Catalogue Descriptif des FossilesNummulitiques de l’Aude et de l’Herault. Annales de l’Universite deLyon, Nouvelle Serie, I. Sciences, Medecine 17: 1–184.

—— 1909. Description des echinides fossiles des terrains mioceniquesde la Sardaigne. Memoires de la Societe Paleontologique Suisse 35(1908): 73–142, 6 pls.

—— 1910. Description des echinides des terrains neogenes du bassinRhone. Fasc. 1 Memoires de la Societe Paleontologique Suisse 37:1–48, 3 pls.

—— 1915. Description des echinides des terrains neogenes du bassinRhone. Fasc. 4 Memoires de la Societe Paleontologique Suisse 41:155–240, 5 pls.

—— 1927a. Revision des echinides fossiles de la Catalogne. Memoriasdel Museo de Ciencias Naturales de Barcelona, Serie Geologica 1:1–102, pls 1–4.

—— 1927b. Revision des echinides fossiles du Bordelais. III. Echinidesdu Miocene. Actes de la Societe Linneenne de Bordeaux 79: 71–132.

—— & Savin, L. 1902. Notes sur deux echinides nouveaux de la MolasseBurdigalienne dite de Vence (Alpes-Maritimes). Bulletin de la SocieteGeologique de France, Serie 4 2: 881–884, 1 pl.

Lambert, J. & Thiery, P. 1924. Essai de Nomenclature Raisonnee desEchinides. L. Ferriere, Chaumont, fasc. 6–7: 385–512, pls 10–11,14 (Dec. 1924).

Laube, G. C. 1869. Uber einige fossile Echiniden von den Murray cliffs inSud-Australien. Sitzungsberichte der Kaiserlichen Akademie der Wis-senschaften, mathematisch-naturwissenschaftliche Classe, AbtheilungI 59(1): 183–198, 1 pl.

Lawrence, J. M. 1987. A Functional Biology of Echinoderms. CroomHelm, London & Sydney, 340 pp.

Leske, N. G. 1778. Jacobi Theodori Klein naturalis dispositioechinodermatum . . . , edita et descriptionibus novisque inventis et syn-onomis auctorem aucta. Addimenta ad I. T. Klein naturalem dis-positionem Echinodermatum. G. E. Beer, Leipzig, XXII+278 pp.

Liao, Y. & Clark, A. M. 1995. The Echinoderms of Southern China.Science Press, Bejing, New York, 614 pp.

Lindley, I. D. 2003. Echinoids of the Kairuku Formation (Lower Plio-cene), Yule Island, Papua New Guinea: Spatangoida. Proceedings ofthe Linnean Society of New South Wales 124: 153–162.

Madon, C. 1994. Phylogenetic versus phenetic analysis in the spatangoidLovenia. Pp. 765–768 in B. David, A. Guille, J.-P. Feral & M. Roux(eds.) Echinoderms through Time – Proceeding of the 8th Interna-tional Echinoderm Conference Dijon/France, 6–10 Sept. 1993. A. A.Balkema, Rotterdam.

Manzoni, A. 1879. Gli echinodemi fossili dello Schlier delle colline di Bo-logna. Denkschriften der kaiserlichen Akademie der Wissenschaften,mathematisch-naturwissenschaftliche Classe 39: 149–164, 4 pls.

Markov, A. V. & Solovjev, A. N. 1995. The adaptions in the superor-der Spatangacea (Echinoidea) and their evolutionary role. Journal ofGeneral Biology 56: 539–553.

Martin, K. 1880. Revision of the fossil Echini from the Tertiary strata ofJava. Notes of the Leyden Museum 2: 73–85.

Mayer, K. 1860. Die Faunula des marinen Sandsteines von Kleinkuhrenbei Konigsberg. Vierteljahrsschrift der Naturforschende Gesellschaftin Zurich 6(2): 1–15.

McCoy, F. 1879. Tertiary Echinodermata. Pp. 33–42 in Prodromus of thePalaeontology of Victoria; or, Figures and Descriptions of the VictorianOrganic Remains. Geological Survey of Victoria, Melbourne.

Michelin, H. 1863. Echinides et Stellerides recueillis par M. Maillard,ingenieur colonial de 1856 a 1860. Pp. 1–7 in L. Maillard (ed.)Notes sur l’Ile de la Reunion (Bourbon). Deuxieme edition. Dentu,Paris.

Mortensen, T. 1948. Contributions to the Biology of the PhilippineArchipelago and adjacent regions. Report on the Echinoidea col-lected by the United States Fisheries Steamer ‘Albatross’ duringthe Philippine Expedition, 1907–1910. Part 3: The Echinoneidae,

Echinolampidae, Clypeastridae, Arachnidae, Laganidae, Fibulariidae,Urechinidae, Echinocorythidae, Palaeostomatidae, Micrasteridae, Pa-laepneustidae, Hemiasteridae, and Spatangidae. Bulletin of the Smith-sonian Institution, U.S. National Museum 100(14/3): 93–140.

—— 1950. A Monograph of the Echinoidea. V, 1. Spatangoida I.Protosternata, Meridosternata, Amphisternata I. Palæopneustidæ,Palæostomatidæ, Aeropsidæ, Toxasteridæ, Micrasteridæ, Hemiaster-idæ. C. A. Reitzel, Copenhagen, 432 pp.

—— 1951. A Monograph of the Echinoidea. V, 2. Spatangoida II. Am-phisternata II. Spatangidæ, Loveniidæ, Pericosmidæ, Schizasteridæ,Brissidæ. C. A. Reitzel, Copenhagen, 593 pp.

Neraudeau, D., David, B. & Madon, C. 1998. Tuberculation in spatang-oid fascioles: Delineating plausible homologies. Lethaia 31: 323–334.

Noetling, F. 1883. Ueber das Alter der samlandischen Tertiarformation.Zeitschrift der deutschen geologischen Gesellschaft 35: 671–694.

Philippe, M. 1998. Les echinides miocenes du Bassin du Rhone: revisionsystematique. Nouvelles Archives du Museum d’Histoire Naturelle deLyon 36(1–2): 3–241, 249–441, 26 pls.

Pomel, A. 1883. Classification methodique et genera des echinides vivanteet fossiles. Theses presentees a la Faculte des Sciences de Paris pourobtenir le Grade de Docteur es Sciences Naturelles 503. AldolpheJourdan, Alger, 131 pp.

—— 1885. Les echinides du Kef Iyhoud. Materiaux pour la Carte geolo-gique de l’Agerie, Serie 1, 1: 1–31, 3 pls, 3 photos.

—— 1887. Paleontologie ou description des animaux fossiles de l’Algerie.Zoophytes. 2e fascicule – Echinodermes. 2e livraison. AdolpheJourdan, Alger, 344 pp.

Richer de Forges, B. & Menou, J. L. 1988. The echinoderms of the softbottoms of the south-western lagoon of New Caledonia. Proceedingsof the 6th International Coral Reef Symposium Australia 2: 21–35.

Rowe, F. W. E. & Gates, J. 1995. Echinodermata. Pp. xiii+510 in A. Wells(ed.) Zoological Catalogue of Australia. Volume 33. CSIRO Australia,Melbourne.

Sanchez Roig, M. 1924. Revision de los Eqiniodos fosilos Cubanos.Memorias de la Sociedad Cubana Historia Natural “Felipe Poey”6(1–2): 6–50, 7 pls.

—— 1926. Contribucion a la paleontologia Cubana – los equinodermosfosiles de Cuba. Boletin de Minas 10: 1–179, 43 pls.

—— 1949. Los equinodermos fosiles de Cuba. Paleontologia Cubana 1:1–302, 50 pls.

Schaffer, F. X. 1912. Die Echiniden der Miocanbildung von Eggen-burg. Abhandlungen der Geologischen Reichsanstalt 22(2): 187–192,pls 58–60.

Seba, A. 1734–1765. Locupletissimi rerum naturalium thesauri accuratadescripto, et iconibus artificiosissimis expressio, collegit, digessit,descripsit et ad pingendum curavit per universam physices historiam,. . .Tome 3, 116 pls., Amstelaedami.

Smith, A. B. (ed.) 2004. The Echinoid Directory. World Wide Webelectronic publication. Available at: http://www.nhm.ac.uk/science-projects/echinoids (accessed 27 July 2004).

—— & Jeffery, C. H. 2000. Maastrichtian and Palaeocene echinoids: akey to world faunas. Special Papers in Palaeontology 63: 1–406.

—— & Stockley, B. 2005. Fasciole pathways in spatangoid echinoids:a new source of phylogenetically informative characters. ZoologicalJournal of the Linnean Society 144(1): 15–35.

Sorignet, L. A. 1850. Oursins fossiles de deux arrondissements dudepartement de l’Eure (Louviers et Andelys). Barbarot, Vernon,83 pp.

Srivastava, D. K. 2004. Fossil spatangoids (Echinodermata) from theIndian Subcontinent – a review. Journal of the Palaeontological Societyof India 49: 125–149, 7 pls.

Stockley, B., Smith, A. B., Littlewood, T., Lessios, H. A., MacKenzie-Dodds, J. A. 2005. Phylogenetic relationships of spatangoid seaurchins (Echinoidea): taxon sampling density and congruence betweenmorphological and molecular estimates. Zoologica Scripta 34: 447–468.

Stockley, G. M. 1927. Neogene Echinoidea from Zanzibar Protectorate.Pp. 103–117 in Report on the palaeontology of the Zanzibar Protect-orate. His Majestys Stationery Office, London.

178 A. Kroh

Sturt, C. 1833. Two expeditions into the interior of Southern Australiaduring the years 1828, 1829, 1830 and 1831, with observations on thesoil, climate and general rersources of the colony of New South Wales,Volume 2. Smith, Elder and Co., Cornhill, London, 271 pp.

Szorenyi, E. 1952. A Maretia hungarica Vadasz mjocen korufaj horvatartozasanak tiszatazsa. Foldtani Kozlony 82: 302–303.

Tate, R. 1894. Critical list of the Tertiary Mollusca and Echinodermataof New Zealand in the collection of the Colonial Museum. Reports ofGeological Exploration New Zealand 1892–1893: 121–127.

USSMDC 1999. Marine Biological Survey, Peacock Point Outfall,Wake Atoll June 1998NODC #0000247 Vers. 1. Department of theArmy, U.S. Army Space and Missile Defense Command, Hunts-ville, Alabama. Available at: http://www.coris.noaa.gov/metadata/records/html/nodc_0000247_wake_atoll_reefs_1998.html.

Vadasz, E. 1915. Die mediterranen Echinodermen Ungarns. GeologicaHungarica 1(2): 79–253, 6 pls.

Villier, L., Neraudeau, D., Clavel, B., Neumann, C. & David, B.2004. Phylogeny of Early Cretaceous spatangoids (Echinodermata:Echinoidea) and taxonomic implications. Palaeontology 47: 265–292.

Wanner, J. 1931. 13. Echinodermata. Pp. 436–460 in De Palaeontologieen stratigraphie van Nedelandsch Oost-Indie. Leidsche GeologischeMededeelingen 5 (K. Martin Festbundel): 436–460.

Weinberg, S. & de Ridder, C. 1998. Asthenosoma marisrubri n. sp.(Echinodermata, Echinoidea) from the Red Sea. Beaufortia 48(3): 27–46.

Woods, J. E. T. 1859. Remarks on a Tertiary deposit in South Australia.Transactions of the Philosophical Institute of Victoria 3: 85–94.

—— 1862. Geological observations in South Australia: principally in thedistrict south-east of Adelaide. Longman, Green, Longman, Roberts& Green / H.T. Dwight, London & Melbourne, xviii-404 pp.

—— 1865. On some Tertiary deposits in the Colony of Victoria, Australia.Quarterly Journal of the Geological Society of London 21: 389–394.

—— 1878. On the Tertiary deposits of Australia. Journal and Proceedingsof the Royal Society of New South Wales 11 (1877): 65–82.

Wright, T. W. 1864. On the fossil Echinidæ of Malta. With additionalnotes on the Miocene beds of the Island, and the stratigraphical distri-bution of the species therein; by A. Leith Adams. Quarterly Journalof the Geological Society of London 20: 471–491, 2 pls.

Zachos, L. G. 1993. Occurrence of the spatangoid echinoid Maretia ar-guta (Clark) in the Middle Eocene of Texas. Journal of Paleontology67: 148–150.

Zittel, K. A. 1867. II. Fossile Mollusken und Echinodermen aus Neu-Seeland. Pp. 15–68 in F. von Hochstetter, M. Hornes & F. R. vonHauer (eds) Palaontologie von Neu-Seeland. Beitrag zur Kenntniss derfossilen Flora und Fauna der Provinzen Auckland und Nelson. NovaraExpedition. Geologischer Theil. k.k. Hof- und Staatsdruckerei, Wien.

Appendix 1 Characters scored in the cladistic analysis

Shape1. Test outline: cordate (0); oval (1).2. Frontal sinus: absent (0); weakly developed (1); well developed (2).

Apical disc3. Apical disc, position: anteriorly displaced (0); subcentral (1); posteriorly displaced (2).4. Number of gonopores: four (0); three (1).

Ambulacra5. 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. IPZ�PZ (0); moderately wide IPZ∼PZ (1); narrow, i.e. IPZ�PZ (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).

Interambulacra13. 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 Sphaeridia23. 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).


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 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 3 3 4 4 4Character 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2

Brissus unicolor (Leske, 1778) 1 0 0 0 0 0 2 0 1 0 1 0 1 0 0 1 3 7 3 3 1 0 0 0 0 0 0 0 0 0 0 0 0 3 3 0 0 1 0 0 1 1Eupatagus valenciennesi Agassiz, 1847 1 0 0 0 0 0 0 0 1 1 0 0 3 2 1 0 3 2 2 2 1 0 2 0 1 1 0 1 1 0 1 0 0 0 0 0 0 2 1 0 2 1Spatangus inermis Mortensen, 1913 0 2 0 0 0 0 0 0 1 0 0 0 1 0 0 0 3 4 4 4 0 0 0 0 1 1 0 1 0 1 1 0 1 3 3 0 0 1 0 0 1 0Spatangus purpureus Muller, 1776 0 2 0 0 0 0 0 0 1 0 1 0 1 0 0 0 3 4 4 4 0 0 0 0 1 1 0 1 1 1 1 0 1 0 0 0 0 1 0 0 1 0Eurypatagus grandiporus Mortensen, 1948 1 0 0 0 0 0 1 0 1 0 1 0 4 2 2 1 4 5 4 4 1 0 1 2 1 1 0 1 1 1 1 0 0 0 0 1 0 2 0 0 0 0Eurypatagus ovalis Mortensen, 1948 1 0 0 0 0 0 1 0 1 0 1 0 4 2 1 1 4 5 5 4 1 0 1 2 1 1 0 1 1 1 1 0 0 0 0 1 0 2 0 0 0 0Eurypatagus parvituberculatus (Clark, 1924) 1 0 0 0 0 0 1 0 1 0 1 0 4 2 2 1 4 4 4 3 1 0 1 2 1 1 0 1 1 1 1 0 0 0 0 1 0 2 0 0 0 0Platybrissus roemeri Grube, 1865 1 0 0 0 0 0 0 0 1 0 1 0 4 2 1 1 4 3 4 3 1 0 2 1 1 1 0 1 0 0 1 0 0 0 0 0 0 2 0 0 0 0Maretia carinata Bolau, 1873 1 0 0 0 0 0 0 0 1 0 0 0 3 2 1 0 ? ? ? ? 1 0 1 1 ? ? 0 1 1 1 1 1 0 0 0 0 ? 2 1 0 2 0Maretia? cordata Mortensen, 1948 0 2 0 0 0 0 0 0 1 0 0 0 3 2 1 0 3 4 3 ? 1 0 2 1 ? 1 0 1 1 1 1 1 0 0 0 0 ? 2 1 0 2 0Maretia planulata (Lamarck, 1816) 1 1 0 0 0 0 0 0 1 0 1 0 3 2 1 0 4 4 3 3 1 0 1 1 1 1 0 1 1 0 1 1 2 0 0 1 1 2 1 0 2 0Hemimaretia elevata (Doderlein, 1906) 1 1 0 1 1 0 1 1 1 ? 0 0 2 2 1 1 2 ? ? ? 1 0 1 1 1 1 0 1 1 0 ? 2 ? ? ? 2 1 1 0 0 1 0Goniomaretia tylota Clark, 1917 1 0 1 1 0 0 0 0 2 1 0 0 2 2 1 1 2 1 2 2 1 0 1 0 1 1 1 1 1 0 3 2 2 2 3 1 1 2 1 0 3 0Pseudomaretia alta Agassiz, 1863 1 0 0 1 0 0 0 0 2 1 0 0 2 2 1 1 2 2 2 2 1 0 1 0 1 1 1 1 1 0 3 2 2 2 2 0 1 1 1 0 3 0Paramaretia multituberculata Mortensen, 1950 1 1 0 0 0 0 1 0 1 0 1 0 4 2 1 1 3 4 4 5 0 0 2 2 1 1 0 1 1 1 3 2 0 0 0 1 1 2 0 0 0 0Breynia australasiae (Leach, 1815) 1 1 0 0 0 1 0 0 2 0 0 1 2 2 1 1 3 3 3 2 1 0 1 0 0 0 1 1 1 0 3 2 2 0 1 0 0 2 1 1 3 1Breynia desorii Gray, 1851 1 1 0 0 0 0 0 0 2 0 0 1 3 2 1 1 3 3 3 2 1 0 1 0 0 0 1 1 1 0 3 2 2 0 0 0 0 2 1 1 3 1Lovenia cordiformis Agassiz, 1872 0 2 2 0 0 1 0 0 2 1 0 1 3 2 1 1 3 2 2 2 0 1 1 1 1 1 1 1 1 0 3 2 2 1 1 2 1 2 2 1 1 0Lovenia elongata Gray, 1845 0 2 0 0 0 1 0 0 2 1 0 1 2 2 1 1 2 1 2 2 0 1 1 1 1 1 1 1 1 0 3 2 2 1 1 2 1 2 2 1 1 0Lovenia gregalis Alcock, 1893 0 3 1 0 0 0 0 0 2 1 0 0 2 2 1 1 2 2 2 2 0 0 1 1 1 1 1 1 1 0 3 2 2 1 2 2 1 2 1 1 1 0Lovenia subcarinata Gray, 1845 0 2 0 0 0 1 0 0 2 1 0 0 3 2 1 1 3 2 2 2 0 0 1 1 1 1 1 1 1 0 3 2 2 2 2 2 1 2 2 1 1 0Lovenia triforis Koehler, 1914 0 2 0 1 0 0 1 0 2 1 0 0 2 2 1 1 2 3 3 2 0 0 1 1 1 1 1 1 1 0 3 2 0 1 2 2 1 2 1 1 1 0Pseudolovenia hirsuta Ag. & Clark, 1907 0 2 0 1 1 0 1 0 1 1 0 0 3 2 1 1 3 3 3 2 ? 0 1 1 1 1 1 1 1 0 3 2 2 1 2 2 1 2 ? 1 1 0Echinocardium cordatum (Pennant, 1777) 0 2 1 0 0 1 1 0 2 1 0 0 2 0 0 1 2 2 2 2 1 0 1 0 1 1 1 0 0 0 0 0 0 3 3 0 0 1 0 1 4 0Echinocardium mortenseni Thiery, 1909 1 1 0 0 0 0 1 0 2 1 0 0 2 1 0 1 2 2 2 2 1 0 1 0 1 1 1 0 0 0 0 0 0 3 3 0 0 2 0 1 1 0Homolampas lovenioides Mortensen, 1948 1 1 0 1 2 0 2 1 1 ? 0 0 3 2 1 1 2 2 3 ? ? 0 1 1 1 1 1 1 1 0 3 2 2 2 2 0 0 2 0 0 1 0Homolampas fragilis (Agassiz, 1869) 1 1 0 1 2 0 2 1 1 ? 0 0 2 2 1 1 2 2 2 2 ? 0 1 1 1 1 1 1 1 0 3 2 2 2 2 0 0 2 0 0 1 0Hemipatagus? argutus Clark, 1915 0 2 0 0 0 0 2 0 1 1 0 ? ? 2 1 1 ? ? ? ? ? 0 1 1 1 1 0 1 1 0 1 1 0 1 1 0 0 2 ? 0 1 0Hemipatagus? carolinensis (Kier, 1997) 0 2 0 0 0 0 0 0 2 1 0 ? 2 2 1 1 ? ? ? ? ? 0 1 1 ? ? 1 1 1 1 ? 1 1 1 2 0 0 2 ? 0 1 0Hemipatagus hoffmanni (Goldfuss, 1826) 0 2 0 0 0 0 0 0 2 1 0 0 2 2 1 1 2 1 2 2 0 0 2 1 1 1 1 1 1 0 2 2 2 1 2 1 1 2 1 0 1 0Hemipatagus ocellata (Defrance, 1827) 0 2 0 0 0 0 0 0 2 1 0 0 2 2 1 1 2 2 2 3 0 0 2 1 1 1 1 1 1 0 3 2 2 1 2 2 1 2 1 0 1 0Hemipatagus? subrostratus Clark, 1915 0 2 0 ? 0 0 2 0 ? ? ? ? ? 2 1 ? ? ? ? ? ? 0 ? 1 1 1 ? 1 1 0 1 1 0 1 2 0 0 2 ? 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 eithergenus (based on Lambert & Thiery 1909–1925 and Kier &Lawson 1978). Systematic decisions were made on the basisof published descriptions and illustrations or taken from theliterature 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 & Thiery: 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 TypeNo. 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 anda 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, figs5, 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) andHenderson (1975: 33), Herklots’ two syntypes represent twodifferent genera/species. One specimen (no. L4292) belongsto the genus Eupatagus. The other specimen (no. L4291) isa 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 atHarvard College, Cambridge.

TYPE LOCALITY. Kagoshima Bay, Japan, ∼ 9 m water depth.

AGE. Recent.

TAXONOMIC PLACEMENT. Type species of PseudomaretiaKoehler, 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 & Philip1985: 173); based, among other features, on the presenceof a peripetalous fasciole, which had been previously recog-nised by Duncan (1877) and subsequent authors, but still thespecies remained in the genus Maretia until the revision byKruse & Philip (1985).

antillarum Cotteau, 1875∗1875 Euspatangus [sic] Antillarum, Cotteau, 1875;

Cotteau: 43; pl. 7, figs 7–11.

SYNTYPES. Collection Cleve, Museum Stockholm, Museumof Upsala.

TYPE LOCALITY. St. Barthelemy Island.

AGE. Eocene.

TAXONOMIC PLACEMENT. Eupatagus (Henderson 1975:33), senior synonym of Maretia clarki Lambert and Maretiatwitchelli 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.


TAXONOMIC PLACEMENT. Leiopneustes (Lambert & Thiery1924: 448); periproct high on the posterior face, visible inaboral 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 likeanything found in Lovenia, Hemipatagus or Maretia. On thewhole the illustrations resemble much more a brissid or somehemiasterids 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-Chateaux, 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, ClarkeCounty, Mississippi, USA.

TYPE OCCURRENCE. Winona Formation, Clairborne Group.

AGE. Middle Eocene.

OCCURRENCE. Middle Eocene of Mississippi and Texas,USA.

TAXONOMIC PLACEMENT. Hemipatagus? Although thisspecies (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, thepresence of rudimentary pores in the anterior poriferouszones of petals II and IV and the distribution of the aboralprimary 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.

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 bilobedsubanal fasciole, distribution of the aboral primary tubercles. . .) in the original publication support an attribution to Hemi-patagus.

buendensis Ebert, 1889∗1889 Spatangus (Maretia) bundensis Ebert n. sp.; Ebert:

69–70; pl. 8, fig. 1a–c.

SYNTYPES. Two specimens in the Museum of Gottingen.

TYPE LOCALITY. Doberg, Germany.

AGE. Late Oligocene.

TAXONOMIC PLACEMENT. Hemipatagus; very similar to thejuveniles of H. martensii, as observed previously by Ebert(1889); Lambert & Thiery (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, figs1–3), collection ‘de la Sorbonne’, Paris.

TYPE LOCALITY. Chaumont (Oise), France.

LITHOSTRATIGRAPHY. Calcaire grossier inferieur.

AGE. Middle Eocene.

TAXONOMIC PLACEMENT. Hemipatagus; very similar to H.grignonensis, possibly a juvenile or subadult specimen ofthat species; the cordiform outline with distinct frontal sinus,the shape and structure of the petals and the distributionof the primary tubercles support an attribution to Hemi-patagus; the original description does not reveal details onthe nature and tuberculation of the plastron and the shape ofthe 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 inalcohol, Viti Islands); Museum Godeffroy (2 dry specimens,Viti Islands; 1 dry specimen, Ellice Islands), Musuem ofGottingen, 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 supportan attribution to the genus Maretia; the shape of the subanalfasciole is unknown and the adapical pores in the anteriorporiferous zones of the anterior paired petals are reduced insize.

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 specimensin the collections of the Geological Survey of Preussen, theMuseum of Lubeck and the Museum of Hamburg.

TYPE LOCALITY. Zarrentin, south-east Holstein, NorthernGermany.

AGE. Erratic boulders of Cenozoic age, presumably Mio-cene.

TAXONOMIC PLACEMENT. Type species of Chuniola Gagel,1905; probably a subjective junior synonym of HemipatagusDesor, 1858 according to Smith (2004: ‘The Echinoid Direct-ory’); differs from typical Hemipatagus by lacking enlargedaboral primary tubercles in interambulacra 2 and 3 and bythe presence of such tubercles in adapical interambulacrum5; due to the poor preservation of the specimens the pres-ence/absence of internal, subanal and peripetalous fasciolescould 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, shortlabrum 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, figs1–2.

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.


TAXONOMIC PLACEMENT. Hemipatagus; although this spe-cies has no truly camellate tubercles (even though the areolesare distinctly sunken), it clearly belongs to Hemipatagusbased on the characteristic cordiform outline, typical shapeof the petals with rudimentary pores in the anterior porifer-ous zones of petal II and IV and the conspicuous field oftubercles in adapical ambulacrum III.

cartagensis Sanchez Roig, 1949∗1949 Hemipatagus cartagensis Sanchez Roig: 216–217

[not seen, fide Kier & Lawson 1978: 121].

AGE. Late Oligocene (Brodermann 1949: 323) or Late Eo-cene (Sanchez Roig 1949).

TAXONOMIC PLACEMENT. Doubtful; not mentioned inKier’s (1984) revision of the fossil spatangoids of Cuba.

clarki Lambert, in Sanchez-Roig, 1924

1924 Maretia clarki Lambert in Sanchez Roig; Lambert& Thiery: 458. [nom. nov. pro specimen B of Eu-patagus floridanus Clark, 1915 (in Clark & Twitchell1915: 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 Eupatagusantillarum (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 pronouncedcordate 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 inDubois’ paper shows the aboral surface of a damaged speci-men with heterogeneous aboral tuberculation. The tuberclesare confined to the area inside the petalodium and seem tohave only slightly sunken areoles. This and the shape of thepetals is strongly reminiscent of Eupatagus and the speciesis tentatively referred to that genus. The illustrations in deLoriol (1880) are rather poor and do not offer additionalinsights. Unfortunately, the present author did not have theopportunity to check the text of Dubois’ work for a descrip-tion of the species.


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-Pyrenees, 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 thespecies to Maretia; based on the presence of a distinct frontalsinus, the distribution of the camellate primary tubercles, thestructure of the petals, and the presence of a bilobed subanalfasciole, the species is here attributed to Hemipatagus.

duncani Lambert, in Lambert & Thiery, 1924∗1924 H. Duncani Lambert; Lambert in Lambert &

Thiery: 457 [nom. nov. pro Eupatagus rostratusDuncan & Sladen (1884b: 267, pl. 43, fig. 6), nond’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 ofSehwan, India.

TYPE OCCURRENCE. Nari Series.

AGE. Middle Oligocene (Srivastava 2004).

OCCURRENCE. ?Eocene–Oligocene and, possibly, Mioceneof India, Japan and Java.

TAXONOMIC PLACEMENT. Eupatagus; junior synonym of E.rostratus rostratus d’Archiac, 1850 according to Henderson(1975: 44).

elevata Doderlein, 1906∗1906 Maretia elevata nov. spec.; Doderlein: 263–265;

pl. 33, figs 5, 5a–b; pl. 48, figs 6a–h.1951 Hemimaretia elevata (Doderlein); Mortensen: 53–

55; pl. 1, fig. 4; pl. 44, figs 5–7, 21.

HOLOTYPE. Badly broken specimen in the Museum ofBerlin (Mortensen 1951).

TYPE LOCALITY. Station 251 (Valdivia-Expedition), off Wit-uland, Eastern Africa (693 m depth).

AGE. Recent.

TAXONOMIC PLACEMENT. Type-species of HemimaretiaMortensen, 1950; this genus has only 3 gonopores (poreof genital plate 2 lacking) and is thus easily distinguishedfrom 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 ona single specimen; according to Mortensen (1950: 281, 1951:26) this species probably belongs to either Platybrissus or,less likely, to Eurypatagus.

estenozi Sanchez Roig, 1926

1984 ? Maretia estenozi Sanchez Roig; Kier: 106–107;pl. 55, fig. 5, pl. 56, figs 1–2.

HOLOTYPE. SRC 5491, figured by Sanchez Roig (1926:pl. 37, fig. 1); Sanchez Roig collection, Cuban Academyof Science.

TYPE LOCALITY. ‘Cervantes’ farm, San Jose de las Lajas,Haban Province, Cuba.

AGE. Early – Middle Miocene.

TAXONOMIC PLACEMENT. Eupatagus (Kier, 1984: 107),possible synonym of E. cubensis (Cotteau, 1875) accordingto Kier (1984: 103–107).

fasciata Lambert, in Lambert & Thiery, 1924∗1924 M. fasciata Lambert; Lambert in Lambert &

Thiery: 458.

HOLOTYPE. Unknown.

TYPE LOCALITY. Philippines.

AGE. Recent.

TAXONOMIC PLACEMENT. Maretia; juvenile of M. planulataaccording 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; neverdescribed or illustrated; on page 79 of the same paper theytentatively 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 HoffmanniSturt 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; McCoy1879; Dennant 1890) referring to this species actuallyrepresent 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 toEarly 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, EasternDesert, Egypt (coll. Blanckenhorn); 11 fragments from WadiEtthal, Sinai, Egypt (coll. Rothpeltz); current whereaboutsunknown.

AGE. Presumably Early or Middle Miocene.

TAXONOMIC PLACEMENT. Spatangus?; classification un-clear; Oppenheim reports that he could not find any tracesof 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 relatedto Spatangus.

gauthieri Lambert, in Lambert & Thiery, 1924

1885 Tuberaster tuberculatus, Peron & Gauthier, 1885;Cotteau et al.: 47–49; pl. 3, figs 1–4 [type-speciesof 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 &Thiery: 457 [nom. nov. pro Tuberaster tuberculatusPeron & 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 thefully tuberculate sternum (if correctly figured in Cotteauet 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 theirmaterial.

girundicus Lambert, 1915∗1915 H. girundicus; Lambert: 191.1924 H. girundicus Lambert; Lambert in Lambert &

Thiery: 457; pl. XIII, fig. 5.1927b Hemipatagus girundicus Lambert, 1915; Lambert:

119–121.

HOLOTYPE. Whereabouts unknown, a cast of the specimenis figured in Lambert & Thiery (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 Rhone Basin.

gottschei Gagel, 1905∗1905 Spatangus ? (Eupatagus ?) Gottschei 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 collectionsof the Geological Survey of Preussen, and the Museum ofHamburg.


AGE. Erratic boulders of Cenozoic age, presumablyMiocene.

TAXONOMIC PLACEMENT. Hemipatagus?; based on the out-line and the distribution of the aboral primary tubercles; typematerial very poor and in need of redescription.

grignonensis Cotteau, 1880

1837 Spatangus grignonensis Desmarest, ined. in litt.;des Moulins: 236–237 [nomen nudum].


∗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.


AGE. Middle Eocene.

TAXONOMIC PLACEMENT. Hemipatagus; Cotteau’s descrip-tion and illustrations partly contradict each other; assumingthe description to be correct this species has a partially tuber-culate sternum, a bilobed subanal fasciole, a cordate outlinewith distinct frontal notch but no rudimentary pores in adap-ical petals II and IV; generic attribution thus is doubtful butHemipatagus 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 desMoulins provides no description or illustration he cannot beconsidered 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. Gattieres, Alpes-Maritimes, France.

AGE. Burdigalian, Early Miocene.

TAXONOMIC PLACEMENT. Hemipatagus; clearly a memberof the genus Hemipatagus with all the typical features de-veloped (outline, tuberculation . . .); very similar to subadultH. 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.


AGE. Middle Eocene.

TAXONOMIC PLACEMENT. Referred to PhymapatagusLambert, 1910 (type-species: S. britannus Tournouer) byLambert & Thiery (1924: 463); this is supported by the fullytuberculate plastron and lack of camellate primary tubercles.

hispanica Cotteau, 1890∗1890 Maretia hispanica, Cotteau, 1889; Cotteau: 5–6;

pl. 22 (1), figs 1–3.

HOLOTYPE. Specimen figured by Cotteau (1890); Cotteaucollection.

TYPE LOCALITY. Callosa, Alicante, Spain.

AGE. ? Eocene.

TAXONOMIC PLACEMENT. Hemipatagus?; the shape of thetest and the distribution and nature of the primary tubercleswould indicate an attribution to Hemipatagus, yet, theplastron is shown as being fully tuberculate in pl. 22 (1), fig.3; it is known, however, that Cotteau’s artist often providedhighly idealized figures, adding plate sutures or tubercleswhere 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 additionalsyntypes in the collection of the Goldfuss Museum (Instituteof Palaeontology, University of Bonn, Germany), number357a–h.

TYPE LOCALITY. Doberg, near Bunde, Germany.

AGE. Chattian, Late Oligocene.

TAXONOMIC PLACEMENT. Type species of Hemipatagus.

hungarica Vadasz, 1915∗1915 Maretia hungarica n.sp.; Vadasz: 237; pl. 9 (3),

figs 17–18.1952 Maretia hungarica Vadasz; Szorenyi: 302, figs a–b.

HOLOTYPE. Specimen MAFI Ech-225, figured by Vadasz(1915: pl. 9 (3), figs 17–18) and Szorenyi (1952: figs a–b);housed at the Museum of the Hungarian Geological Survey,Budapest.

TYPE LOCALITY. Matraverebely, Nograd, Hungary.

AGE. Late Badenian (Early Serravallian), Middle Miocene.

REMARKS. Specimen damaged since the photograph ofSzorenyi (1952), petal V now missing.

TAXONOMIC PLACEMENT. Hemipatagus; Lambert & Thiery(1924: 458) tentatively referred this species to Atelospatan-gus Koch, 1885 because no anterior poriferous zones of petalsII and IV are visible in Vadsz’s drawing; the correctness ofthis action was questioned by Mortensen (1951: 79), arguingthat the drawing might be incorrect; Mortensen was indeedcorrect, as shown by Szorenyi (1952: 302), only the adapicalpores of the anterior poriferous zones of petals II and IV arerudimentary; no internal fasciole could be observed in thetype.

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.


AGE. Middle Eocene.

TAXONOMIC PLACEMENT. Doubtful; published descriptionsare 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 theUniversity of Gottingen; others in the collections of the Uni-versity of Munich, the Museum of Breslau, the Natural His-tory Museum of Osnabruck and the Geological Survey inBerlin.



TAXONOMIC PLACEMENT. Similar to H. hoffmanni, but withfully tuberculate sternal plates and non-camellate primarytubercles, 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. Drome, France.

AGE. Burdigalian, Early Miocene.

TAXONOMIC PLACEMENT. ? Lovenia; Cotteau’s fig. 7provides a magnification of the aboral primary tubercles,showing distinct crenulations; no extant species of Love-nia with crenulated primary tubercles are known (Mortensen1951); although Cotteau explicitly stated that an internal anda subanal fasciole are present, Lambert & Savin (1902: 884)and Lambert & Thiery (1924: 467) state that this speciescould be a Hemipatagus, an opinion not shared by any sub-sequent author; lately Philippe (1998: 225) suggested thatthe type specimen might be a juvenile Maretia ocellata (hereHemipatagus); he argued that he could find no trace of an in-ternal fasciole in the cast of the holotype studied by him andthat the general appearance was much more like H. ocellatusthan like a Lovenia.

madurae Bohm, 1882∗1882 Hemipatagus Madurae n. sp.; Bohm: 366–367;

pl. 2, fig. 3a–c.

HOLOTYPE. Specimen figured by Bohm (1882: pl. 2,figs 3a–c); whereabouts unknown.

TYPE LOCALITY. Sepoloc, northern coast of Madura, N ofJava.

AGE. ? Neogene.

TAXONOMIC PLACEMENT. ? Lovenia (referred to the genusVasconaster by Lambert & Thiery (1924: 466), today con-sidered a junior synonym of Lovenia); generic attributionunclear, type material needs to be re-examined to verify thepresence or absence of an internal fasciole and to study themorphological 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 ofthe genus Hemipatagus with the characteristic cordiform out-line with distinct frontal sinus, typical distribution of theaboral primary tubercles (restricted to the anterior part nearthe interradial suture in the interambulacra 1 and 4) andtypical shape of the petals, with rudimentary pores in theadapical 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 ofGottingen, Germany.



TAXONOMIC PLACEMENT. Hemipatagus; this species exhib-its all the features typical for Hemipatagus and is actuallyrather 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 Loveniaduncani (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. Reunion Island, Indian Ocean.

AGE. Recent.

TAXONOMIC PLACEMENT. Maretia; typical Maretia withlong, straight petals, characteristic distribution of the aboralprimary tubercles (also present in the posterior part of the


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. ObviouslyMichelin was not aware of the genus Maretia, as he states‘Le spatangus planulatus de Lamarck, qui se trouve dansla mer Rouge, et subfossile sur ses bords, est une especed’hemispatangus, vivant aujourd’hui’ (1853: A 6); accordingto Mortensen (1951: 38–39) this species is a junior synonymof M. planulata (Lamarck).

mauritanica Pomel, 1885∗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, Dept. d’Alger, Algeria.

AGE. Eocene or Oligocene.

TAXONOMIC PLACEMENT. Lovenia or Hemipatagus;without re-examining the types or topotypic material it isimpossible to decide to which of the two genera the speciesbelongs; none of the authors that have studied the specimenscould confirm the presence or absence of a internal fasciole(e.g. Cotteau et al. 1885: 37–38); Contrary to the figures inCotteau’s Paleontologie Francaise (1893; pl. 362, figs 1–6),the plastron is not fully tuberculate (compare description andillustration in Cotteau et al. 1885); Cotteau et al. (1885) stateexplicitly that the aboral primary tubercles are crenulated inthis species (which can also be seen on their pl. 1, fig. 5), afeature which does not occur in extant Lovenia according toMortensen (1951); however, it seems that many ambulacralplates 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 Geologieen 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 Bohm, 1882∗1882 Brissomorpha mojsvari n. sp.; Bohm: 364–365;

pl. 2, figs 1a–d.

HOLOTYPE. Specimen figured by Bohm (1882: pl. 2,figs 1a–d); whereabouts unknown.

TYPE LOCALITY. Sepocloc, northern coast of Madura, N ofJava.

AGE. ? Neogene.

TAXONOMIC PLACEMENT. ? Eupatagus; this species hasbeen 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 inBohm; as Bohm (1882: 365) explicitly mentions the presenceof a peripetalous fasciole this species is tentatively referred toEupatagus here (contrary to Jeannet & Martin (1937: 277),who assume that a peripetalous fasciole is lacking based onthe distribution of the aboral primary tubercles in Bohm’sfigure).

nicklesi Cotteau, 1890∗1890 Maretia Nicklesi, Cotteau, 1889; Cotteau: 6–7;

pl. 33 (12), figs 7–10.

HOLOTYPE. Specimen figured by Cotteau (1890); Nicklescollection.

TYPE LOCALITY. Callosa, Alicante, Spain.

AGE. ? Eocene.

TAXONOMIC PLACEMENT. Eupatagus; based on labrumshape, 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 synonymof H. ocellatus according to Agassiz & Desor (1847: 7) andPhilippe (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-Chateaux, Rhone Basin,France.

AGE. Burdigalian.

TAXONOMIC PLACEMENT. Hemipatagus; this well-knownspecies shows all features characteristic of Hemipatagus: acordate 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 ofMaretia 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 toMortensen (1951: 37) the identity of this species isunclear; Leske’s illustrations were based on those fromSeba’s Thesaurus III (pl. 15, figs 27–28), which accordingto Mortensen are rather poor and of unknown affinity; Hestrongly argues against synonymising this species withMaretia planulata as proclaimed by other authors (e.g. Clark1925).

pareti Manzoni, 1879∗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 MazettiaLambert & Thiery in Lambert, 1915 (replacement for Manzo-nia Pomel, 1883 preoccupied by Manzonia Brusina, 1870),regarded as subgenus of Prospatangus Lambert, 1902 byLambert (1915); current status unclear, but certainly notclosely related to either Hemipatagus or Maretia (based onthe fully tuberculate sternal plates, the en-chevron arrange-ment of the aboral primary tubercles, the presence of primarytubercles 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-Pyrenees, France.

TYPE OCCURRENCE. Couche a Eupatagus ornatus.

AGE. Late Eocene.

TAXONOMIC PLACEMENT. Hemipatagus (this paper); basedon 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 anddistribution of the primary tubercles; single known specimenis rather small and very similar to juvenile H. desmoulinsifrom 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.

TYPE LOCALITY. 26 miles south-west of cape Everard,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 deLoriol 1880).

TYPE AREA. Sinai Peninsula, Egypt.

AGE. ? Eocene.

TAXONOMIC PLACEMENT. ? Eupatagus; the specimenfigured by de Loriol (1880) clearly seems to be a Eupatagusbased on the shape of the petals, the size and distribution ofthe enlarged primary tubercles; the presence of a peripetal-ous fasciole is not confirmed but seems highly probable; theidentity 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 inSchaffer (1912: pl. 59, figs 4–6); housed at the Krahuletz-Museum, Eggenburg, Austria.


TYPE LOCALITY. Eggenburg (Kremserberg), lower Austria.

TYPE OCCURRENCE. Zogelsdorf Formation.

AGE. Late Eggenburgian (Early Burdigalian), Early Mio-cene.

TAXONOMIC PLACEMENT. Hemipatagus; junior synonym ofH. ocellatus (Desmarest) (see Kroh 2005).

planulatus Lamarck, 1816∗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 NaturelleParis.

TYPE AREA. ‘les mers australes’.

AGE. Recent.

OCCURRENCE. South-western Red Sea; Indian Ocean andwestern Pacific, from Eastern Africa to Fiji and GilbertIslands, and from Sagami Bay (Japan) to Port Jackson(Australia)

TAXONOMIC PLACEMENT. Type-species of Maretia; IndianOcean populations are usually whitish to yellow, Malayanspecimens are dark coloured, but show no morphologicaldifferences according to Mortensen (1951: 36).

planulata abbassi Ali, 1985∗1985 Maretia planulata abbassi n. sp.; Ali: 294–295;

figs 12A–B.

TYPE MATERIAL. USNM 339748.

TYPE LOCALITY. Wadi Abu Abraiki, Western Egypt.

AGE. Early Pliocene.

TAXONOMIC PLACEMENT. Maretia; very similar to extantM. planulata.

praelongus Herklots, 1854∗1854 Spatangus praelongus, nouv. esp.; Herklots: 11–12;

pl. 2, fig. 6.1858 [Hemipatagus] praelongus; Desor: 418.1880 Maretia planulata Gray; Martin: 81.1924 Atelospatangus praelongus; Lambert & Thiery:

458.1937 Maretia planulata (Lamarck); Jeannet & Martin:

277–278.

HOLOTYPE. The specimen figured by Herklots (pl. 2, fig. g),no. 433.


AGE. Pliocene.

TAXONOMIC PLACEMENT. Maretia; junior synonym of theextant Maretia planulata according to Jeannet & Martin

(1937: 277); Herklots (1854: 11) mentioned a second speci-men (no. 452b; pl. 4, figs 5, 5a) which he tentatively referredto 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–

b.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).


AGE. Pliocene.

TAXONOMIC PLACEMENT. Eupatagus; Lambert & Thiery(1924: 451, 458) listed this species both under the pre-Linnaean genus Brissoides and under Maretia; the oval,anteroposteriorly elongated outline, lack of a frontal sinus,shape of the petals, fully tuberculate plastron, restriction ofthe aboral primary tubercles to the area between the petalsand the presence of a peripetalous fasciole indicate an attri-bution to the genus Eupatagus.

regiomontanus Mayer, 1860∗1860 Hemispatangus [sic] Regiomontanus Mayer;

Mayer: 11 [pagination of the reprint].

TYPE MATERIAL. Current whereabouts unknown.

TYPE LOCALITY. Kleinkuhren, north-west of Konigsberg,Samland, Russia.

AGE. Late Eocene.

TAXONOMIC PLACEMENT. Hemipatagus?; poorly known;a junior synonym of H. sambiensis according to Cotteau(1885).

rotundus Duncan & Sladen, 1884∗1884a Macropneustes roundus, Duncan & Sladen;

Duncan & Sladen: 232–233; pl. 38, figs 6–7.1924 Hemipatagus rotundus Duncan & Sladen (Mac-

ropneustes); Lambert & Thiery: 457.2004 Macropneustes (Macropneustes) roundus (Duncan

& Sladen) n. comb.; Srivastava: 147–148; pl. 7, figs12–13.

HOLOTYPE. Geological Survey of India no. G302/100 (GSIType No. 2685 (Srivastava 2004)).

TYPE LOCALITY. Teyon (or Tiyun), east of Chorla, Sind,India.

LITHOSTRATIGRAPHY. Kithar Series.

AGE. Middle Eocene.

190 A. Kroh

TAXONOMIC PLACEMENT. Eupatagus or Macropneustes(Macropneustes); Duncan & Sladen (1884a) expressed theirproblems concerning the generic attribution of this speciesand named both genera mentioned above as potential candi-dates. They choose Macropneustes because of the numerousand prominent development of the aboral primary tuberclesand the inferred absence of the subanal fasciole (specimendamaged posteriorly). Neither Henderson (1975: 33) norSrivastava (2004: 147–148) state why they changed the gen-eric attribution and, apparently, no additional specimens havebeen found since the discovery of the types. Until more spe-cimens are described the generic attribution of this speciesremains 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 aboralprimary tubercles (missing in posterior part of interambulac-ral columns 1a and 4b), typical petals with rudimentary poresin 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 wouldhave 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 Konigsberg, Samland, Russia.

AGE. Late Eocene.

TAXONOMIC PLACEMENT. Hemipatagus ?; poorly known;H. regiomontanus Mayer, 1860 is a junior synonym of thisspecies 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. Vallee du Trapel, Aude, France.

AGE. Middle Eocene.

TAXONOMIC PLACEMENT. Doubtful; referred to Leiop-neustes by Lambert & Thiery (1924: 448); type materialpoorly preserved, large parts of the aboral and oral surfaceare missing; referred to Maretia by Cotteau (1893) on thebasis of the missing peripetalous fasciole; the features thatcan be observed, however, are in favour of Eupatagus (oval,elongated outline; very shallow frontal sinus; non-camellateprimary 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 Setif, Dept. 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 absenceof an internal fasciole, but mention in their discussion thatthis species is similar to H. ocellatus and explicitly state thatthe latter lacks an internal fasciole, even in perfectly pre-served specimens; the cordate shape, the pronounced frontalsinus, the rudimentary pores in the adapical poriferous zoneof ambulacra II and IV, as well as the distribution of theprimary 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 tuberclesand shape of the petals suggest the attribution to Hemi-patagus; according to Cooke (1959: 81) this species is verysimilar 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).


TAXONOMIC PLACEMENT. Hemipatagus; according toCotteau et al. (1891) no traces of an internal fasciole couldbe 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 primarytubercles suggest an attribution to Hemipatagus.

tuberculata Agassiz & Clark, 1907∗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 anattribution to Lovenia is also not improbable (the holotypehas 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 planulatathe 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; inLovenia scaber, by contrast, the gonopores open at 30 mm).

tuberculatus Zittel, 1867∗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.


OCCURRENCE. Whaingaroan to Altonian (Oligocene toEarly Miocene) of New Zealand.

TAXONOMIC PLACEMENT. Lovenia.

tuberculatus Peron & Gauthier, in Cotteau et al.,1885∗1885 Tuberaster tuberculatus, Peron & Gauthier, 1885;

Cotteau et al.: 47–49; pl. 3, figs 1–4 [= H. gauthieriLambert in Lambert & Thiery, 1924: 457 (nom. nov.pro Tuberaster tuberculatus Peron & Gauthier, inCotteau et al., 1885 non Zittel, 1867); see above].

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 thefully tuberculate sternum (if correctly figured in Cotteauet 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 theirmaterial.

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 likelyIV) and the adjoining interambulacra which bear camelatetubercles; currently impossible to decide whether it is a Love-nia, Hemipatagus or something else.

twitchelli Lambert, in Lambert & Thiery, 1924

1924 Maretia twitchelli Lambert in Sanchez Roig;Lambert & Thiery: 458 [nom. nov. pro specimenC 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 ofEupatagus 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-seum, London.

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∗1878 Breynia vicentina nov. sp.; Dames: 75–76; pl. 7,

figs 7a–b.

HOLOTYPE. Specimen figured by Dames (1878: pl. 7, figs7a–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, butinferred the presence of an internal fasciole on the basis ofthe 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 accordingto the figure, whereas he mentions details of the adapical areain the description; likewise he reports a naked plastron, whilethe figures show that the plastron is largely missing); it is un-clear why Lambert & Thiery (1924: 466) refer this speciesto Hemipatagus; the available information rather suggestsan attribution to Lovenia, providing Dames’ description iscorrect.

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,Victoria, Australia.

TYPE OCCURRENCE. Black Rock Sandstone, BrightonGroup.

AGE. Cheltenhamian (Messinian), Late Miocene.

OCCURRENCE. Early (?) Miocene – Pliocene (Zanclean) ofthe Port Phillip Basin, Victoria, Australia; possibly also inTasmania.

TAXONOMIC PLACEMENT. Lovenia.

zeisei Gagel, 1905∗1905 Maretia Zeisei 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 Museumof Lubeck, Germany.


AGE. Erratic boulders of Cenozoic age, presumably Mio-cene.

TAXONOMIC PLACEMENT. ? Hemipatagus; based on theoutline and the distribution of the aboral primary tubercles;type material very poor and in need of re-description; juniorsynonym of H. gottschei according to Lambert & Thiery(1924: 457); or juveniles of H. hoffmanni as discussed byGagel (1905: 543; he rejected this hypothesis because of thenaked plastron of H. zeisei, which could, however, also berelated to the poor preservation).

Hemipatagus, amisinterpreted Loveniid(Echinodermata: … · Hemipatagus, amisinterpreted...

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