bordering the anterior paired petals. Their number ranges from Discussion: 4 to 60 in interambulacra 1 and 4 and 5 to 35 in interambulacra The generic assignment of this species was subject to consider- 2 and 3. The surface between and around these enlarged pri- able debate: originally referred to the genus Spatangus (e.g. mary tubercles is densely crowded with miliary tubercles. Sec- DEFRANCE, 1827; DE LORIOL, 1876) it was later placed into the ondary tubercles with distinct areoles are scattered among genus Maretia (COTTEAU et al., 1891), only to be transferred to them, but are generally rather rare on the apical surface of the the genus Hemipatagus by LAMBERT (1909) soon afterwards. test. Only in interambulacrum 5 along the interradial suture During this time the question arose, whether Maretia and and along the adradial sutures to ambulacrum III in interambu- Hemipatagus should be considered synonymous (AGASSIZ, lacra 2 and 3 a larger numbers of secondary tubercles is pres- 1872: 568, 1873; COTTEAU, 1885-89: 24; FOURTAU, 1920: 83) or ent. not (e.g. DUNCAN, 1889: 222, 252; LAMBERT, 1909: 107, 1915a: In the interambulacral columns 2b and 3a a conspicuous field 188, 1927b: 87), or even if Hemipatagus is synonymous with of coarse tubercles is developed adjacent to the adradial suture Lovenia (DUNCAN, 1877: 56). In his “Monograph of the Echi- to the adapical part of ambulacrum III (see Pl. 81, Fig. 2b) noidea” MORTENSEN (1951: 23-27) lengthily discussed this mat- On the oral surface interambulacra 1, 2, 3 and 4 are covered ter and finally considered Hemipatagus DESOR,1858 a junior densely with camellate primary tubercles arranged in distinct synonym of Maretia GRAY, 1855. In the “Treatise on Inverte- rows. They have deeply sunken, asymmetric areoles and heli- brate Paleontology” FISCHER (1966: U609) adopted MORTENSEN’s cally spiralled parapets. These tubercles are largest next to the view and thus it was soon widely accepted in the scientific adradial sutures and decrease in size from the centre towards community, although some authors (e.g. PHILIPPE, 1998: 222) the margin. The plastron is ultramphisternous with a narrow expressed their doubts. contact between labrum and sternum (Fig. 94). The labrum is Based on material of the type-species of the genera in ques- triangular and projects slightly into the peristome. Up to two tion the present author (KROH, in prep) was able to show that third of the sternal plates are naked, only in the posterior part Hemipatagus is clearly different from Maretia and in fact be- secondary tubercles are found, which are arranged in a fan- longs to the Loveniidae, not the Spatangidae as the latter ge- shaped pattern. nus. The features with which the two genera can be separated Peristome: The peristome is kidney shaped, transversely elon- are outlined above under the remarks to the genus. gated and lies about 25-28 % TL from the anterior margin. It The studied specimens clearly belong into the genus is moderately large, being about 15 to 16 % TL wide in adult Hemipatagus, and are conspecific with H. ocellatus (DEFRANCE, specimens. 1827) recently re-described and illustrated by PHILIPPE (1998: Periproct: The periproct lies inframarginally, high on the verti- 220-223; pl. 22, figs. 6, 7, 8a-b, 9a-b, 10a-c; under the name cally truncated posterior end of the test. It is oval, transversely Maretia ocellata). elongated and is with approximately 10 % TL distinctly smaller Based on the examination of the type material housed at the than the peristome. There is a depressed subanal area, which is Krahuletz-Museum (Eggenburg, NÖ, Austria) the species Spa- free of primary and secondary tubercles. tangus (Maretia) perornatus SCHAFFER, 1912a has to be placed Fascioles: The only fasciole present is a wide bilobed subanal into the synonymy of H. ocellatus. According to AGASSIZ & fasciole. The fasciole band is moderately wide and corresponds DESOR (1847b: 7) and PHILIPPE (1998: 222) Spatangus nicoleti to the othofasciole type of NÉRAUDEAU et al. (1998b). The exis- AGASSIZ, 1839 is also a junior synonym of H. ocellatus. Al- tence of an internal fasciole in adult specimens can be exclud- though, surprisingly similar and originally determined at first as ed. Even in specimens with extremely well preserved aboral Spatangus hoffmanni (WRIGHT, 1855: 176) and later as S. ocel- tuberculation no traces of fasciole bands were located around latus (WRIGHT, 1864: 487), the Maltese species Lovenia dun- the apical disc. cani (GREGORY, 1891), is clearly a loveniid, based on the pres- ence of an internal fasciole as shown by CHALLIS (1980). Differential diagnosis: The material of VADÁSZ (1914, 1915) attributed to Hemipata- The closely related Hemipatagus girundicus LAMBERT, 1915a gus ocellatus is rather poorly preserved. The figured specimen from the Burdigalian to Langhian of Bordeaux differs from this could not be located at the MAFI. The only specimens that are species by a broader keel in interambulacrum 5, which is more preserved there are three weathered fragments with camellate flattened and more densely tuberculate than in H. ocellatus aboral tubercles (MAFI Ech 237) that are not identifiable at (compare the descriptions in LAMBERT, 1915a: 191; LAMBERT & genus level. THIÉRY, 1909-25: 457, pl. 13, fig. 5; LAMBERT, 1927b: 119-121). If this sole feature is enough to keep these species separated is Occurrence: still a matter of debate. CHAVANON (1974: 271-272) and PHILIPPE Austria: Early to Late Eggenburgian (Early Burdigalian) (1998: 222-223) did not synonymise them, although the latter Molasse Zone: Achberg (Scutellensande, Loibersdorf expressed his doubts and stated that more material is needed Fm.), near Maria Dreieichen, NÖ (STEININGER, 1971a, b); to solve this question. Eggenburg (Brunnstube), NÖ (STEININGER, 1971a, c); Eggen- Hemipatagus hoffmanni (GOLDFUSS, 1829) differs by its burg (Kremserberg, Zogelsdorf Fm.), NÖ (SCHAFFER, 1912a; higher test (test height ranging from 42 to 50 % TL), the [KM]); Fels am Wagram (Fels Fm.), NÖ (STEININGER, 1971a); bluntly pointed, obliquely truncated posterior end, the laterally Gösing (Fels Fm.), NÖ ([KM, NHMW]); Grübern (Zogels- flexed anterior petals, the lower number of camellate primary dorf Fm.), NÖ (SCHAFFER, 1912a; [NHMW]); Unternalb tubercles, which are, moreover, never as large and closely (Retz Fm.), NÖ (HARZHAUSER & KROH, 1999; KROH & HAR- spaced as in H. ocellatus [based on the description in GOLDFUSS, ZHAUSER, 1999; LUKENEDER & HARZHAUSER, 2002 [NHMW]) 1826-44: 152-153, pl. 47, figs. 3a-c, the type-material in the collection of the Goldfuss Museum (Univ. Bonn) and speci- Paratethys (non-Austrian occurrences): Burdigalian, ? Badenian mens from the type locality in the collection of the NHMW and (Langhian-Early Serravallian) the NKM Berlin]. Swiss Molasse: La Chaux-de-Fonds, Jura (AGASSIZ, 1839; Spatangus ornatus (CUV.) (as described in GOLDFUSS, 1829: DESOR, 1858; MANZONI, 1873; DE LORIOL, 1876; LAMBERT, 152; pl. 47, figs. 2a-c), which was the name-inspiring species 1915; LAMBERT & JEANNET, 1928); Les Verrières, Neuchâtel for SCHAFFER’s S. perornata, is an Eupatagus. Its plastron is (DE LORIOL, 1876; LAMBERT, 1915a); Neuchâtel (AGASSIZ & fully covered with tubercles and a peripetalous fasciole is DESOR, 1847b; D’ORBIGNY, 1852; PICTET, 1857) present according to AGASSIZ & DESOR (1847b: 9). Moreover, Great Hungarian Basin (Pannonian Basin): ? Mátra- albeit generally similar it has no camellate primary tubercles verebély, Nógrád, Hungary (VADÁSZ, 1914, 1915) and cannot easily be confused with the species discussed Transylvanian Basin: ? Gârbova de Sus (= Felsö-Orbó), here. Romania (VADÁSZ, 1914, 1915) 192 SYSTEMATIC PART SYSTEMATIC PART 193 Mediterranean: Burdigalian, ? Langhian Egerian (Chattian-Aquitanian) – Pucking or Ebelsberg or Western Mediterranean: Italy, mainland (AGASSIZ, Weikerlsee or Asten (Ebelsberg Fm.), OÖ, Austria 1840b; PICTET, 1846): ? Monteorsello (MAZZETTI & PANTA- NHMW: 19 spine aggregates (NHMW 2003z0026/1375- NELLI, 1883), ? Monte Titano (MANZONI, 1873), ? Rocca S. 1381, 2003z0026/1383-1393) Maria (MAZZETTI & PANTANELLI, 1883), ? Sasso Bolognese Egerian (Chattian-Aquitanian) – Weikerlsee (Ebelsberg Fm.), (MANZONI, 1873); Rhône Basin, France (PICTET, 1846, 1857): Linz, OÖ, Austria Beaucaire (PHILIPPE, 1998), Bonnieux (PHILIPPE, 1998), Cap NHMW: 2 slabs with accumulated spatangoid spines Couronne, near Martigues, Bouches-du-Rhône (NEGRETTI, (NHMW 2003z0026/1229-1230) 1984; PHILIPPE et al., 1990), Carry, Bouches-du-Rhône Late Eggenburgian (Early Burdigalian) – Eggenburg, NÖ, (GOURRET, 1892; LAMBERT, 1915a), Châteauneuf-Miravail Austria (PHILIPPE, 1998), Clansayes (LAMBERT, 1915a; PHILIPPE, 1998), NHMW: 1 spine fragment (NHMW 2004z0110/0013) Couremes, near Vence, Alpes-Maritimes (LAMBERT, 1915a), Late Eggenburgian (Early Burdigalian) – Unternalb (Retz Fm.), Crest (PHILIPPE, 1998), Grignan (PHILIPPE, 1998), Istres SE Retz, NÖ, Austria (PHILIPPE, 1998), Lacoste (PHILIPPE, 1998), Martigues, Bouch- NHMW: numerous spine fragments (NHMW 1999z0051/ es-du-Rhône (PHILIPPE, 1998), Ménerbes (PHILIPPE, 1998), 0026) Montségur-sur-Lauzon (PHILIPPE, 1998), E of Port des Tam- Ottnangian (Late Burdigalian) – Höbmannsbach, Taufkirchen aris, Bouches-du-Rhône (NEGRETTI, 1984), Reillanne, Basse- Bay, ESE Schärding, OÖ, Austria Alpes (LAMBERT, 1915a; PHILIPPE, 1998), Saint-Mitre-les- NHMW: 8 spine fragments (NHMW 1978/1966/12i) Remparts (PHILIPPE, 1998), Saint-Paul-Trois-Châteaux, Karpatian (Late Burdigalian) – Teiritzberg, near Stetten, Drôme (AGASSIZ & DESOR, 1847b;