Journal of Systematic Palaeontology 5 (2): 163–192 Issued 25 May 2007 doi:10.1017/S1477201906002021 Printed in the United Kingdom C The Natural History Museum Hemipatagus, a misinterpreted Loveniid (Echinodermata: Echinoidea)
Andreas Kroh∗ Naturhistorisches Museum Wien, Burgring 7, A-1010 Wien, Austria
SYNOPSIS The echinoid genus Hemipatagus is a poorly understood fossil spatangoid taxon that is now usually treated as a subjective synonym of the extant genus Maretia, but was originally subject to considerable dispute within the scientific community. Restudy of the species attributed to Hemipatagus and a range of presumably related spatangoids including Lovenia and Maretia has been carried out to solve the problem of its relationships. Cladistic analysis shows that Hemipatagus is not close to Maretia, but is closely related to Lovenia and should be placed in the Loveniidae, which is here confirmed as a monophyletic group. Characters of the adapical tuberculation suggest that in Hemipatagus an internal fasciole is present in early ontogeny, but lost in the adults. A clade comprising the genera Eurypatagus, Paramaretia and Platybrissus, and here named Eurypataginae, appears in all resulting trees as sister group to Maretia.
KEY WORDS Echinoidea, Hemipatagus, Loveniidae, Cenozoic, Eurypataginae nov.
Contents Introduction 163 History of nomenclature 164 Material and methods 164 Abbreviations and repositories 167 Results 167 Status of Hemipatagus 167 Phylogenetic affinities of Hemipatagus 169 Cladistic analysis 171 Spatial and temporal distribution of Hemipatagus and Maretia 172 Systematic Palaeontology 173 Order Spatangoida Claus, 1876 173 Family Loveniidae Lambert, 1905 173 Genus Hemipatagus Desor, 1858 173 Family Spatangidae Gray, 1825 174 Subfamily Eurypataginae nov. 174 Acknowledgements 174 References 174 Appendix 1 Characters scored in the cladistic analysis 178 Appendix 2 Data matrix 179 Appendix 3 Annotated list of the nominal species of Hemipatagus and Maretia 180
Introduction exploiting organic detritus as their prime food resource. Due to their life habit and evolutionary success they are Spatangoids are echinoids characterised by their bilaterally the most diverse of all extant sea urchin groups and one of symmetrical corona and highly modified ambulacra form- the echinoid clades with the best fossil record (Kier 1977). ing a complex respiratory apparatus. They are well adapted Their highly adapted test morphology provides many fea- to a life as burrowers and ploughers in mobile sediments, tures that can be employed in classification. Consequently * Corresponding author. E-mail address: [email protected] spatangoid taxonomy is, in general, rather straightforward. 164 A. Kroh
Particular features, however, have recently been shown (2) the broader, bilobed subanal fasciole in Hemipatagus (he to display high levels of convergence and homoplasy also disregarded this feature, although noted that it is im- (Markov & Solovjev 1995; Neraudeau et al. 1998; Villier portant for generic classification of some brissids). In the et al. 2004; Smith & Stockley 2005). Indeed, spatangoid ‘Treatise on Invertebrate Paleontology’ Fischer (1966: U609) phylogeny and higher-rank taxonomy has been heavily re- adopted Mortensen’s view and thus it was soon widely accep- liant on fascioles. These are narrow bands of small cili- ted in the scientific community, although some authors (e.g. ate spines secreting mucus that consolidates the walls of Philippe 1998: 222; Kroh & Harzhauser 1999: 163) still con- their burrows and protects the respiratory tube feet from fine tinued to express their doubts. Smith (2004: ‘The Echinoid mud (see Lawrence 1987). Unfortunately, fascioles were de- Directory’) noted the high similarity between Hemipatagus veloped several times during spatangoid evolution and can and Lovenia and proposed its inclusion in the Loveniidae. be secondarily lost (Villier et al. 2004; Smith & Stockley Part of the confusion surrounding Hemipatagus and 2005) and the emphasis placed on these structures has led to its relationship to Maretia and Lovenia resulted from considerable confusion and problems in spatangoid system- Desor’s (1858) statement that no fascioles are present in atics. The problem investigated here represents a case where Hemipatagus. Laube (1869) and Etheridge (1875), however, too much emphasis has been placed on presence and absence showed that a ‘spectacle-shaped’ (bilobed) subanal fasciole of fascioles. is present (see Fig. 2). Another major reason for the differing Hemipatagus (Pl. 1) is a genus widely distributed in the opinions resulted from the habit of basing conclusions on fea- Eocene to Miocene of Europe, occurring abundantly in the tures observed in various non-type species attributed to either famous Doberg section in northern Germany (Oligocene) genus (e.g. Duncan 1877). The type species of Hemipatagus and in Early Miocene strata of the Rhoneˆ Basin (Philippe and Maretia, in contrast, were rarely investigated with regard 1998), the Aquitaine Basin and the central Paratethys (Kroh to this question. Furthermore, most authors placed a heavy 2005). It is a spatangoid characterised by its heart-shaped emphasis on the presence or absence of an internal fasciole, outline and heterogeneous aboral tuberculation. The phylo- a feature which is often not readily observed in fossil speci- genetic and systematic relationship of Hemipatagus within mens. Moreover, it is known now that fascioles may be lost the spatangoids is uncertain, although it has commonly been during ontogeny, being present in juveniles and absent in the synonymised with Maretia (Pl. 2, figs 1 & 2), a Pliocene to adults of many spatangoids (e.g. Meoma (partial loss of the extant inhabitant of the Indo-West Pacific. The complicated subanal fasciole: Kier & Grant 1965; Chesher 1969), Para- nomenclatorial history of both genera is outlined below. maretia, Platybrissus and Eurypatagus (complete loss of the In this paper, the relationship between Hemipatagus, subanal fasciole: Mortensen 1950, 1951); see also Neraudeau´ Lovenia and Maretia is investigated using a cladistic ap- et al. 1998; Smith & Stockley 2005). proach based on morphological analysis. This generates a Although discussed at length it is not really clear why phylogeny and revised taxonomy for Hemipatagus and its Mortensen (1951: 23–27) finally chose to synonymise Hemi- relatives. patagus with Maretia. It would have been understandable if he had synonymised Hemipatagus with Lovenia, as he stated himself on p. 27 that ‘On the whole, the distinction between Maretia [here actually referring to Hemipatagus]andLove- History of nomenclature nia is, in case of the fossils, very difficult, the presence or absence of an inner fasciole – the only fully reliable distinc- The genus Hemipatagus was established in 1858 by Desor tion between these two genera – being, rather impossible to for the species Spatangus hoffmanni Goldfuss, 1829 (Fig. 1). ascertain beyond doubt, if the preservation is not very fine.’ It Subsequently, Agassiz (1873: 568) placed Hemipatagus in is difficult to understand why he rejected the shape of the sub- synonymy with Maretia. In the following years this place- anal fasciole as a diagnostic feature in Hemipatagus, while ment was subject to some debate. Some echinologists suppor- widely using it in brissid genera, particularly, as this feature ted Agassiz’s view (Cotteau 1885: 24–25; Fourtau 1920: 83), allows a very clear separation, with no known intermediate while others disagreed (Duncan 1877: 56–58, 1889: 222, 252; species. Mortensen (1951) also made a strong case against Lambert 1909: 107, 1915: 188, 1927a: 87). In his ‘A Mono- the use of the outline and the depth of the frontal sinus, graph of the Echinoidea’ Mortensen (1951: 23–27) discussed arguing that he would be forced to split Lovenia into sev- this matter at length and finally considered Hemipatagus eral genera when using these features to distinguish between Desor, 1858 a junior synonym of Maretia Gray, 1855. His Hemipatagus and Maretia. Yet, it is well known that features opinion was based on the observation that only one of the that are useful to separate some taxa might be highly variable three features discussed in the literature seemed important in others, not allowing a clear separation. to him (the presence of internal ampullae associated with spine tubercles in Hemipatagus). He argued that one feature alone was not enough to justify separation of the two gen- Material and methods era. Moreover, he had also observed internal swellings that he assumed were ampullae in juveniles of Maretia planu- Two specimens of the extant Maretia planulata (Lamarck, lata (Lamarck, 1816) and saw them as proof for his point of 1816) from the Gulf of Siam, Mauritius (NHMW, 2nd view. In addition, Mortensen noted that the presence of am- Zoological Department, collection Invertebrata-Varia no. pullae was not verified in the type species of Hemipatagus. 12715), three specimens of Lovenia elongata (Gray, 1845) The other two features listed by him were: (1) a more heart- from the Red Sea (NHMW, 2nd Zoological Department, col- shaped outline with deeper frontal notch (he disregarded this lection Invertebrata-Varia no. 12714) and 24 well preserved feature, due to the fact that large differences in the depth of the specimens of Hemipatagus hoffmanni (Goldfuss, 1829) frontal notch are also present in other genera, e.g. in Lovenia); 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, near B¨unde, 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 Gr¨ubern, Austria. 11–12, NHMW 1914.VII.22b; 13–14, 1914.VII.22a. Figs 15, Hemipatagus martensii (Ebert, 1889), Late Oligocene of Doberg, near B¨unde, 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 B¨unde, Germany, MB.E4840. Hemipatagus – a misinterpreted Loveniid 167
Figure 1 Hemipatagus hoffmanni (Goldfuss, 1829), syntype and figured specimen of Goldfuss (1829); specimen PIUB 357a. A,aboralview; B, right lateral view; C,oralview.
Germany (NHMW, Geological-Palaeontological Depart- morphological differentiation amongst the taxa investigated ment 1852.II.1500, 1852.II,1501, 1855.I.800, 1855.I.801, many characters were multi-state. Characters have neither 1862.XLIII.43, 1874.XXIX.1178 and MB, Palaeontology been polarised nor differentially weighted. Department MB.4839–4843, plus 6 unregistered speci- mens) were examined. In addition, the type-material of Hemipatagus hoffmanni was located (PIUB 357a-h) and Abbreviations and repositories a photograph of Goldfuss’s original specimen (PIUB 357a) is presented for the first time (Fig. 1). Furthermore, GSI, Geological Survey of India; KM, Krahuletz Museum, the type material of Hemipatagus perornatus (Schaffer Eggenburg, Austria; NHMW, Naturhistorisches Museum 1912:2 syntypes – KM F/0127 and F/0128) was located and Wien, Austria; MB, Naturkunde Museum Berlin, Germany; reinvestigated, along with some new specimens (NHMW PIUB, Institute of Palaeontology, University of Bonn, Geology-Palaeontology Department 1914.VII.22a-c, Germany (Goldfuss Museum). 1999z0049/0003, 2002z0159/0001; see also Kroh 2005). Cladistic analysis was carried out using the programme PARS from the Phylogeny Inference Package PHYLIP (Felsenstein 2002), using the ‘Randomize input order of Results species option’ (1000 replicates). The program CONSENS from the same package was used to obtain majority rule Status of Hemipatagus consensus trees. The analysis involved 30 species from the Spatangidae and Loveniidae together with Eupatagus valen- Material from the type species of Hemipatagus, Lovenia and ciennesi Agassiz, in Agassiz & Desor, 1847. A total of 50 Maretia was investigated in detail, focusing on tubercula- characters were scored based on test shape, apical disc struc- tion types and arrangement, gross morphology, pore morpho- ture, plating pattern, tuberculation, sphaeridia, pore and peri- logy, fascioles (Fig. 2) and, most importantly, plating patterns proct structure. A number of additional, phylogenetically (Figs 3 & 4, Table 1). uninformative characters (basal synapomorphies) as well as When comparing H. hoffmanni with M. planulata, the some highly variable characters were excluded. The data following differences were observed in M. planulata: (1) the matrix and description of the remaining 42 characters are frontal sinus is shallower and often indistinct; (2) the lab- found in Appendices 1 and 2. Due to the high degree of 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, Gr¨ubern, 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, Gr¨ubern, Austria. Interambulacra shaded, fasciole finely stippled, small stippled circles are sphaeridial pits.
end is bluntly pointed not transversely truncated; (4) even in in H. hoffmanni (compare Figs 3 and 4A); (11) a conspicuous large specimens the primary tubercles do not develop true field of coarse tubercles adjacent to adapical ambulacrum III ampullae or strongly sunken areoles, especially not on the is well developed in H. hoffmanni but lacking in M. planu- oral surface; (5) the plastron is much more strongly con- lata (Pl. 3, figs 1–5). These differences provide ample means stricted between the first and second pair of episternal plates to separate those two genera. Lovenia and Hemipatagus are (Fig. 4A); (6) the outline of the test is oval, antero-posteriorly more difficult to separate. Hemipatagus differs mainly by the elongated, not cordiform; (7) the subanal fasciole is oval and absence of the internal fasciole and its different petal struc- rather high instead of bilobed, very wide and low (Table ture (never forming continuous arcs as in most species of 1); (8) the labrum and the primordial plates of the interam- Lovenia). bulacra extend much further (Fig. 4A); (9) there are more When other species (e.g. H. ocellatus, H. grignonensis, phyllodal pores (14–15) in ambulacra II and IV (only 6–7 H. girundicus, M. planulata pliocenica, M. ? cordata: see Ap- in H. hoffmanni); (10) ambulacra II and IV are only slightly pendix 3) attributed to the three genera in question are taken constricted ambitally, whereas they are strongly constricted into account, it becomes apparent that most of the features
Figure 4 Camera lucida drawings of the oral plating. A, Maretia planulata (Lamarck, 1816) NHMW-EV 12715, Recent, Gulf of Siam, Mauritius; B, Lovenia elongata (Gray, 1845) NHMW-EV 12714, Recent, Red Sea. Interambulacra shaded, fasciole finely stippled, small stippled circles are sphaeridial pits. Hemipatagus – a misinterpreted Loveniid 169
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. out by various authors recently (e.g. Jeffery 1998; Neraudeau´ Concerning the structure of the primary tubercles, however, et al. 1998; Smith & Jeffery 2000), in the past too much em- Mortensen (1951) was quite correct to question their value phasis was placed on the presence/absence of certain types as a diagnostic feature. While some species of Hemipatagus of fascioles in the adult animal in developing a supra-generic have truly camellate tubercles (i.e. tubercles with associated classification of the spatangoids. It has been clearly estab- pouch-like ampullae forming a bulge on the interior of lished that fascioles may be lost during ontogeny, a fact that the test) on the oral and aboral surface (e.g. H. ocellatus, has to be taken into account when trying to resolve this H. perornatus, H. girundicus), others lack them either over question (e.g. Mortensen 1951; Kier & Grant 1965; Chesher the oral surface (e.g. H. hoffmanni) or completely (H. carolin- 1969). ensis, H. argutus). In those species where internal ampullae Here the conspicuous field of tubercles in adapical in- are lacking, the areoles are, nevertheless, strongly sunken. terambulacral columns 2b and 3a (Pl. 3, figs 2–5) comes However, the other features, especially the distribution of the into play. In all genera (except Hemipatagus) where this fea- aboral primary tubercles, number of phyllodal pores and all ture is present it is invariably associated with an internal features related to the plating pattern, are quite stable and fasciole (Breynia, Echinocardium, Lovenia, Pseudolovenia allow a consistent separation (see ‘Emended diagnosis’ and and Vasconaster). It is, therefore, possible to assume that ‘Remarks’, below, for details). an internal fasciole was present during the early ontogeny of Hemipatagus. The fact that this arrangement is not well developed in some species of Hemipatagus does not neces- Phylogenetic affinities of Hemipatagus sarily mean that these species never possessed an internal fasciole. There are some species of Lovenia in which this The systematic placement of Hemipatagus remains difficult. special arrangement is not very conspicuous (e.g. L. sub- Most spatangoid families are based on the type of primary carinata). By contrast, such an arrangement has never been tuberculation, differentiation of fascioles, structure of the ap- observed in any member of the family Spatangidae (including ical disc and plastron structure (see Fischer 1966). As pointed Maretia). 170 A. Kroh
Plate 3 Adapical tuberculation and phyllodes of Hemipatagus, Lovenia and Maretia. Figs 1 & 9, Maretia planulata (Lamarck, 1816), extant, Gulf of Siam, Mauritius, NHMW, 2nd Zoological Department, collection Invertebrata-Varia no. 12715. Figs 2 & 8, Lovenia elongata (Gray, 1845), extant, Red Sea, NHMW, 2nd Zoological Department, collection Invertebrata-Varia no. 12714. Figs 3–4, 6, Hemipatagus hoffmanni (Goldfuss, 1829), all Late Oligocene of Doberg, near B¨unde, 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 Gr¨ubern, Austria. 5, 1914.VII.22a; 7, NHMW 1914.VII.22b. Hemipatagus – a misinterpreted Loveniid 171
Figure 5 Result of the cladistic analysis based on the full dataset (A) and the reduced dataset (B). Italic numbers beside the nodes indicate bootstrap values obtained (1000 replicates). Numbers in normal font beside the nodes in the 50% majority consensus tree (A) indicate the percentage of times that this node occurred in the full set of most parsimonious trees.
On the basis of its strong similarity to Lovenia and the between a clade containing Eurypatagus, Maretia, Para- inferred loss of the internal fasciole during early ontogeny, maretia, Platybrissus and Spatangus (clade A) and a second Hemipatagus is here placed in the family Loveniidae. clade containing the classical Loveniidae (Breynia, Ech- inocardium, Homolampas, Lovenia, Pseudolovenia) plus Hemipatagus and Pseudomaretia (the latter traditionally Cladistic analysis placed in the Spatangidae: see Fischer 1966) (clade B). Due to the presence of several missing characters in Maretia cordata To test the inferences based on the subjective evaluation and two old species tentatively placed in Hemipatagus (H.? of the morphological features of Hemipatagus, Lovenia and argutus and H.? carolinus, from the Middle Eocene and the Maretia, a cladistic analysis was carried out. In all analyses Oligocene, respectively) 30 equally parsimonious trees were Hemipatagus hoffmanni and H. ocellatus form a sister group found. The majority consensus tree of these trees is shown taxon to the extant species of Lovenia. Exclusion of the spe- in Fig. 5A. The placement of M. cordata switches between cies with a high number of unknown characters (Maretia just two positions (at the base of clade A versus as sister- carinata, Hemimaretia elevata, Hemipatagus? subrostratus) group taxon to the two spatangids). Hemipatagus? argu- results in a very stable tree topology with a deep split tus and H.? carolinus, in contrast, are resolved either as 172 A. Kroh
Figure 6 Spatial distribution of the genera Hemipatagus and Maretia. Open symbols show taxa of doubtful generic affinity. Distribution of extant and fossil M. planulata is based on data in: Stockley 1927; Brighton 1931; Mortensen 1951; Richer de Forges & Menou 1988; Baker & Rowe 1990; Kerr et al. 1992; Liao & Clark 1995; Rowe & Gates 1995, USSMDC 1999; Lindley 2003.
stem-group loveniids (in the majority of the trees) or as trees (42), the majority consensus trees of which show a very crown-group loveniids forming a polyphyletic clade with similar topology to the trees discussed above. Pseudolovenia, H. hoffmanni, H. ocellatus and the extant The present analysis clearly shows that Hemipatagus species Lovenia. In just one of the 30 trees H.? argutus and is a loveniid that evolved during the Middle or Late Eocene H.? carolinus are resolved as sister group to Echinocardium. from a common ancestor with Lovenia. Despite the superfi- Exclusion of M.? cordata, H.? argutus and H.? carolinus cial similarities with Maretia it is clearly not closely related from the analysis results in a single most parsimonious tree to that genus. This is also reflected by their temporal and (Fig. 5B) very similar to the majority consensus tree of the spatial distributions, which are disjunct with respect to each former analysis but with better resolution within clade B. It other (see below). Earlier statements that Maretia and Hemi- is apparent that the loveniids as a whole are a very robust patagus are synonymous (e.g. Mortensen 1951) were based group with well-defined monophyletic genera. The only ex- on misinterpretation and erroneous weighting of a small set ception is Lovenia, which is monophyletic, but paraphyletic of characters. if Pseudolovenia is excluded. The present analysis suggests In a recent paper on spatangoid phylogeny, Stockley that Pseudolovenia should either be synonymised with Love- et al. (2005) present the results of a cladistic analysis in- nia or that Lovenia should be split into two different genera. volving several of the genera discussed here. In the strict The topology of the Lovenia clade resulting from the present consensus tree resulting from their morphological analysis analysis is very similar to that published by Madon (1994) (Stockley et al. 2005: fig. 2) the Loveniidae are a poly- using a different dataset. Another new result is the placement phyletic clade. According to co-author A.B. Smith (pers. of the Pseudomaretia-clade in the loveniids. Earlier Pseudo- comm., 01.11.2005), however, their analysis produced a very maretia (and the junior synonym Goniomaretia) were con- similar topology to the one presented here when restricted sidered to be spatangids (Fischer 1966) or maretiids. The to the micrasterine clade alone. Smith relates the differences genera Eurypatagus, Platybrissus and Paramaretia form a between their fig. 2 and the present tree (Fig. 5) to the global distinct clade in the present analysis and are a sister group to parsimony analysis employed in Stockley et al. (2005), which Maretia planulata in most trees. This clade appears also in fails to group the loveniid taxa into a single clade. the strict consensus tree published by Stockley et al. (2005: fig. 2), where it also forms the sister group of Maretia.The Eurypatagus–Platybrissus–Paramaretia-clade is here named Spatial and temporal distribution of Hemipatagus Eurypataginae (see ‘Systematic palaeontology’ below, for a and Maretia formal definition). Inclusion of a more distant outgroup (Brissus unicolor) Hemipatagus and Maretia are well separated in time and results in a distinctly higher number of most parsimonious space. The former occurs most commonly in the Eocene to Hemipatagus – a misinterpreted Loveniid 173
Middle Miocene of the Mediterranean region. In addition, mens (Figs 3A, B); (7) moderately elongated labral plate not there are a few species on the east coast of North America extending beyond the second ambulacral plates (Figs 3A, B); (see Fig. 6). Maretia, by contrast, occurs first in the Plio- (8) prominent oral tubercles arranged in distinct rows, in cene and is restricted to the Indian Ocean, the Indo–Malayan some species with deeply sunken areoles and helically spir- Archipelago and the West Pacific Islands (farthest east alled parapet (e.g. Pl. 3, fig. 7); (9) three to four ambulacral occurrences are on Hawaii). The only potential record of plates extending into the subanal fasciole in ambulacrum I Hemipatagus from the Pliocene and from the Indo–West Pa- and V, respectively. cific region is H.? bandaensis. The data available for this taxon are not sufficient to firmly establish its generic affinity. INCLUDED SPECIES. Apart from the type species H. hoff- It might also be a misidentified Lovenia. A re-description and manni (Goldfuss, 1829) the following species can be con- new illustrations are urgently needed. fidently assigned to Hemipatagus based on the published Another issue apparent in Fig. 6 is the disjunct distri- information: H. calvimontanus (Cotteau, 1885); H. car- butional pattern of the extant Maretia planulata. Apparently olinensis (Kier, 1997); H. desmoulinsi (Cotteau, 1863); H. there are no records from Western India, Pakistan, the Persian girundicus Lambert, 1915; H. guebhardi (Lambert & Savin, Gulf or the Indian Ocean coast of the Arabian Peninsula. In 1902); H. hungarica (Vadsz, 1915); H. marianii (Airaghi, view of this the colouration differences noted by Mortensen 1903); H. martensii (Ebert, 1889); H. ocellatus (Defrance, (1951: 36) may be more meaningful than he considered they 1827) (=H. nicoleti (Agassiz, 1839) = H. perornata (Schaf- were. He described a whitish to yellow colouration in Indian fer, 1912)); H. pellati Cotteau 1863; H. saccoi (Airaghi, Ocean populations, versus Malayan specimens, which are 1905); H. soubellensis (Peron & Gauthier, in Cotteau et al., usually dark. As Mortensen (1951: 36) could find no apparent 1891); H. tenuis (Peron & Gauthier, in Cotteau et al., 1891). morphological differences, he considered this phenomenon In addition, there are several species that might also belong to be local variation. Further investigations are needed to to Hemipatagus, but which are too poorly known at present judge whether there might actually be two species. A sim- to establish their generic affinities with certainty: H.? ar- ilar situation has recently been discovered in Asthenosoma gutus Clark, in Clark & Twitchell, 1915; H.? bandaensis by Weinberg & de Ridder (1998), who discovered that the Martin, in Jeannet & Martin, 1937; H.? carolinae (Gagel, echinoids previously classified as A. varium belong to two 1905); H.? gauthieri (Lambert, in Lambert & Thiery,´ 1924); distinct species, which are geographically separated. H.? gottschei (Gagel, 1905); H.? grignonensis (Cotteau, 1880); H.? hispanica (Cotteau, 1890); H.? mauritanica (Pomel, 1885); H.? H.? regiomontanus (Mayer, 1860); H.? sambiensis (Beyrich, 1848); H.? subrostratus Clark, in Systematic Palaeontology Clark & Twitchell, 1915; H.? tuberculatus (Peron & Gauthier, in Cotteau et al., 1885); H.? zeisei (Gagel, 1905). Order SPATANGOIDA Claus, 1876 See Appendix 3 for original generic assignments and more data on the individual species listed here. Family LOVENIIDAE Lambert, 1905 Genus HEMIPATAGUS Desor, 1858 REMARKS. There are a number of genera that show superfi- cial similarities to Hemipatagus. All of them, however, can TYPE SPECIES. Spatangus hoffmanni Goldfuss, 1829; by ori- be clearly separated on the basis of structural features (see ginal designation (Desor 1858: 416). (nomen correctum pro Table 1). Lovenia differs by the presence of an internal fas- Spatangus hofmanni Goldfuss, 1829 – Grateloup (1836: 175) ciole, high number of ambulacral plates extending into the (presumably intentionally) changed the name to S. hoffmanni. subanal fasciole (up to 11, although in some species there According to the ICZN rules this is an unjustified emenda- are only three or four as in Hemipatagus) and less lanceol- tion. However, as the name S. hoffmanni has been in prevail- ate petals; in most species of Lovenia ambulacral columns ing usage since 1836 and is attributed to the original author Ib–IIa and IVb–Va form continuous arcs (not observed in and date of the original spelling (e.g. des Moulins 1837: any known Hemipatagus species); in some species of Hemi- 244; Agassiz & Desor 1847: 7), it has to be deemed to be a patagus the aboral and/or oral tubercles are not camellate, justified emendation according to the ICZN (1999: 4th edn while this is always the case in Lovenia (see also Fig. 4B; Pl. Article 33.2.3.1.)). 2, figs 3–4; Pl. 3, fig. 8). Maretia differs by possessing an extremely shallow RANGE. Eocene to Middle Miocene (Serravallian). frontal notch, the distribution of the aboral primary tubercles EMENDED DIAGNOSIS. Cordiform spatangoid differing from (primary tubercle reach the tips of the posterior paired petals), other loveniids by its (1) heterogeneous aboral tubercula- the lack of strongly sunken areoles or camellate aboral and tion with large crenulate or non-crenulate primary tubercles oral primary tubercles, the strongly crenulate nature of the (Table 1); (2) conspicuous aboral primary tubercles in inter- aboral primary tubercles, the lack of a conspicuous field ambulacra 1–4, but missing in interambulacrum 5 and the of coarse tubercles in adapical interambulacral columns 2b posterior part of interambulacral columns 1a and 4b (see and 3a, its oval to shield-shaped subanal fasciole, the longer Table 1); (3) strongly sunken areoles or even the presence of primordial plates of the paired interambulacra (plates 1a1, camellae in some species; (4) presence of a subanal fasciole 2a1, 3a1 and 4a1: which extend to the third or fourth am- in adult specimens, which is strongly bilobed (Figs 2A, B); bulacral plate of the adjacent ambulacra in Maretia, while (5) presence of a conspicuous field of coarse tubercles adja- they do not extend beyond the second ambulacral plate in cent to adapical ambulacrum III in interambulacral columns Hemipatagus or Lovenia), the labrum which extends to the 2b and 3a (Pl. 3, figs 3–5); (6) strong constriction of ambu- third or fourth ambulacral plate (not extending beyond the lacra II and IV near the margin on the oral side in adult speci- second ambulacral plate in Hemipatagus and Lovenia)and 174 A. Kroh the less constricted adoral ambulacra 2 and 3 (Fig. 4A; Pl. 2, (partially tuberculate in Maretiinae), (3) structure of the figs 1–2). petals (narrowed, distally open, extending almost to the am- Breynia differs by its fully tuberculate sternum, pres- bitus) and (4) labrum extending to the 4th plates of ambulacra ence of an internal and a peripetalous fasciole, shield-shaped I and V. Differing from the Spatanginae by their (1) strongly form of the subanal fasciole, shape of the petals (with am- elongated labrum, (2) naked sternal plates (fully tuberculate bulacral columns Ib–IIa and IVb–Va forming a continuous in Spatanginae), (3) crowding of the adoral plates of ambu- arc in some species), by the labrum and primordial plates of lacra II and IV and (4) the structure of the petals (narrowed, the paired interambulacra, which extend to the third ambu- distally open, extending almost to the ambitus). lacral plate, the presence of primary tubercles on the primor- dial plates of the paired interambulacra; by the smaller oral REMARKS. Additional features that are characteristic, but primary tubercles, which are never sunken or camellate; and not exclusive, for this family are: (1) an oval outline with the high number of ambulacral plates extending into the sub- only feeble frontal sinus if present at all; (2) heterogeneous anal fasciole (usually six to eight), by the distribution of the aboral tuberculation (enlarged primary tubercles developed aboral primary tubercles (confined to the area within the peri- in most forms; where developed, they are arranged without petalous fasciole, but reaching the tips of the posterior paired distinct pattern). petals) and the near-straight contact between the sternal and The genus Platybrissus is another potential member of episternal plates. the Eurypataginae. In some analyses with a different out- Eupatagus differs by its fully tuberculate sternum, pres- group (Brissus unicolor), however, this genus forms the sis- ence of a peripetalous fasciole, oval shape of the subanal ter group to Eurypatagus, Paramaretia and Maretia.There- fasciole, the different nature of the aboral and oral primary fore it is only tentatively included in the Eurypataginae at tubercles (never camellate or strongly sunken), different dis- present. tribution of the aboral primary tubercles (see Table 1), by the An independent cladistic analysis presented by labrum and primordial plates of the paired interambulacra (Stockley et al. 2005: fig. 2) likewise shows a monophyletic which extend to the third ambulacral plate and by the lack of clade containing Eurypatagus, Paramaretia,andPlatybris- any frontal notch. sus. As in most trees resulting from the present analysis, Pseudolovenia differs by its narrow, poorly-defined Stockley et al. (2005) found a sister-group relationship petals with widely spaced, less well developed pores which between this clade and Maretia. continue to the ambitus, restriction of the aboral primary tubercles to the interambulacral columns adjacent to ambu- lacra II and IV, its labrum which extends to the third am- bulacral plates, the presence of an internal fasciole and the Acknowledgements higher number of plates extending into the subanal fasciole This study was supported by the Austrian Science Fund (five). (FWF) via project no. P-13466-Bio to Werner E. Piller (Uni- Pseudomaretia and Goniomaretia differ by possessing versity of Graz). The critical reviews and helpful comments an ethmolytic apical disc with only three gonopores (no gono- of Andrew B. Smith, Rich Mooi and another anonymous re- pore in genital plate 2), lack of a frontal notch, more reduced viewer greatly improved the paper. Furthermore, I would like nature of the anterior poriferous zones of the anterior paired to express my sincere thanks to the staff of the Geological De- petals (well developed pores only in the most distal plates), partment of the Natural History Museum Vienna for provid- oval to shield-shaped subanal fasciole and the absence of ing scientific facilities. Special thanks to Johannes Tuzar and most or all aboral primary tubercles. Andreas Thinschmidt (both Krahuletz Museum, Eggenburg) Paramaretia differs by the more reduced nature of the for helping to locate the type-material of H. peronatus and to anterior poriferous zones of the anterior paired petals (no Manuel Kunz, Martin Sanders and Georg Oleschinski (all at large, well developed pores present at all), its labrum which the Goldfuss Museum, Bonn) for locating the type material extends to the fourth ambulacral plates, the primordial plates of H. hoffmanni and providing photographs of it. Thanks also extend to the fourth to sixth ambulacral plates, the loss of the to Alice Schumacher (NHM Vienna) for taking the photo- subanal fasciole in adults, having a plastron which is only graphs. weakly indented behind the first pair of episternal plates, its naked sternal plates and, most obviously, by the distribu- tion of the aboral primary tubercles which are present in all interambulacra, without obvious pattern. References Family SPATANGIDAE Gray, 1825 Agassiz, A. 1864. Synopsis of the echinoids collected by Dr. W. Stimpson on the North Pacific Exploring Expedition under the command of Cap- tains Ringgold and Rodgers. Proceedings of the Academy of Natural Subfamily EURYPATAGINAE nov. Sciences of Philadelphia 15 (1863): 352–361. TYPE GENUS. Eurypatagus Mortensen, 1948: 133. —— 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. OTHER GENERA INCLUDED. Paramaretia Mortensen, 1950, —— & Clark, H. L. 1907. Preliminary report on the Echini collected ? Platybrissus Grube, 1866 (see Fig. 5). 1906, from May to December, among the Aleutian Islands, in Ber- ing Sea, and along the coasts of Kamchatka, Saghalin, Korea, and RANGE. Miocene to Recent. Japan, by U.S. Fish Commission steamer “Albatross”, in charge of Lieut.-Commander L. M. Garrett, U.S.N., commanding. Bulletin of the DIAGNOSIS. Differing from the Maretiinae by their (1) lack Museum of Comparative Zoology at Harvard College 51(3): 107– of any fascioles in adult individuals, (2) naked sternal plates 139. Hemipatagus – a misinterpreted Loveniid 175
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Lambert, J. 1902. Description des echinides´ de la Province de Barcelone. Echinolampidae, Clypeastridae, Arachnidae, Laganidae, Fibulariidae, Fasc. 1. Memoires´ de la Societ´ eG´ eologique´ de France, Paleontologie´ Urechinidae, Echinocorythidae, Palaeostomatidae, Micrasteridae, Pa- 9: 1–57, 4 pls. laepneustidae, Hemiasteridae, and Spatangidae. Bulletin of the Smith- —— 1905. Notes sur quelques Echinides´ eoc´ eniques´ de l’Aude et de sonian Institution, U.S. National Museum 100(14/3): 93–140. l’Herault.´ Pp. 129–164, pl. 6 in Catalogue Descriptif des Fossiles —— 1950. A Monograph of the Echinoidea. V, 1. Spatangoida I. Nummulitiques de l’Aude et de l’Herault´ . Annales de l’Universitede´ Protosternata, Meridosternata, Amphisternata I. Palæopneustidæ, Lyon, Nouvelle Serie,´ I. Sciences, Medecine´ 17: 1–184. Palæostomatidæ, Aeropsidæ,¨ Toxasteridæ, Micrasteridæ, Hemiaster- —— 1909. Description des echinides´ fossiles des terrains mioceniques´ idæ. C. A. 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Essai de Nomenclature Raisonnee´ des —— 1887. Paleontologie´ ou description des animaux fossiles de l’Algerie.´ Echinides. L. Ferriere,` Chaumont, fasc. 6–7: 385–512, pls 10–11, Zoophytes. 2e fascicule – Echinodermes.´ 2e livraison. Adolphe 14 (Dec. 1924). Jourdan, Alger, 344 pp. Laube,G.C.1869. Uber¨ einige fossile Echiniden von den Murray cliffs in Richer de Forges, B. & Menou, J. L. 1988. The echinoderms of the soft Sud-Australien.¨ Sitzungsberichte der Kaiserlichen Akademie der Wis- bottoms of the south-western lagoon of New Caledonia. Proceedings senschaften, mathematisch-naturwissenschaftliche Classe, Abtheilung of the 6th International Coral Reef Symposium Australia 2: 21–35. I 59(1): 183–198, 1 pl. Rowe,F.W.E.&Gates,J.1995. Echinodermata. Pp. xiii+510 in A. Wells Lawrence, J. M. 1987. A Functional Biology of Echinoderms. Croom (ed.) Zoological Catalogue of Australia. Volume 33. CSIRO Australia, Helm, London & Sydney, 340 pp. Melbourne. Leske, N. G. 1778. 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Abhandlungen der Geologischen Reichsanstalt 22(2): 187–192, Madon, C. 1994. Phylogenetic versus phenetic analysis in the spatangoid pls 58–60. Lovenia. Pp. 765–768 in B. David, A. Guille, J.-P. Feral´ & M. Roux Seba, A. 1734–1765. Locupletissimi rerum naturalium thesauri accurata (eds.) Echinoderms through Time – Proceeding of the 8th Interna- descripto, et iconibus artificiosissimis expressio, collegit, digessit, tional Echinoderm Conference Dijon/France, 6–10 Sept. 1993.A.A. descripsit et ad pingendum curavit per universam physices historiam, Balkema, Rotterdam. ...Tome 3, 116 pls., Amstelaedami. Manzoni, A. 1879. Gli echinodemi fossili dello Schlier delle colline di Bo- Smith, A. B. (ed.) 2004. The Echinoid Directory. World Wide Web logna. Denkschriften der kaiserlichen Akademie der Wissenschaften, electronic publication. Available at: http://www.nhm.ac.uk/science- mathematisch-naturwissenschaftliche Classe 39: 149–164, 4 pls. projects/echinoids (accessed 27 July 2004). 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Fossil spatangoids (Echinodermata) from the Palaeontology of Victoria; or,Figures and Descriptions of the Victorian Indian Subcontinent – a review. Journal of the Palaeontological Society Organic Remains. Geological Survey of Victoria, Melbourne. of India 49: 125–149, 7 pls. Michelin, H. 1863. Echinides´ et Stellerides´ recueillis par M. Maillard, Stockley, B., Smith, A. B., Littlewood, T., Lessios, H. A., MacKenzie- ingenieur´ colonial de 1856 a 1860. Pp. 1–7 in L. Maillard (ed.) Dodds, J. A. 2005. Phylogenetic relationships of spatangoid sea Notessurl’IledelaReunion´ (Bourbon). Deuxieme` edition´ . Dentu, urchins (Echinoidea): taxon sampling density and congruence between Paris. morphological and molecular estimates. Zoologica Scripta 34: 447– Mortensen, T. 1948. Contributions to the Biology of the Philippine 468. Archipelago and adjacent regions. Report on the Echinoidea col- Stockley,G.M.1927. Neogene Echinoidea from Zanzibar Protectorate. lected by the United States Fisheries Steamer ‘Albatross’ during Pp. 103–117 in Report on the palaeontology of the Zanzibar Protect- the Philippine Expedition, 1907–1910. Part 3: The Echinoneidae, orate. His Majestys Stationery Office, London. 178 A. Kroh
Sturt, C. 1833. Two expeditions into the interior of Southern Australia Weinberg, S. & de Ridder, C. 1998. Asthenosoma marisrubri n. sp. during the years 1828, 1829, 1830 and 1831, with observations on the (Echinodermata, Echinoidea) from the Red Sea. Beaufortia 48(3): 27– soil, climate and general rersources of the colony of New South Wales, 46. Volume 2. Smith, Elder and Co., Cornhill, London, 271 pp. Woods, J. E. T. 1859. Remarks on a Tertiary deposit in South Australia. Szor¨ enyi,´ E. 1952. A Maretia hungarica Vadasz´ mjocen´ koru´ Transactions of the Philosophical Institute of Victoria 3: 85–94. faj horvatartozas´ anak´ tiszat´ azsa.´ Foldtani¨ Kozl¨ ony¨ 82: 302– —— 1862. Geological observations in South Australia: principally in the 303. district south-east of Adelaide. Longman, Green, Longman, Roberts Tate, R. 1894. Critical list of the Tertiary Mollusca and Echinodermata & Green / H.T. Dwight, London & Melbourne, xviii-404 pp. of New Zealand in the collection of the Colonial Museum. Reports of —— 1865. On some Tertiary deposits in the Colony of Victoria, Australia. Geological Exploration New Zealand 1892–1893: 121–127. Quarterly Journal of the Geological Society of London 21: 389–394. USSMDC 1999. Marine Biological Survey, Peacock Point Outfall, —— 1878. On the Tertiary deposits of Australia. Journal and Proceedings Wake Atoll June 1998NODC #0000247 Vers. 1. Department of the of the Royal Society of New South Wales 11 (1877): 65–82. Army, U.S. Army Space and Missile Defense Command, Hunts- Wright,T.W.1864. On the fossil Echinidæ of Malta. With additional ville, Alabama. Available at: http://www.coris.noaa.gov/metadata/ notes on the Miocene beds of the Island, and the stratigraphical distri- records/html/nodc_0000247_wake_atoll_reefs_1998.html. bution of the species therein; by A. Leith Adams. Quarterly Journal Vadasz,´ E. 1915. Die mediterranen Echinodermen Ungarns. Geologica of the Geological Society of London 20: 471–491, 2 pls. Hungarica 1(2): 79–253, 6 pls. Zachos,L.G.1993. Occurrence of the spatangoid echinoid Maretia ar- Villier, L., Neraudeau,´ D., Clavel, B., Neumann, C. & David, B. guta (Clark) in the Middle Eocene of Texas. Journal of Paleontology 2004. Phylogeny of Early Cretaceous spatangoids (Echinodermata: 67: 148–150. Echinoidea) and taxonomic implications. Palaeontology 47: 265– Zittel, K. A. 1867. II. Fossile Mollusken und Echinodermen aus Neu- 292. Seeland. Pp. 15–68 in F. von Hochstetter, M. Hornes¨ & F. R. von Wanner, J. 1931. 13. Echinodermata. Pp. 436–460 in De Palaeontologie Hauer (eds) Palaontologie¨ von Neu-Seeland. Beitrag zur Kenntniss der en stratigraphie van Nedelandsch Oost-Indie.¨ Leidsche Geologische fossilen Flora und Fauna der Provinzen Auckland und Nelson. Novara Mededeelingen 5 (K. Martin Festbundel): 436–460. Expedition. Geologischer Theil. k.k. Hof- und Staatsdruckerei, Wien.
Appendix 1 Characters scored in the cladistic analysis
Shape 1. Test outline: cordate (0); oval (1). 2. Frontal sinus: absent (0); weakly developed (1); well developed (2). Apical disc 3. Apical disc, position: anteriorly displaced (0); subcentral (1); posteriorly displaced (2). 4. Number of gonopores: four (0); three (1). Ambulacra 5. Shape of the paired ambulacra: petaloid (0); subpetaloid (1); non-petaloid (2). 6. Anterior PZ of PPP and posterior PZ of APP: form an angle (0); form a crescentic line (1). 7. Petals: broad, i.e. IPZPZ (0); moderately wide IPZ∼PZ (1); narrow, i.e. IPZPZ (2). 8. Pores in the paired petals: elongate isopores (0); other (1). 9. Rudimentary pores in the anterior PZ of the APP: none (0); in the adapical plates, gradual change to normal-sized pores (1); in the adapical plates, abrupt change (2); in all but the most distal plates (3). 10. Adoral ambulacra II and IV between phyllodes and margin: only slightly constricted, with roughly isometric plates (0); strongly constricted, with sinuous, elongated plates (1). 11. Phyllodal plates: plates roughly isometric and few plates included in phyllodes II and IV (up to plates II.4a and IV.4a, respectively) (O); plates elongated, crowded and numerous plates in phyllodes II and IV (usually extending beyond plates II.5a and IV. 5a, respectively) (1). 12. Number of pores within the subanal fasciole (adult condition): up to five (0); usually more than 5 (1). Interambulacra 13. Labrum, posterior extension to the: 1st (1); 2nd (2); 3rd (3); 4th (4) ambulacral plates. 14. Labrum, shape: broad, short crescent, length<
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 11111 11112 22222 22223 33333 33334 44 Character 12345 67890 12345 67890 12345 67890 12345 67890 12 Brissus unicolor (Leske,1778) 10000 02010 10100 13733 10000 00000 00033 00100 11 Eupatagus valenciennesi Agassiz,1847 10000 00011 00321 03222 10201 10110 10000 00210 21 Spatangus inermis Mortensen,1913 02000 00010 00100 03444 00001 10101 10133 00100 10 Spatangus purpureus M¨uller,1776 02000 00010 10100 03444 00001 10111 10100 00100 10 Eurypatagus grandiporus Mortensen,1948 10000 01010 10422 14544 10121 10111 10000 10200 00 Eurypatagus ovalis Mortensen,1948 10000 01010 10421 14554 10121 10111 10000 10200 00 Eurypatagus parvituberculatus (Clark,1924) 10000 01010 10422 14443 10121 10111 10000 10200 00 Platybrissus roemeri Grube,1865 10000 00010 10421 14343 10211 10100 10000 00200 00 Maretia carinata Bolau,1873 10000 00010 00321 0???? 1011? ?0111 11000 0?210 20 Maretia? cordata Mortensen,1948 02000 00010 00321 0343? 1021? 10111 11000 0?210 20 Maretia planulata (Lamarck,1816) 11000 00010 10321 04433 10111 10110 11200 11210 20 Hemimaretia elevata (D¨oderlein,1906) 11011 0111? 00221 12??? 10111 10110 ?2??? 21100 10 Goniomaretia tylota Clark,1917 10110 00021 00221 12122 10101 11110 32223 11210 30 Pseudomaretia alta Agassiz,1863 10010 00021 00221 12222 10101 11110 32222 01110 30 Paramaretia multituberculata Mortensen,1950 11000 01010 10421 13445 00221 10111 32000 11200 00 Breynia australasiae (Leach,1815) 11000 10020 01221 13332 10100 01110 32201 00211 31 Breynia desorii Gray,1851 11000 00020 01321 13332 10100 01110 32200 00211 31 Lovenia cordiformis Agassiz,1872 02200 10021 01321 13222 01111 11110 32211 21221 10 Lovenia elongata Gray,1845 02000 10021 01221 12122 01111 11110 32211 21221 10 Lovenia gregalis Alcock,1893 03100 00021 00221 12222 00111 11110 32212 21211 10 Lovenia subcarinata Gray,1845 02000 10021 00321 13222 00111 11110 32222 21221 10 Lovenia triforis Koehler,1914 02010 01021 00221 12332 00111 11110 32012 21211 10 Pseudolovenia hirsuta Ag.&Clark,1907 02011 01011 00321 13332 ?0111 11110 32212 212?1 10 Echinocardium cordatum (Pennant,1777) 02100 11021 00200 12222 10101 11000 00033 00101 40 Echinocardium mortenseni Thi´ery,1909 11000 01021 00210 12222 10101 11000 00033 00201 10 Homolampas lovenioides Mortensen,1948 11012 0211? 00321 1223? ?0111 11110 32222 00200 10 Homolampas fragilis (Agassiz,1869) 11012 0211? 00221 12222 ?0111 11110 32222 00200 10 Hemipatagus? argutus Clark,1915 02000 02011 0??21 1???? ?0111 10110 11011 002?0 10 Hemipatagus? carolinensis (Kier,1997) 02000 00021 0?221 1???? ?011? ?1111 ?1112 002?0 10 Hemipatagus hoffmanni (Goldfuss,1826) 02000 00021 00221 12122 00211 11110 22212 11210 10 Hemipatagus ocellata (Defrance,1827) 02000 00021 00221 12223 00211 11110 32212 21210 10 Hemipatagus? subrostratus Clark,1915 020?0 020?? ???21 ????? ?0?11 1?110 11012 002?0 ?0
Note: In characters 17–20 there may be some variation in character state between the paired interambulacra, here the most common state was used. 180 A. Kroh
Appendix 3 Annotated list of the 1951 Pseudomaretia alta (A. Agassiz); Mortensen: 58– nominal species of Hemipatagus 62; pl. 3, figs 12–14, 16–20; pl. 46, figs 1–6, 8–11, and Maretia 13, 19, 20. TYPE MATERIAL. Museum of Comparative Zoological at Also included are the species subsequently placed in either Harvard College, Cambridge. genus (based on Lambert & Thiery´ 1909–1925 and Kier & Lawson 1978). Systematic decisions were made on the basis TYPE LOCALITY. Kagoshima Bay, Japan, ∼ 9 m water depth. of published descriptions and illustrations or taken from the literature where modern revisions were available. AGE. Recent. TAXONOMIC PLACEMENT. Type species of Pseudomaretia affinis Duncan & Sladen, 1883 Koehler, 1914. ∗1883 Euspatangus [sic] affinis, Duncan & Sladen; Duncan & Sladen: 46–47, pl. 12, fig. 2. anomala Duncan, 1877 1924 Hemipatagus affinis Duncan et Sladen (Euspatan- ∗1877 Maretia anomala, sp. nov.; Duncan: 53–53; pl. 4, gus [sic]); Lambert & Thiery:´ 457. figs 1–4. 2004 Eupatagus (Eupatagus) affinis (Duncan & Sladen) 1985 Eupatagus anomalus (Duncan, 1877); Kruse & n. comb; Srivastava: 145–146; pl. 6, figs 5–6. Philip: 173–174; figs. 9a–b; pl. 2, fig. 5; pl. 3, figs 1–3; pl. 5, figs 4–5. HOLOTYPE. The specimen figured by Duncan & Sladen HOLOTYPE. Natural History Museum, London 14072, (1883); Geological Survey of India no. C 043 B (GSI Type figured by Duncan (1877: pl. 4, figs 1–4). No. 2867 (Srivastava 2004)). TYPE LOCALITY. Cliffs at the mouth of the Sherbrooke River, TYPE LOCALITY. One mile east of Goir, near Narainsir, Victoria, Australia. Kachchh, Pakistan. TYPE OCCURRENCE. Ruthledge Creek Member. OCCURRENCE. Numulitic Series of Kachchh. AGE. Balcombian, Middle Miocene. AGE. Middle Eocene (Srivastava 2004). TAXONOMIC PLACEMENT. Eupatagus (Kruse & Philip TAXONOMIC PLACEMENT. Eupatagus Duncan & Sladen 1985: 173); based, among other features, on the presence (1883); on the basis of the occurrence of a peripetalous and of a peripetalous fasciole, which had been previously recog- a subanal fasciole. nised by Duncan (1877) and subsequent authors, but still the species remained in the genus Maretia until the revision by affinis Herklots, 1854 Kruse & Philip (1985). ∗1854 Spatangus affinis, nouv. esp.; Herklots: 12; pl. 2, figs 5, 5a. antillarum Cotteau, 1875 1937 Eupatagus (s. Brissoides) affinis (Herklots); ∗1875 Euspatangus [sic] Antillarum, Cotteau, 1875; Jeannet & Martin: 274–275, fig. 51. Cotteau: 43; pl. 7, figs 7–11.
LECTOTYPE. The specimen figured by Herklots (pl. 2, figs SYNTYPES. Collection Cleve, Museum Stockholm, Museum 5, 5a), no. 477 (the same number is mentioned under his S. of Upsala. pulchellus); no. L4292 according to Jeannet & Martin (1937: 274), Rijksmuseum van Geologie en Mineralogie, Leiden. TYPE LOCALITY. St. Barthelemy´ Island.
TYPE LOCALITY. Tjidamar, Java. AGE. Eocene.
AGE. Pliocene. TAXONOMIC PLACEMENT. Eupatagus (Henderson 1975: 33), senior synonym of Maretia clarki Lambert and Maretia TAXONOMIC PLACEMENT. Eupatagus (Henderson 1975: twitchelli Lambert according to Henderson (1975: 33). 33). According to Jeannet & Martin (1937: 274) and Henderson (1975: 33), Herklots’ two syntypes represent two different genera/species. One specimen (no. L4292) belongs aragonensis Cotteau, 1887 to the genus Eupatagus. The other specimen (no. L4291) is ∗1887 Maretia aragonensis Cotteau, 1887; Cotteau: 93– a misidentified Maretia planulata (Lamarck) (Martin 1880: 95; pl. 11, figs 9–12; No. 57. 81; Gerth 1922: 512). From Herklots’ paper alone, however, it is not obvious that there were two specimens and it is un- LECTOTYPE. Specimen figured by Cotteau (1887: pl. 11, clear which of the two specimens mentioned by Jeannet & figs 9–12), collection Maurice Gourdon. Martin (1937) is the figured specimen. PARALECTOTYPES. Not figured, collection Cotteau. alta A. Agassiz, 1864 TYPE LOCALITY. Pobla de Roda, Aragon, Spain. ∗ 1864 Maretia alta A. Ag.; Agassiz: 360. AGE. Eocene. Hemipatagus – a misinterpreted Loveniid 181
TAXONOMIC PLACEMENT. Leiopneustes (Lambert & Thiery´ TYPE LOCALITY. Banda Islands. 1924: 448); periproct high on the posterior face, visible in aboral view, sternal plates fully tuberculate, no naked peri- AGE. ? Pliocene. plastronal areas (providing Cotteau’s illustrations are correct; TAXONOMIC PLACEMENT. Hemipatagus?; all features the plating of the oral side, shown in fig. 11, however, is ob- (cordiform outline, frontal sinus, lack of internal or peri- viously wrong), petals are strongly divergent, not at all like petalous fascioles, shape of the petals, presence of a bilobed anything found in Lovenia, Hemipatagus or Maretia.Onthe subanal fasciole, distribution of the aboral primary tubercles whole the illustrations resemble much more a brissid or some ...) in the original publication support an attribution to Hemi- hemiasterids than a spatangid or loveniid. patagus. archiaci Agassiz, in Agassiz & Desor, 1847 buendensis Ebert, 1889 ∗ 1847 Spatangus Archiaci Agass.; Agassiz & Desor: 8. ∗1889 Spatangus (Maretia) bundensis¨ Ebert n. sp.; Ebert: 69–70; pl. 8, fig. 1a–c. TYPE MATERIAL. Collection d’Archiac. SYNTYPES. Two specimens in the Museum of Gottingen.¨ TYPE LOCALITY. Oulchy-le-Chateaux,ˆ France. TYPE LOCALITY. Doberg, Germany. LITHOSTRATIGRAPHY. Calcaire grossier. AGE. Late Oligocene. AGE. Eocene. TAXONOMIC PLACEMENT. Hemipatagus; very similar to the TAXONOMIC PLACEMENT. Doubtful; according to Cotteau juveniles of H. martensii, as observed previously by Ebert (1885) this species is synonymous with ‘Spatangus’ grignon- (1889); Lambert & Thiery´ (1924: 457) consider H. buenden- ensis Cotteau, 1880. sis to be a junior synonym of H. hoffmanni. argutus Clark, in Clark & Twitchell, 1915 calvimontanus Cotteau, 1885 ∗ 1915 Hemipatagus argutus Clark, n. sp.; Clark & ∗1885 Maretia calvimontana, Cotteau, 1885; Cotteau: 36– Twitchell: 150; pl. 49, figs 1a–d. 38; pl. 5, figs 1–3. 1959 Maretia arguata (Clark); Cooke: 81; pl. 34, figs 1–4. HOLOTYPE. Specimen figured by Cotteau (1885: pl. 5, figs 1993 Maretia arguata (Clark, 1915); Zachos: 148–150; 1–3), collection ‘de la Sorbonne’, Paris. figs 1.1–3. TYPE LOCALITY. Chaumont (Oise), France. HOLOTYPE. USNM 141107; U.S. National Museum, Smith- sonian Institution, Washington, USA (according to Kier LITHOSTRATIGRAPHY. Calcaire grossier inferieur.´ (1980) only a single specimen known, although Cooke (1959) AGE. Middle Eocene. listed three localities). TAXONOMIC PLACEMENT. Hemipatagus; very similar to H. TYPE LOCALITY. Chickasawhay River at Enterprise, Clarke grignonensis, possibly a juvenile or subadult specimen of County, Mississippi, USA. that species; the cordiform outline with distinct frontal sinus, the shape and structure of the petals and the distribution TYPE OCCURRENCE. Winona Formation, Clairborne Group. of the primary tubercles support an attribution to Hemi- AGE. Middle Eocene. patagus; the original description does not reveal details on the nature and tuberculation of the plastron and the shape of OCCURRENCE. Middle Eocene of Mississippi and Texas, the subanal fasciole. USA.
TAXONOMIC PLACEMENT. Hemipatagus? Although this carinata Bolau, 1873 species (like the other New World species of Hemipatagus) ∗1873 [Maretia] carinata n. sp. Bolau: 6. has no camellate tubercles, it is tentatively attributed to Hemi- 1951 Maretia carinata Bolau; Mortensen: 39–41; pl. 1, patagus based on the characteristic cordiform outline, the figs 5–7; pl. 46, figs 14–16. presence of rudimentary pores in the anterior poriferous zones of petals II and IV and the distribution of the aboral SYNTYPES. Museum of Hamburg, Germany (2 specimens in primary tubercles; according to Cooke (1959: 81) and Kier alcohol, Viti Islands); Museum Godeffroy (2 dry specimens, (1980: 50) this species might be a junior synonym of H. Viti Islands; 1 dry specimen, Ellice Islands), Musuem of subrostratus Clark. Gottingen,¨ Germany (1 specimen in alcohol, Bay of Bengal).
TAXONOMIC PLACEMENT. Maretia; the elongate outline, bandaensis Martin, in Jeannet & Martin, 1937 lack of a frontal sinus, long and straight petals and the distri- ∗1937 ? Hemipatagus bandaensis nov. spec. (R. Martin).; bution of the non-camellate aboral primary tubercles support Jeannet & Martin: 276; fig. 53a–c. an attribution to the genus Maretia; the shape of the subanal fasciole is unknown and the adapical pores in the anterior HOLOTYPE. Specimen figured by Jeannet & Martin (1937: poriferous zones of the anterior paired petals are reduced in fig. 53a–c); private coll. Mieg, Basel. size. 182 A. Kroh carolinae Gagel, 1905 clarki Lambert, in Sanchez-Roig, 1924 ∗1905 Chuniola Carolinae gen. nov. sp. n. Gagel: 531– 1924 Maretia clarki Lambert in Sanchez´ Roig; Lambert 537; fig. 1; pl. 24, figs 1a–c, 2a–c, 3, 4a–b, 5. &Thiery:´ 458. [nom. nov. pro specimen B of Eu- patagus floridanus Clark, 1915 (in Clark & Twitchell SYNTYPES. Six whole and several fragmentary specimens 1915: 176–177; pl. 83, figs 1a–c)]. in the collections of the Geological Survey of Preussen, the Museum of Lubeck¨ and the Museum of Hamburg. HOLOTYPE. Specimen figured by Clark & Twitchell (1915: pl. 83, figs 1a–c); Wagner Free Institute of Science. TYPE LOCALITY. Zarrentin, south-east Holstein, Northern Germany. TYPE LOCALITY. Levy County, Florida, USA. AGE. Early Oligocene. AGE. Erratic boulders of Cenozoic age, presumably Mio- cene. TAXONOMIC PLACEMENT. Eupatagus; based on the pres- ence of a peripetalous fasciole; junior synonym of Eupatagus TAXONOMIC PLACEMENT. Type species of Chuniola Gagel, antillarum (Cotteau) according to Henderson (1975: 33). 1905; probably a subjective junior synonym of Hemipatagus Desor, 1858 according to Smith (2004: ‘The Echinoid Direct- ory’); differs from typical Hemipatagus by lacking enlarged cordata Mortensen, 1948 aboral primary tubercles in interambulacra 2 and 3 and by ∗1948 Maretia cordata, new species; Mortensen: 132– the presence of such tubercles in adapical interambulacrum 133. 5; due to the poor preservation of the specimens the pres- 1951 Maretia cordata Mrtsn.; Mortensen: 41–45; pl. 1, ence/absence of internal, subanal and peripetalous fascioles figs 8, 11–16; pl. 44, figs 9–10, 20, 25, 28, 30–31. could not be ascertained (a peripetalous fasciole, however, HOLOTYPE. USNM E7158 (one adult from the ‘Albatross’- seems to be lacking if Gagel’s figures are correct); Gagel ◦ ◦ (1905: pl. 24, fig. 5) illustrated a plastron with broad, short Expedition Stat. 5192 (11 09 N, 123 50 E, 59 m)). labrum very unlike anything observed in Maretia, Hemi- ADDITIONAL MATERIAL. Four specimens from the patagus,orLovenia. ‘Albatross’-expedition (one juvenile from Stat. 5104 (14◦4548 N, 120◦12E, 60 m), 1 adult from Stat. 5257, ◦ ◦ carolinensis Kier, 1997 Mindanao (7 22 N, 124 12 E, 51 m), two adults from Stat. 5431 (10◦3845N, 120◦1245E, 93m). ∗1997 Maretia carolinensis, new species, Kier: 11–13; fig. 6; pl. 9, figs 8–9; pl. 10, figs 1–7; pl. 11, figs TYPE AREA. Philippines. 1–2. AGE. Recent.
HOLOTYPE. USNM 398338, figured by Kier (1997: pl. 10, OCCURRENCE. Philippines; Palawan Islands; Bali Sea; ? figs 1–4), U.S. National Museum, Smithsonian Institution, ‘Siboga’-expedition Stat. 207 (Flores Sea). Washington, USA. TAXONOMIC PLACEMENT. Maretia?; more pronounced PARATYPES. USNM 398337, 398339–398341, 492101– cordate outline than M. planulata with distinct frontal notch; 492124. the subanal fasciole is oval in outline as in M. planulata.
TYPE LOCALITY. Polocksville state quarry, North Carolina, USA. depressus Dubois, 1843 ∗1843 Spatangus depressus; Dubois: pl. 1, fig. 16. TYPE OCCURRENCE. Trent Formation. 1847 Spatangus depressus Dub.; Agassiz & Desor: 8. AGE. Late Oligocene. 1858 Hemipatagus depressus; Desor: 417. 1880 Hemispatangus [sic] depressus (Dubois), Desor.; TAXONOMIC PLACEMENT. Hemipatagus; although this spe- de Loriol: 135–136; pl. 11, fig. 6, 6a–c. cies has no truly camellate tubercles (even though the areoles are distinctly sunken), it clearly belongs to Hemipatagus HOLOTYPE. The specimen figured by Dubois (1843). based on the characteristic cordiform outline, typical shape TYPE AREA. Crimea, Ukraine. of the petals with rudimentary pores in the anterior porifer- ous zones of petal II and IV and the conspicuous field of AGE. ? Eocene. tubercles in adapical ambulacrum III. TAXONOMIC PLACEMENT. Eupatagus?; the illustration in Dubois’ paper shows the aboral surface of a damaged speci- cartagensis Sanchez´ Roig, 1949 men with heterogeneous aboral tuberculation. The tubercles ∗ are confined to the area inside the petalodium and seem to 1949 Hemipatagus cartagensis Sanchez´ Roig: 216–217 have only slightly sunken areoles. This and the shape of the [not seen, fide Kier & Lawson 1978: 121]. petals is strongly reminiscent of Eupatagus and the species AGE. Late Oligocene (Brodermann 1949: 323) or Late Eo- is tentatively referred to that genus. The illustrations in de cene (Sanchez´ Roig 1949). Loriol (1880) are rather poor and do not offer additional insights. Unfortunately, the present author did not have the TAXONOMIC PLACEMENT. Doubtful; not mentioned in opportunity to check the text of Dubois’ work for a descrip- Kier’s (1984) revision of the fossil spatangoids of Cuba. tion of the species. Hemipatagus – a misinterpreted Loveniid 183 desmoulinsi Cotteau, 1863 TAXONOMIC PLACEMENT. Type-species of Hemimaretia ∗ Mortensen, 1950; this genus has only 3 gonopores (pore 1863 Eupatagus Des Moulinsi, Cotteau, 1863; Cotteau: of genital plate 2 lacking) and is thus easily distinguished 148–149; pl. 7, fig. 11. from Hemipatagus, Lovenia and Maretia. 1885 Maretia Des Moulinsi (1863), Cotteau, 1885; Cotteau: 26–28; pl. 2, figs 1–6. elliptica Bolau, 1874 HOLOTYPE. The specimen figured by Cotteau (1863: pl. 7, ∗1874 Maretia elliptica n. sp.; Bolau: 175–177; pl. 6, fig. 11); collection of Cotteau. figs 1–2.
TYPE LOCALITY. La Goureppe, near Biarritz, Basses- HOLOTYPE. Figured by Bolau (1874: pl. 6, figs 1–2), Mu- Pyren´ ees,´ France. seum of Hamburg.
AGE. Late Eocene. TYPE LOCALITY. Malden Island, Pacific Ocean.
TAXONOMIC PLACEMENT. Hemipatagus; initially Cotteau AGE. Recent. (1863: 148) assumed the presence of a peripetalous fasciole, TAXONOMIC PLACEMENT. Insufficiently known; based on but stated that he could not observe it in the specimens. a single specimen; according to Mortensen (1950: 281, 1951: In 1885 he renounced this assumption and transferred the 26) this species probably belongs to either Platybrissus or, species to Maretia; based on the presence of a distinct frontal less likely, to Eurypatagus. sinus, the distribution of the camellate primary tubercles, the structure of the petals, and the presence of a bilobed subanal estenozi Sanchez´ Roig, 1926 fasciole, the species is here attributed to Hemipatagus. 1984 ? Maretia estenozi Sanchez´ Roig; Kier: 106–107; pl. 55, fig. 5, pl. 56, figs 1–2. duncani Lambert, in Lambert & Thiery,´ 1924 ∗ HOLOTYPE. SRC 5491, figured by Sanchez´ Roig (1926: 1924 H. Duncani Lambert; Lambert in Lambert & pl. 37, fig. 1); Sanchez´ Roig collection, Cuban Academy Thiery:´ 457 [nom. nov. pro Eupatagus rostratus of Science. Duncan & Sladen (1884b: 267, pl. 43, fig. 6), non d’Archiac]. TYPE LOCALITY. ‘Cervantes’ farm, San JosedelasLajas,´ 1975 Eupatagus rostratus rostratus d’Archiac; Haban Province, Cuba. Henderson: 44–45. AGE. Early – Middle Miocene. 2004 Eupatagus (Eupatagus) rostratus (d’Archiac) n. comb.; Srivastava: 146–147; pl. 7, fig. 8. TAXONOMIC PLACEMENT. Eupatagus (Kier, 1984: 107), possible synonym of E. cubensis (Cotteau, 1875) according HOLOTYPE. The specimen figured by Duncan & Sladen to Kier (1984: 103–107). (1884b: 267, pl. 43, fig. 6) as Eupatagus rostratus d’Archiac; Geological Survey of India no. G226/76. fasciata Lambert, in Lambert & Thiery,´ 1924 TYPE LOCALITY. West side of Bagothoro Hill, south of ∗1924 M. fasciata Lambert; Lambert in Lambert & Sehwan, India. Thiery:´ 458.
TYPE OCCURRENCE. Nari Series. HOLOTYPE. Unknown.
AGE. Middle Oligocene (Srivastava 2004). TYPE LOCALITY. Philippines.
OCCURRENCE. ?Eocene–Oligocene and, possibly, Miocene AGE. Recent. of India, Japan and Java. TAXONOMIC PLACEMENT. Maretia; juvenile of M. planulata according to Mortensen (1951: 38). TAXONOMIC PLACEMENT. Eupatagus; junior synonym of E. rostratus rostratus d’Archiac, 1850 according to Henderson Pomel, 1887 (1975: 44). ficheuri ∗1887 Hemipatagus Ficheuri; Pomel: 26 [figured on the elevata Doderlein,¨ 1906 unpublished plate A.23 according to Cotteau et al. (1891: 83)]. ∗1906 Maretia elevata nov. spec.; Doderlein:¨ 263–265; pl. 33, figs 5, 5a–b; pl. 48, figs 6a–h. TAXONOMIC PLACEMENT. Nomen dubium; Cotteau et al. 1951 Hemimaretia elevata (Doderlein);¨ Mortensen: 53– (1891: 83) state that this species is unknown to them; never 55; pl. 1, fig. 4; pl. 44, figs 5–7, 21. described or illustrated; on page 79 of the same paper they tentatively refer to it in the synonymy of Maretia tenuis Peron HOLOTYPE. Badly broken specimen in the Museum of & Gauthier, in Cotteau et al., 1891. Berlin (Mortensen 1951). forbesii Woods, 1862 TYPE LOCALITY. Station 251 (Valdivia-Expedition), off Wit- ∗ uland, Eastern Africa (693 m depth). 1862 Spatangus Forbesii; Woods: 75, 83, 105; fig. on p. 75 and 83 [nom. nov. pro Spatangus Hoffmanni AGE. Recent. Sturt 1833 non Goldfuss]. 184 A. Kroh
1864 Hemipatagus Forbesi, Woods & Duncan; Duncan: gauthieri Lambert, in Lambert & Thiery,´ 1924 165, pl. 6, fig. 3. 1877 Lovenia Forbesi, Woods and Duncan; Duncan: 44, 1885 Tuberaster tuberculatus, Peron & Gauthier, 1885; 56–61, pls 4, figs 5–8. Cotteau et al.: 47–49; pl. 3, figs 1–4 [type-species 1994 Lovenia forbesii (Woods, 1862); Irwin & Archbold: of Tuberaster Peron & Gauthier in Cotteau et al., 2–5, figs 2A-K [cum syn.]. 1885]. 1893 Tuberaster tuberculatus, Peron & Gauthier, 1885; LECTOTYPE. Specimen figured by Sturt (1833: pl. 3, fig. 10), Cotteau: 660–662; pl. 360, figs 5–10. designated by Irwin & Archbold (1994: 2); current where- ∗1924 H. Gauthieri Lambert; Lambert in Lambert & abouts unknown. Thiery:´ 457 [nom. nov. pro Tuberaster tuberculatus Peron´ & Gauthier, 1885 non Zittel, 1867]. TYPE LOCALITY. Murray cliffs at Mannum, South Australia. HOLOTYPE. Specimen figured by Cotteau et al. (1885: pl. 3, TYPE OCCURRENCE. Mannum Formation. figs 1–4); collection Gauthier.
AGE. ?Longfordian (Burdiagalian), Early Miocene. TYPE LOCALITY. Kef Iroud, Algeria.
TAXONOMIC PLACEMENT. Lovenia; some references AGE. Oligocene. (Woods 1859, 1862, 1865, 1878; Etheridge 1878; McCoy 1879; Dennant 1890) referring to this species actually TAXONOMIC PLACEMENT. Hemipatagus?; apart from the represent another species (Lovenia bagheerae Irwin in Irwin fully tuberculate sternum (if correctly figured in Cotteau & Archbold, 1994) according to Irwin & Archbold (1994). et al. 1885) this species is very similar to typical Hemi- patagus species (like H. hoffmanni); although Cotteau et al. formosus Zittel, 1867 (1885) debate much about the presence and absence of fasci- oles, none are ascertained due to the heavy abrasion of their ∗1867 Hemipatagus formosus Zitt.; Zittel: 63; pl. 12, material. figs 2a–c. 1894 Eupatagus formosus; Tate: 126 [not seen]. girundicus Lambert, 1915 1975 Eupatagus rostratus zitteli nomen nov.; Henderson: ∗ 45–46; fig. 10; pl. 11, figs 5–6; pl. 12, figs 1–3. 1915 H. girundicus; Lambert: 191. 1924 H. girundicus Lambert; Lambert in Lambert & LECTOTYPE. NHMW 1970/1336, designated by Henderson Thiery:´ 457; pl. XIII, fig. 5. (1975: 45); Natural History Museum Vienna, Austria. 1927b Hemipatagus girundicus Lambert, 1915; Lambert: 119–121. PARALECTOTYPE. NHMW 1959/335/50, 1970/1337. HOLOTYPE. Whereabouts unknown, a cast of the specimen TYPE LOCALITY. Cape Farewell, west Nelson, New Zealand. is figured in Lambert & Thiery´ (1924: pl. 13, fig. 5). AGE. ? Abel Head Fm., Late Oligocene. TYPE AREA. Saucats, Bordelaise, France. OCCURRENCE. Whaingaroan to Altonian (Oligocene to Early Miocene) of New Zealand. AGE. Early Miocene. TAXONOMIC PLACEMENT. Hemipatagus; very similar to H. TAXONOMIC PLACEMENT. Eupatagus; E. formosus (Zittel, ocellatus from the Burdigalian of the Rhoneˆ Basin. 1867) is a secondary homonym of E. formosus de Loriol, 1863 and was renamed E. rostratus zitteli by Henderson (1975: 45–46). gottschei Gagel, 1905 ∗1905 Spatangus ?(Eupatagus ?) Gottschei¨ sp. n.; Gagel: fuchsi Oppenheim, in Blanckenhorn, 1901 539–541; pl. 24, fig. 7; pl. 25, figs 1a–c, 2, 8. ∗ 1901 Maretia Fuchsi Oppenheim; Oppenheim in Blanck- SYNTYPES. 3 natural moulds and one cast in the collections enhorn: 114–117, fig. 9. of the Geological Survey of Preussen, and the Museum of Hamburg. SYNTYPES. 1 specimen from Mirsa Badia, Cyrenaica, Libya (coll. Schweinfurth; figured by Oppenheim in Blancken- TYPE LOCALITY. Zarrentin, south-east Holstein, Northern horn, 1901: fig. 9); 1 fragment from Gebel Geneffe, Eastern Germany. Desert, Egypt (coll. Blanckenhorn); 11 fragments from Wadi Etthal, Sinai, Egypt (coll. Rothpeltz); current whereabouts AGE. Erratic boulders of Cenozoic age, presumably unknown. Miocene.
AGE. Presumably Early or Middle Miocene. TAXONOMIC PLACEMENT. Hemipatagus?; based on the out- line and the distribution of the aboral primary tubercles; type TAXONOMIC PLACEMENT. Spatangus?; classification un- material very poor and in need of redescription. clear; Oppenheim reports that he could not find any traces of an internal or peripetalous fasciole, but records the pres- grignonensis Cotteau, 1880 ence of a subanal fasciole and a fully tuberculate plastron; these features (if correct) suggest that the species is related 1837 Spatangus grignonensis Desmarest, ined. in litt.; to Spatangus. des Moulins: 236–237 [nomen nudum]. Hemipatagus – a misinterpreted Loveniid 185
∗ 1885 Maretia grignonensis (Desmarets) [sic], Cotteau, HOLOTYPE. Specimen figured by Cotteau (1890); Cotteau 1880; Cotteau: 30–36; pl. 3, figs 1–8; pl. 4, 1–4. collection.
TYPE MATERIAL. Unknown. TYPE LOCALITY. Callosa, Alicante, Spain.
TYPE AREA. Paris Basin (Grignon, Cahumont, Fours, AGE. ? Eocene. Chenet ...); France. TAXONOMIC PLACEMENT. Hemipatagus?; the shape of the LITHOSTRATIGRAPHY. Calcaire grossier. test and the distribution and nature of the primary tubercles would indicate an attribution to Hemipatagus,yet,the AGE. Middle Eocene. plastron is shown as being fully tuberculate in pl. 22 (1), fig. TAXONOMIC PLACEMENT. Hemipatagus; Cotteau’s descrip- 3; it is known, however, that Cotteau’s artist often provided tion and illustrations partly contradict each other; assuming highly idealized figures, adding plate sutures or tubercles the description to be correct this species has a partially tuber- where there were none originally. Unfortunately the descrip- culate sternum, a bilobed subanal fasciole, a cordate outline tion gives no further clues. with distinct frontal notch but no rudimentary pores in adap- ical petals II and IV; generic attribution thus is doubtful but hoffmanni Goldfuss, 1829 Hemipatagus seems to fit best; according to Cotteau (1885) ∗ this species is synonymous with Spatangus omali Galeotti, 1829 Spatangus hofmanni [sic] nobis; Goldfuss: 152– 1837 and S. archiaci Agassiz & Desor, 1847. 153; pl. 47, figs 3a–c. TYPE MATERIAL. Figured specimen and seven additional REMARKS. Although frequently cited, Desmarest never pub- lished this species; it is first mentioned in des Moulins (1837) syntypes in the collection of the Goldfuss Museum (Institute referring to an unpublished manuscript by Desmarest; as des of Palaeontology, University of Bonn, Germany), number Moulins provides no description or illustration he cannot be 357a–h. considered the valid author either. TYPE LOCALITY. Doberg, near Bunde,¨ Germany.
AGE. Chattian, Late Oligocene. guebhardi Lambert & Savin, 1902 ∗ AXONOMIC PLACEMENT Type species of Hemipatagus. 1902 Maretia Guebhardi Lambert et Savin nov. sp.; T . Lambert & Savin: 882–889; pl. 59, figs 1–5. 1915 Hemipatagus Guebhardi Lambert & Savin (Mare- hungarica Vadasz,´ 1915 tia), 1905; Lambert: 189. ∗1915 Maretia hungarica n.sp.; Vadasz:´ 237; pl. 9 (3), HOLOTYPE. Specimen figured by Lambert & Savin (1902: figs 17–18. pl. 59, figs 1–5), Savin collection. 1952 Maretia hungarica Vadasz;´ Szor¨ enyi:´ 302, figs a–b.
TYPE LOCALITY. Gattieres,` Alpes-Maritimes, France. HOLOTYPE. Specimen MAFI Ech-225, figured by Vadasz´ (1915: pl. 9 (3), figs 17–18) and Szor¨ enyi´ (1952: figs a–b); AGE. Burdigalian, Early Miocene. housed at the Museum of the Hungarian Geological Survey, Budapest. TAXONOMIC PLACEMENT. Hemipatagus; clearly a member of the genus Hemipatagus with all the typical features de- TYPE LOCALITY. Matraverebely,´ Nograd,´ Hungary. veloped (outline, tuberculation ...); very similar to subadult H. ocellatus. AGE. Late Badenian (Early Serravallian), Middle Miocene. REMARKS. Specimen damaged since the photograph of heberti Cotteau, 1885 Szor¨ enyi´ (1952), petal V now missing. ∗ 1885 Maretia Heberti, Cotteau, 1885; Cotteau: 38–40; TAXONOMIC PLACEMENT. Hemipatagus; Lambert & Thiery´ pl. 5, figs 4–8. (1924: 458) tentatively referred this species to Atelospatan- gus Koch, 1885 because no anterior poriferous zones of petals TYPE MATERIAL. Sorbonne collection. II and IV are visible in Vadsz’s drawing; the correctness of TYPE LOCALITY. La Mortagne, Oise, France. this action was questioned by Mortensen (1951: 79), arguing that the drawing might be incorrect; Mortensen was indeed LITHOSTRATIGRAPHY. Calcaire grossier. correct, as shown by Szor¨ enyi´ (1952: 302), only the adapical AGE. Middle Eocene. pores of the anterior poriferous zones of petals II and IV are rudimentary; no internal fasciole could be observed in the TAXONOMIC PLACEMENT. Referred to Phymapatagus type. Lambert, 1910 (type-species: S. britannus Tournouer) by Lambert & Thiery´ (1924: 463); this is supported by the fully integer Sorignet, 1850 tuberculate plastron and lack of camellate primary tubercles. ∗1850 Spatangus integer; Sorignet: 49 [not seen, fide hispanica Cotteau, 1890 Desor, 1858: 417]. 1858 Hemipatagus integer; Desor: 417. ∗1890 Maretia hispanica, Cotteau, 1889; Cotteau: 5–6; 1885 Maretia integer (Sorignet), Cotteau, 1885; Cotteau: pl. 22 (1), figs 1–3. 40–41. 186 A. Kroh
HOLOTYPE. Unique specimen in the collection Sorignet; TYPE LOCALITY. Sepoloc, northern coast of Madura, N of whereabouts unknown. Java.
TYPE LOCALITY. Fours, Eure, France. AGE. ? Neogene.
LITHOSTRATIGRAPHY. Calcaire grossier. TAXONOMIC PLACEMENT. ? Lovenia (referred to the genus Vasconaster by Lambert & Thiery´ (1924: 466), today con- AGE. Middle Eocene. sidered a junior synonym of Lovenia); generic attribution unclear, type material needs to be re-examined to verify the TAXONOMIC PLACEMENT. Doubtful; published descriptions presence or absence of an internal fasciole and to study the are insufficient. morphological details. koeneni Ebert, 1889 marianii Airaghi, 1903 ∗ 1889 Spatangus (Maretia) Koeneni Ebert, n.sp.; Ebert: ∗ 1903 Maretia Marianii n. sp.; Airaghi: 423–424; fig. 10. 60–62: pl. 8, figs 2a–d. HOLOTYPE. Specimen figured by Airaghi (1903: fig. 10), TYPE MATERIAL. Figured specimen in the collection of the Museo civico Milano, Italy. University of Gottingen;¨ others in the collections of the Uni- versity of Munich, the Museum of Breslau, the Natural His- TYPE LOCALITY. Tirolo, Veneto, Italy. tory Museum of Osnabruck¨ and the Geological Survey in Berlin. AGE. Eocene.
TYPE LOCALITY. Doberg, near Bunde,¨ Germany. TAXONOMIC PLACEMENT. Hemipatagus; typical member of the genus Hemipatagus with the characteristic cordiform out- AGE. Late Oligocene. line with distinct frontal sinus, typical distribution of the aboral primary tubercles (restricted to the anterior part near TAXONOMIC PLACEMENT. Similar to H. hoffmanni, but with the interradial suture in the interambulacra 1 and 4) and fully tuberculate sternal plates and non-camellate primary typical shape of the petals, with rudimentary pores in the tubercles, probably fits best in Spatangus (Phymapatagus) adapical anterior poriferous zones of ambulacra II and IV. ? (based on the description of Ebert; re-examination neces- sary). martensii Ebert, 1889 ∗ lorioli Cotteau, 1878 1889 Spatangus (Maretia) Martensii Ebert n.sp.; Ebert: 67–69; pl. 9, fig. 2a–d. ∗1878 Lovenia Lorioli, Cotteau, 1878; Cotteau: 216–217; pl. 30, figs 2–7; No. 120. HOLOTYPE. Specimen figured by Ebert (1889), Museum of Gottingen,¨ Germany. HOLOTYPE. Specimens figured by Cotteau (1878: pl. 30, figs 2–7); Gauthier collection. TYPE LOCALITY. Doberg, near Bunde,¨ Germany.
TYPE LOCALITY. Drome,ˆ France. AGE. Late Oligocene.
AGE. Burdigalian, Early Miocene. TAXONOMIC PLACEMENT. Hemipatagus; this species exhib- its all the features typical for Hemipatagus and is actually TAXONOMIC PLACEMENT. ? Lovenia; Cotteau’s fig. 7 rather similar to H. hoffmanni; it differs from the latter spe- provides a magnification of the aboral primary tubercles, cies by its lower profile, prominent keel in interambulac- showing distinct crenulations; no extant species of Love- rum 5, more eccentric peristome and higher number of en- nia with crenulated primary tubercles are known (Mortensen larged primaries compared to H. hoffmanni of similar size; 1951); although Cotteau explicitly stated that an internal and Ebert (1889) also saw similarities with H. ocellatus (however, a subanal fasciole are present, Lambert & Savin (1902: 884) part of the illustrations he refers to actually depict Lovenia and Lambert & Thiery´ (1924: 467) state that this species duncani (namely those of Wright 1864)). could be a Hemipatagus, an opinion not shared by any sub- sequent author; lately Philippe (1998: 225) suggested that Michelin, 1863 the type specimen might be a juvenile Maretia ocellata (here mascareignarum Hemipatagus); he argued that he could find no trace of an in- ∗1863 Hemipatagus Mascareignarum N.; Michelin in ternal fasciole in the cast of the holotype studied by him and Maillard: A 6; pl. 16, fig. 2a–i. that the general appearance was much more like H. ocellatus than like a Lovenia. HOLOTYPE. Specimen figured by Michelin (1863: pl. 16, figs 2a–d); whereabouts unknown. madurae Bohm,¨ 1882 TYPE LOCALITY. Reunion´ Island, Indian Ocean. ∗ 1882 Hemipatagus Madurae n. sp.; Bohm:¨ 366–367; AGE. Recent. pl. 2, fig. 3a–c. TAXONOMIC PLACEMENT. Maretia; typical Maretia with HOLOTYPE. Specimen figured by Bohm¨ (1882: pl. 2, long, straight petals, characteristic distribution of the aboral figs 3a–c); whereabouts unknown. primary tubercles (also present in the posterior part of the Hemipatagus – a misinterpreted Loveniid 187 interambulacra 1a and 4b, along the adradial suture), oval, TYPE LOCALITY. Sepocloc, northern coast of Madura, N of outline with only faint frontal notch, low profile, oval sub- Java. anal fasciole, well developed phyllodes and so on. Obviously Michelin was not aware of the genus Maretia, as he states AGE. ? Neogene. ‘Le spatangus planulatus de Lamarck, qui se trouve dans TAXONOMIC PLACEMENT. ? Eupatagus; this species has la mer Rouge, et subfossile sur ses bords, est une espece` been referred to a variety of genera by different authors: Eu- d’hemispatangus, vivant aujourd’hui’ (1853: A 6); according patagus by Gerth (1922: 512), Brissoides by Wanner (1931: to Mortensen (1951: 38–39) this species is a junior synonym 455) and Maretia by Jeannet & Martin (1937: 277); these dif- of M. planulata (Lamarck). ferent opinions obviously result from the poor illustration in Bohm;¨ as Bohm¨ (1882: 365) explicitly mentions the presence mauritanica Pomel, 1885 of a peripetalous fasciole this species is tentatively referred to ∗ Eupatagus here (contrary to Jeannet & Martin (1937: 277), 1885 Sarsella mauritanica; Pomel: 16, pl. 1, figs 6–7; who assume that a peripetalous fasciole is lacking based on pl. 2, figs 5–6 [not seen]. the distribution of the aboral primary tubercles in Bohm’s¨ 1885 Sarsella mauritanica, Pomel, 1885; Cotteau et al.: figure). 36–39; pl. 1, figs 4–8. 1893 Sarsella mauritanica, Pomel, 1885; Cotteau: 667– 670; pl. 362, figs 1–6. nicklesi Cotteau, 1890 ∗ TYPE LOCALITY. Kef Iroud, Dept.´ d’Alger, Algeria. 1890 Maretia Nicklesi, Cotteau, 1889; Cotteau: 6–7; pl. 33 (12), figs 7–10. AGE. Eocene or Oligocene. HOLOTYPE. Specimen figured by Cotteau (1890); Nickles` TAXONOMIC PLACEMENT. Lovenia or Hemipatagus; collection. without re-examining the types or topotypic material it is impossible to decide to which of the two genera the species TYPE LOCALITY. Callosa, Alicante, Spain. belongs; none of the authors that have studied the specimens AGE. ? Eocene. could confirm the presence or absence of a internal fasciole (e.g. Cotteau et al. 1885: 37–38); Contrary to the figures in TAXONOMIC PLACEMENT. Eupatagus; based on labrum Cotteau’s Paleontologie´ Franc¸aise (1893; pl. 362, figs 1–6), shape, fully tuberculate sternum and poorly developed phyl- the plastron is not fully tuberculate (compare description and lodes (Mortensen, 1951: 26). illustration in Cotteau et al. 1885); Cotteau et al. (1885) state explicitly that the aboral primary tubercles are crenulated in nicoleti Agassiz, 1839 this species (which can also be seen on their pl. 1, fig. 5), a ∗ feature which does not occur in extant Lovenia according to 1839 Spatangus Nicoleti Ag.; Agassiz: 33; pl. 4, figs 7–8. Mortensen (1951); however, it seems that many ambulacral plates reach within the subanal fasciole (5 or 6, Cotteau et al. HOLOTYPE. Specimen figured by Agassiz (1839). 1885: pl. 1, fig. 6), a common feature in Lovenia. TYPE LOCALITY. La Chaux-de Fonds, Jura, Switzerland. martini Gerth, 1922 AGE. Burdigalian. ∗1922 Eupatagus martini; Gerth: 511–512; pl. 52, figs 1, TAXONOMIC PLACEMENT. Hemipatagus; a junior synonym 1a. of H. ocellatus according to Agassiz & Desor (1847: 7) and 1975 Eupatagus rostratus rostratus d’Archiac; Philippe (1998: 222). Henderson: 44–45. ocellatus Defrance, 1827 HOLOTYPE. Specimen St.4286 (figured by Gerth 1922); pa- ∗ laeontological collection of the Rijksmuseum van Geologie 1827 Spatangus ocellatus nov. sp.; Defrance in Blainville en Mineralogie, Leiden. et al.: vol. 50, 96–97 [not seen, fide Philippe, 1998: 220]. TYPE LOCALITY. Goenoeng Spolong, Java. 1998 Maretia ocellata (Defrance, 1827); Philippe: 220– 223; pl. 22, figs 6, 7, 8a–b, 9a–b, 10a–c. OCCURRENCE. ? Eocene, Oligocene to Miocene of India, Japan and Java. 2005 Hemipatagus ocellatus (Defrance, 1827); Kroh: 191–194, fig. 94, pl. 81, figs 1–4, pl. 82, figs 1– TAXONOMIC PLACEMENT. Eupatagus; junior synonym of E. 2. rostratus rostratus d’Archiac, 1850 according to Henderson (1975: 44). HOLOTYPE. Whereabouts unknown. TYPE LOCALITY. Saint-Paul-Trois-Chateaux,ˆ Rhoneˆ Basin, mojsvari Bohm,¨ 1882 France. ∗ 1882 Brissomorpha mojsvari n. sp.; Bohm:¨ 364–365; AGE. Burdigalian. pl. 2, figs 1a–d. TAXONOMIC PLACEMENT. Hemipatagus; this well-known HOLOTYPE. Specimen figured by Bohm¨ (1882: pl. 2, species shows all features characteristic of Hemipatagus:a figs 1a–d); whereabouts unknown. cordate outline, camellate primary tubercles on the aboral 188 A. Kroh and oral side, a conspicuous field of tubercles in adapical in- pellati Cotteau 1863 terambulacra 2 and 3, a bilobed subanal fasciole...; Philippe ∗ (1998) recognised the differences to ‘true’ representatives of 1863 Hemipatagus Pellati, Cotteau, 1863; Cotteau: 150. Maretia and expressed his reservations concerning that gen- 1885 Maretia Pellati (1863), Cotteau, 1885; Cotteau: 28– eric attribution. 30; pl. 2, figs 7–11. HOLOTYPE. The specimen described by Cotteau (1863); col- omali Galeotti, 1837 lection of Pellat.
TYPES AND PROVENANCE. Unknown. TYPE LOCALITY. Biarritz, Basses-Pyren´ ees,´ France.
TAXONOMIC PLACEMENT. Doubtful; according to Cotteau TYPE OCCURRENCE. Couche a` Eupatagus ornatus. (1885) this species is synonymous with ‘Spatangus’ grignon- ensis Cotteau, 1880. AGE. Late Eocene.
TAXONOMIC PLACEMENT. Hemipatagus (this paper); based ovata Leske, 1778 on the cordate outline, the presence of a distinct frontal notch, ∗1778 Spatangus ovatus; Leske: 188; pl. 49, figs 12–13 rudimentary pores in the adapical part of the anterior porifer- [not seen; fide Mortensen, 1951]. ous zones of the anterior paired petals, and the presence and distribution of the primary tubercles; single known specimen TYPES AND PROVENANCE. Unknown. is rather small and very similar to juvenile H. desmoulinsi from the same area. TAXONOMIC PLACEMENT. Doubtful; according to Mortensen (1951: 37) the identity of this species is unclear; Leske’s illustrations were based on those from peloria Clark, 1916 Thesaurus III ∗ Seba’s (pl. 15, figs 27–28), which according 1916 Maretia peloria; Clark: 121; pl. 44, figs 1–3 [not to Mortensen are rather poor and of unknown affinity; He seen]. strongly argues against synonymising this species with 1917 Maretia peloria; Clark: 248–249; pl. 146, fig. 25 Maretia planulata as proclaimed by other authors (e.g. Clark [illustration of pedicellaria]. 1925). 1951 Paramaretia peloria (H. L. Clark); Mortensen: 51– 52. pareti Manzoni, 1879 ∗ TYPE LOCALITY. 26 miles south-west of cape Everard, 1879 Maretia Pareti Manzoni; Manzoni: 158–159; pl. 1, Victoria, Australia; ∼ 165 m water depth. figs 1–2; pl. 2, fig. 18; pl. 4, figs 33–39. AGE. Recent. TYPE MATERIAL. Specimens figured by Manzoni (1879). TAXONOMIC PLACEMENT. Paramaretia (Mortensen 1951). TYPE AREA. ‘Colline de Bologna e di Aucona’, Italy.
AGE. ? Miocene. pendulus Agassiz, in Agassiz & Desor, 1847 TAXONOMIC PLACEMENT. Type species of Mazettia ∗1847 Spatangus pendulus Agass.; Agassiz & Desor: 8. Lambert & Thiery´ in Lambert, 1915 (replacement for Manzo- 1880 Hemispatagus [sic] pendulus (Agassiz), Desor; de nia Pomel, 1883 preoccupied by Manzonia Brusina, 1870), Loriol: 133–135; pl. 11, figs 7, 7a–c. regarded as subgenus of Prospatangus Lambert, 1902 by Lambert (1915); current status unclear, but certainly not TYPE MATERIAL. Two poorly preserved syntypes in the Mu- closely related to either Hemipatagus or Maretia (based on seum of Paris (probably not conspecific according to de the fully tuberculate sternal plates, the en-chevron arrange- Loriol 1880). ment of the aboral primary tubercles, the presence of primary tubercles in all aboral interambulacra and the different shape TYPE AREA. Sinai Peninsula, Egypt. (presence of a frontal sinus in combination with an oval, AGE. ? Eocene. elongated outline and long, straight petals)). TAXONOMIC PLACEMENT. ? Eupatagus; the specimen parvituberculata H. L. Clark, 1924 figured by de Loriol (1880) clearly seems to be a Eupatagus based on the shape of the petals, the size and distribution of ∗ 1924 Maretia parvituberculata; Clark: 13–15; pl. 4, the enlarged primary tubercles; the presence of a peripetal- figs 1–5 [not seen]. ous fasciole is not confirmed but seems highly probable; the 1925 Maretia parvituberculata; Clark: 227. identity of the second syntype remains dubious. 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. perornata Schaffer, 1912 ∗1912 Spatangus (Maretia) perornatus Schff.; Schaffer: TYPE LOCALITY. South Africa. 190–191; pl. 59, fig. 4–6. AGE. Recent. SYNTYPES. 2 syntypes (KM F/0127, F/0128), figured in TAXONOMIC PLACEMENT. Eurypatagus (Mortensen 1950: Schaffer (1912: pl. 59, figs 4–6); housed at the Krahuletz- 260). Museum, Eggenburg, Austria. Hemipatagus – a misinterpreted Loveniid 189
TYPE LOCALITY. Eggenburg (Kremserberg), lower Austria. (1937: 277); Herklots (1854: 11) mentioned a second speci- men (no. 452b; pl. 4, figs 5, 5a) which he tentatively referred TYPE OCCURRENCE. Zogelsdorf Formation. to his S. praelongus, but which, in fact is indeterminable. AGE. Late Eggenburgian (Early Burdigalian), Early Mio- cene. pulchellus Herklots, 1854 ∗ TAXONOMIC PLACEMENT. Hemipatagus; junior synonym of 1854 Spatangus pulchellus, nouv. esp.; Herklots: 12; H. ocellatus (Desmarest) (see Kroh 2005). pl. 4, figs 7, 7a–b. 1858 [Hemipatagus] pulchellus; Desor: 418. 1880 Maretia? pulchella Herkl. spec.; Martin: 81. planulatus Lamarck, 1816 1922 Maretia? pulchella (Herkl.); Gerth: 512, figs 9, 9a– ∗1816 Spatangus planulatus; Lamarck: 31. b. 1951 Maretia planulata (Lamarck); Mortensen: 27–39; 1937 Eupatagus (s. Brissoides) pulchellus (Herklots); pl. 4, figs 1–5, 10–11; pl. 5, figs 3–17; pl. 44, Jeannet & Martin: 273–274, figs 50a–b. figs 1–4, 18, 26–27, 29, 32–33. HOLOTYPE. The specimen figured by Herklots (1854: pl. 4, TYPE MATERIAL. Probably at the Museum Histoire Naturelle figs 7, 7a–b), no. 447 (the same number is listed under S. Paris. affinis).
TYPE AREA. ‘les mers australes’. TYPE LOCALITY. Tjidamar, Java.
AGE. Recent. AGE. Pliocene.
OCCURRENCE. South-western Red Sea; Indian Ocean and TAXONOMIC PLACEMENT. Eupatagus; Lambert & Thiery´ western Pacific, from Eastern Africa to Fiji and Gilbert (1924: 451, 458) listed this species both under the pre- Islands, and from Sagami Bay (Japan) to Port Jackson Linnaean genus Brissoides and under Maretia;theoval, (Australia) anteroposteriorly elongated outline, lack of a frontal sinus, shape of the petals, fully tuberculate plastron, restriction of TAXONOMIC PLACEMENT. Type-species of Maretia; Indian the aboral primary tubercles to the area between the petals Ocean populations are usually whitish to yellow, Malayan and the presence of a peripetalous fasciole indicate an attri- specimens are dark coloured, but show no morphological bution to the genus Eupatagus. differences according to Mortensen (1951: 36). regiomontanus Mayer, 1860 planulata abbassi Ali, 1985 ∗1860 Hemispatangus [sic] Regiomontanus Mayer; ∗ 1985 Maretia planulata abbassi n. sp.; Ali: 294–295; Mayer: 11 [pagination of the reprint]. figs 12A–B. TYPE MATERIAL. Current whereabouts unknown. TYPE MATERIAL. USNM 339748. TYPE LOCALITY. Kleinkuhren, north-west of Konigsberg,¨ TYPE LOCALITY. Wadi Abu Abraiki, Western Egypt. Samland, Russia.
AGE. Early Pliocene. AGE. Late Eocene.
TAXONOMIC PLACEMENT. Maretia; very similar to extant TAXONOMIC PLACEMENT. Hemipatagus?; poorly known; M. planulata. a junior synonym of H. sambiensis according to Cotteau (1885). praelongus Herklots, 1854 ∗1854 Spatangus praelongus, nouv. esp.; Herklots: 11–12; rotundus Duncan & Sladen, 1884 pl. 2, fig. 6. ∗1884a Macropneustes roundus, Duncan & Sladen; 1858 [Hemipatagus] praelongus; Desor: 418. Duncan & Sladen: 232–233; pl. 38, figs 6–7. 1880 Maretia planulata Gray; Martin: 81. 1924 Hemipatagus rotundus Duncan & Sladen (Mac- 1924 Atelospatangus praelongus; Lambert & Thiery:´ ropneustes); Lambert & Thiery:´ 457. 458. 2004 Macropneustes (Macropneustes) roundus (Duncan 1937 Maretia planulata (Lamarck); Jeannet & Martin: & Sladen) n. comb.; Srivastava: 147–148; pl. 7, figs 277–278. 12–13.
HOLOTYPE. The specimen figured by Herklots (pl. 2, fig. g), HOLOTYPE. Geological Survey of India no. G302/100 (GSI no. 433. Type No. 2685 (Srivastava 2004)).
TYPE LOCALITY. Tjidamar, Java. TYPE LOCALITY. Teyon´ (or Tiyun),´ east of Chorla, Sind, India. AGE. Pliocene. LITHOSTRATIGRAPHY. Kithar Series. TAXONOMIC PLACEMENT. Maretia; junior synonym of the extant Maretia planulata according to Jeannet & Martin AGE. Middle Eocene. 190 A. Kroh
TAXONOMIC PLACEMENT. Eupatagus or Macropneustes TAXONOMIC PLACEMENT. Doubtful; referred to Leiop- (Macropneustes); Duncan & Sladen (1884a) expressed their neustes by Lambert & Thiery´ (1924: 448); type material problems concerning the generic attribution of this species poorly preserved, large parts of the aboral and oral surface and named both genera mentioned above as potential candi- are missing; referred to Maretia by Cotteau (1893) on the dates. They choose Macropneustes because of the numerous basis of the missing peripetalous fasciole; the features that and prominent development of the aboral primary tubercles can be observed, however, are in favour of Eupatagus (oval, and the inferred absence of the subanal fasciole (specimen elongated outline; very shallow frontal sinus; non-camellate damaged posteriorly). Neither Henderson (1975: 33) nor primary tubercles; arrangement of primary tubercles). Srivastava (2004: 147–148) state why they changed the gen- eric attribution and, apparently, no additional specimens have soubellensis Peron & Gauthier, in Cotteau et al., been found since the discovery of the types. Until more spe- 1891 cimens are described the generic attribution of this species remains tentative. ∗1891 Maretia soubellensis, Peron & Gauthier; Cotteau et al.: 81–83; pl. 1, fig. 3. saccoi Airaghi, 1905 HOLOTYPE. Specimen figured by Cotteau et al. (1891: pl. 1, ∗1905 Maretia Saccoi n.sp.; Airaghi: 53–54; pl. figs 21– fig. 3); collection Peron, current whereabouts unknown. 22. TYPE LOCALITY. Foum-Soubella, south of Setif,´ Dept.´ Con- HOLOTYPE. The specimen figured by Airaghi (1905: pl. stantine, Algeria. fig. 21–22); Museo geologico Torino, Italy. AGE. ? Middle Miocene (‘Langhien’ of Cotteau et al., 1891). TYPE LOCALITY. C. Dogana, Umbria, Italy. TAXONOMIC PLACEMENT. Hemipatagus; Cotteau et al. AGE. Miocene. (1891) provide no information on the presence or absence of an internal fasciole, but mention in their discussion that TAXONOMIC PLACEMENT. Hemipatagus; typical Hemi- this species is similar to H. ocellatus and explicitly state that patagus with the characteristic arrangement of the aboral the latter lacks an internal fasciole, even in perfectly pre- primary tubercles (missing in posterior part of interambulac- served specimens; the cordate shape, the pronounced frontal ral columns 1a and 4b), typical petals with rudimentary pores sinus, the rudimentary pores in the adapical poriferous zone in the anterior poriferous zones of the anterior paired petals, of ambulacra II and IV, as well as the distribution of the cordiform outline and strong frontal sinus; in Airaghi’s de- primary tubercles suggest an attribution to Hemipatagus. scription no fascioles are mentioned, should an internal fas- ciole be discovered in the type material this species would have to be transferred to Lovenia. subrostratus Clark, in Clark & Twitchel, 1915 ∗1915 Hemipatagus subrostratus Clark, n. sp.; Clark & sambiensis Beyrich, 1848 Twitchel: 151; pl. 49, figs 2a–b. 1959 Maretia subrostrata (Clark); Cooke: 81; pl. 34, figs ∗1848 Spatangus Sambiensis n. sp.; Beyrich: 100 5–6. [pagination of the reprint]. 1883 Maretia Sambiensis Beyr. sp.; Noetling: 688. HOLOTYPE. USNM 164652; U.S. National Museum, Smith- 1885 Maretia sambiensis (Beyrich), Noetling, 1883; sonian Institution, Washington, USA. Cotteau: 43. TYPE LOCALITY. Wilmington, North Carolina, USA. TYPE MATERIAL. Current whereabouts unknown. TYPE OCCURRENCE. Castle Hayne Limestone. TYPE LOCALITY. Between Warnicken and Groß-Kuhren, north-west of Konigsberg,¨ Samland, Russia. AGE. Late Middle Eocene. TAXONOMIC PLACEMENT. Hemipatagus?; generic attribu- AGE. Late Eocene. tion uncertain but outline, distribution of primary tubercles TAXONOMIC PLACEMENT. Hemipatagus ?; poorly known; and shape of the petals suggest the attribution to Hemi- H. regiomontanus Mayer, 1860 is a junior synonym of this patagus; according to Cooke (1959: 81) this species is very species according to Cotteau (1885). similar to H. desmoulinsi Cotteau (1885: 26; pl. 2, figs 1–6). savini Cotteau, 1893 tenuis Peron & Gauthier, in Cotteau et al.,1891 ∗ ∗1893 Maretia Savini, Cotteau, 1893; Cotteau: 637–639; 1891 Maretia tenuis, Peron & Gauthier; Cotteau et al.: pl. 354, figs 1–4. 79–81; pl. 1, fig. 2.
TYPE MATERIAL. Two poorly preserved specimens; collec- HOLOTYPE. Specimen figured by Cotteau et al. (1891: pl. 1, tion Savin, current whereabouts unknown. fig. 2); collection Peron, current whereabouts unknown.
TYPE LOCALITY. Vallee´ du Trapel, Aude, France. TYPE LOCALITY. Oued Sebt, near Tizi-Ouzou, Algeria.
AGE. Middle Eocene. AGE. ? Middle Miocene (‘Langhien’ of Cotteau et al., 1891). Hemipatagus – a misinterpreted Loveniid 191
TAXONOMIC PLACEMENT. Hemipatagus; according to TYPE LOCALITY. Kef Iroud, Algeria. Cotteau et al. (1891) no traces of an internal fasciole could be found; the cordate shape, the pronounced frontal sinus, AGE. Oligocene. the rudimentary pores in the adapical poriferous zone of am- TAXONOMIC PLACEMENT. Hemipatagus?; apart from the bulacra II and IV, as well as the distribution of the primary fully tuberculate sternum (if correctly figured in Cotteau tubercles suggest an attribution to Hemipatagus. et al. (1885)) this species is very similar to typical Hemi- patagus species (like H. hoffmanni); although Cotteau et al. tuberculata Agassiz & Clark, 1907 (1885) debate much about the presence and absence of fasci- ∗ oles, none are ascertained due to the heavy abrasion of their 1907 Maretia tuberculata A. Ag. & Clark; Agassiz & material. Clark: 134. 1917 Maretia tuberculata; Clark: 246; pl. 160, figs 5–7. tuberosus Fraas, 1867 HOLOTYPE. Single immature specimen. ∗1867 Eupatagus tuberosus Frs.; Fraas: 279; pl. 6, fig. 8. TYPE LOCALITY. Eastern channel, Korea Strait, west of Shi- 1880 Euspatangus [sic] tuberosus, Fraas; de Loriol: 141– monoseki, Japan; 108 m water depth (59 fathoms). 142; pl. 11, fig. 5.
AGE. Extant. HOLOTYPE. Fragmentary specimen figured by Fraas (1867); Museum Stuttgart, Germany. TAXONOMIC PLACEMENT. Doubtful (juvenile specimen); Mortensen (1951: 45) refers this species to Maretia,butan TYPE LOCALITY. Ouadi-el-Tih, near Caire, Egypt. attribution to Lovenia is also not improbable (the holotype has 7 ambulacral plates reaching into the subanal fasciole AGE. ? Eocene. (2×5 tube feet within the subanal fasciole) and no gono- pores developed at a size of 26 mm; in Maretia planulata TAXONOMIC PLACEMENT. Doubtful; the holotype is a frag- the gonopores open at a size of 18 mm and it has 4 ambulac- ment of the aboral side preserving a petal (I, or more likely ral plates reaching into the subanal fasciole (2×2 tube feet); IV) and the adjoining interambulacra which bear camelate in M. cordata the gonopores open at 13 mm test length; in tubercles; currently impossible to decide whether it is a Love- Lovenia scaber, by contrast, the gonopores open at 30 mm). nia, Hemipatagus or something else. tuberculatus Zittel, 1867 twitchelli Lambert, in Lambert & Thiery,´ 1924 ∗1867 Hemipatagus tuberculatus Zitt.; Zittel: 63–64; pl. 1924 Maretia twitchelli Lambert in Sanchez Roig; 12, figs 1a–c. Lambert & Thiery:´ 458 [nom. nov. pro specimen 1894 Lovenia tuberculata; Tate: 126 [not seen]. CofEupatagus floridanus Clark, 1915 (in Clark & 1975 Lovenia tuberculata (Zittel); Henderson: 31–32; Twitchell 1915: 176–177; pl. 83, figs 2a–d)]. pl. 4, figs 5–6; pl. 5, figs 3–6. HOLOTYPE. Specimen figured by Clark & Twitchell (1915: LECTOTYPE. NHMW 1959/335/51, figured by Zittel (1867: pl. 83, figs 2a–d); Museum of Comparative Zoology, Har- pl. 12, figs 1a–c), designated by Henderson (1975: 31); Nat- vard, USA. ural History Museum Vienna, Austria. TYPE LOCALITY. Levy County, Florida, USA. PARALECTOTYPE. NHMW 1970/1338. TYPE OCCURRENCE. Vicksburg Group. TYPE LOCALITY. Cape Farewell, west Nelson, New Zealand. AGE. Early Oligocene. TYPE OCCURRENCE. ? Abel Head Formation. TAXONOMIC PLACEMENT. Eupatagus; junior synonym of AGE. Late Oligocene. Eupatagus antillarum (Cotteau) according to Henderson (1975: 33). OCCURRENCE. Whaingaroan to Altonian (Oligocene to Early Miocene) of New Zealand. variegata Gray, 1866 TAXONOMIC PLACEMENT. Lovenia. ∗1866 Spatangus (Maretia) variegatus, n.sp.; Gray: 170– 171. tuberculatus Peron & Gauthier, in Cotteau et al., 1885 TYPE MATERIAL. Two specimens in the Natural History Mu- seum, London. ∗1885 Tuberaster tuberculatus, Peron & Gauthier, 1885; Cotteau et al.: 47–49; pl. 3, figs 1–4 [= H. gauthieri TYPE LOCALITY. Pulo Taya, China Sea. Lambert in Lambert & Thiery,´ 1924: 457 (nom. nov. pro Tuberaster tuberculatus Peron´ & Gauthier, in AGE. Recent. Cotteau et al., 1885 non Zittel, 1867); see above]. TAXONOMIC PLACEMENT. Maretia; according to Mortensen HOLOTYPE. Specimen figured by Cotteau et al. (1885: pl. 3, (1951: 36) M. variegata is a dark coloured variant of M. figs 1–4); collection Gauthier. planulata and synonymous with that species. 192 A. Kroh vicentina Dames, 1878 TYPE LOCALITY. Beaumaris, east shore of Port Phillip Bay, Victoria, Australia. ∗1878 Breynia vicentina nov. sp.; Dames: 75–76; pl. 7, figs 7a–b. TYPE OCCURRENCE. Black Rock Sandstone, Brighton Group. HOLOTYPE. Specimen figured by Dames (1878: pl. 7, figs 7a–b); current whereabouts unknown. AGE. Cheltenhamian (Messinian), Late Miocene.
TYPE LOCALITY. Longio, Vicentin, Northern Italy. OCCURRENCE. Early (?) Miocene – Pliocene (Zanclean) of the Port Phillip Basin, Victoria, Australia; possibly also in AGE. ? Eocene. Tasmania. TAXONOMIC PLACEMENT. ? Lovenia; Dames could not ob- serve any fascioles on the single specimen available, but TAXONOMIC PLACEMENT. Lovenia. inferred the presence of an internal fasciole on the basis of the absence of any pore pairs in the adapical ambulacra; zeisei Gagel, 1905 the figures provided by Dames do not fully fit his descrip- ∗ tion (the whole adapical area seems to be missing according 1905 Maretia Zeisei¨ spec. nov.; Gagel: 541–543; pl. 24, to the figure, whereas he mentions details of the adapical area fig. 6; pl. 25, figs 3a–c, 4, 5. in the description; likewise he reports a naked plastron, while SYNTYPES. Four poorly preserved specimens in the collec- the figures show that the plastron is largely missing); it is un- tions of the Geological Survey of Preussen, and the Museum clear why Lambert & Thiery´ (1924: 466) refer this species of Lubeck,¨ Germany. to Hemipatagus; the available information rather suggests an attribution to Lovenia, providing Dames’ description is TYPE LOCALITY. Zarrentin, south-east Holstein, Northern correct. Germany.
AGE. Erratic boulders of Cenozoic age, presumably Mio- woodsii Etheridge, 1875 cene. ∗1875 Hemipatagus Woodsii, sp. nov.; Etheridge: 445– 447, pl. 21, figs 1–7. 1877 Lovenia Forbesi, var. Woodsi, Etheridge; Duncan: TAXONOMIC PLACEMENT. ? Hemipatagus; based on the 44, 56–61. outline and the distribution of the aboral primary tubercles; 1994 Lovenia woodsii (Etheridge, 1875); Irwin & Arch- type material very poor and in need of re-description; junior bold: 6–10, figs 4A–C, 6A–D, 7A–M [cum syn]. synonym of H. gottschei according to Lambert & Thiery´ (1924: 457); or juveniles of H. hoffmanni as discussed by LECTOTYPE. Specimen F17500, figured by Etheridge (1875: Gagel (1905: 543; he rejected this hypothesis because of the pl. 21, figs 1–7), designated by Irwin & Archbold (1994: 6, naked plastron of H. zeisei, which could, however, also be fig. 6A–D); Australian Musuem, Sydney. related to the poor preservation).