BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE, SCIENCES DE LA TERRE, 57: 201-215, 1987 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN, AARDWETENSCHAPPEN, 57: 201-215, 1987
The genus Typocidaris (Cidaroida; Echinoidea) in the Upper Cretaceous of the Maastricht area (Belgium and the Netherlands)
by Joris F. GEYS
Abstract Unfortunately, isolated skeletal remains of Cidaroida are rarely identifiable. In the past, this has often Compilation of earlier work shows that 19 names have been used for resulted in a profusion of "species", many of which Cidaroida from the Belgian and Dutch Campanian and Maastrichtian. cannot be accepted. In earlier literature, I recorded Most of these have to be rejected or are doubtful, because of the no less than 19 names for Cidaroida from the Maas¬ poor state of préservation of the specimens involved. Typocidaris is trichtian of the area. reintroduced as a separate genus. Four species are described and Maastricht Three of these are discussed; T. ubaghsi is new. characteristic for Danian, rather than Maastrichtian Key-words: Cidaroida, Echinoidea, Maastrichtian, Cretaceous, Bel¬ strata: a frequent error of earlier authors, due to insuf¬ gium, The Netherlands. ficiënt stratigraphical knowledge at that time. Nine taxa have to be rejected, for reasons explained herein. The presence of another four could not be confirmed. Résumé Two species, one of them new, had to be added to the list. This means that the original 19 species are reduced to a mere Pour la description des échinides cidaroides du Campanien et du five, four of them belonging to the Maastrichtien belge et néerlandais, 19 dénominations ont été utilisées genus Typocidaris. A survey of former and present dans le passé. La plupart de celles-ci doivent être rejetées ou ont opinions on the composition of the cidaroid faunas of trait à des espèces insuffisamment connues, ayant été intoduites pour the Upper Cretaceous in the Maastricht area, is given des spécimens de conservation médiocre. Typocidaris est réintroduit in table 1. comme genre indépendant. Quatre espèces sont décrites, dont une, T. ubaghsi, est nouvelle. Mots-clefs: Cidaroida, Echinoidea, Maastrichtien, Crétacé, Belgique, Pays-Bas. Abbreviations used in the text
D : diameter of the corona, at the ambitus Introduction h : height of the corona ds : diameter of the apical System dp : diameter of the peristome Fossil remains of Cidaroid Echinoids are by no means rare in Upper Cretaceous deposits. In the Low Coun- Lg : province of Liège NLb province of Dutch Limburg tries, as well as elsewhere, they are frequently found BLb : province of Belgian Limburg as isolated and interambulacral plates. Many hundreds NL : the Netherlands of such fossils are present in almost every important B : Belgium collection, public or private. On the other hand, com¬ DDR : German Democratie Republic plete coronas of Cidaroida are exceedingly rare. Even DK : Denmark D : Fédéral a few plates, remaining in anatomical connection, are Republic of Germany E : Spain an exceptional, and thus valuable fossil. This pheno- F : France menon is partly due to the life habits of Cidaroida, GB United Kingdom of Great Britain and Northern Ireland browsing on hard substrates and hence living in erosio- PL : People's Republic of Poland nal environments (KlER, 1977). The structural SU : Union of Socialist Soviet Republics weakness of the cidaroid test does not favor its préser¬ KBIN : vation either. The stereomes of adjacent plates hardly Royal Belgian Institute for Natural Sciences, Brussels, B. NHMM : Natural History Museum, Maastricht, NL. possess any interlocking processes, so that the plates fall easily apart, after the animal's death and after In the synonymy lists, the conventional signs of A.V. Dhondt (1972) decay of the Connecting tissues (Smith, 1984). have been adopted. 202 Joris F. GEYS
Table 1 Cidaroida from the Upper Cretaceous in the Maastricht area, recorded by different authors.
Geys herein
1857Bosquet, 1874cotteau, 1897Lambert, 1881Mourlon, 1911Lambert, 1935Smiser, 1965Meijer, Gu Ma Da
- Cidaris arenata x x - baylei x ciplyensis x x x — — danica x x x x faujasi x x x x x x — — filamentosa x — — forchhammeri x x x x hardouini x x x x x x lingualis x x o o ornatissima x o o
— perornata x — pistillum x x x pseudohirudo x x — — regalis x x o o sceptrifera x x x schlueteri x x — — serrata x x x x x sorigneti x o o subvesiculosa x x x — — ubaghsi x venulosoides x x — —
x: recorded; — : doubtful or rejected; O: unconfirmed. Gu: Gulpen Chalk; Ma: Maastricht Chalk; Da: Danian.
Typocidaris: independent genus orjunior synonym of Stereocidaris? 05 o E S oî m o 05 a genus, ever JJ in Typocidaris has been well established _c c 0) o o C en — o since it has been erected by Pomel Animais eu en (1883). 05 en >N 'c en CD
In this context, Stereocidaris, Temnocidaris and Typo¬ Gulpen Formation, Maastrichtian: 1 impression in cidaris should be considered as separate genera. If flint (coll. L. Indeherberghe). Brunssumer Heide ancestors of Stereocidaris should belong to Typocida¬ (NLb, NL); reworked from Gulpen Formation, Maas¬ ris, the former genus should have to be considered a trichtian: 1 impression in flint (coll. NHMM, n° Boers- subgenus of the latter. This view has already been ma 503). supported by Lambert & Thierry (1910). Our present state of knowledge is insufficiënt however, to Dimensions: permit a final statement. Both flint impressions are fragments. The dimensions Phalacrocidaris does not belong to the Stereocidari- of the J. Jagt specimen are given below: D = 27 mm; nae: it misses the characteristic suturai grooves. In my h = 12 mm; ds = 10 mm; dp = 7 mm; d/D-ratio = opinion, its similarity to Stereocidaris is due to conver¬ 0.44. ds/D-ratio = 0.37; dp/D-ratio = 0.26. gence, rather than to relationship. Description : Moderately small Typocidaris, with a circular peri- Systematic descriptions stome, which is not sunken. Gill slits are absent. Ambulacra are sinuous and wide. Poriferous zones Class Echinoidea Leske, 1778 are slightly depressed. Pores are not conjugate; neural Subclass Perischoechinoidea M'Coy, 1854 grooves are present. The pores of each pair are sepa- Order CIDAROIDA Claus, 1880 rated by a tiny granule, which shows a small, trans- Family CIDARIDAE Gray, 1825 verse groove. The pore pairs are very slightly oblique. Subfamily STEREOCIDARINAE lambert, 1900 Interporiferous areas are fairly wide and densely gra- Genus Typocidaris pomel, 1883 nulated. Vertical, regular series of granules, one on every ambulacral plate, adjoin the poriferous zones. Type-species: Perradially, between these series of larger granules, Cidaris malum Gras, 1848 (original désignation). the remaining space is covered by a very dense, very fine granulation. These small, perradial granules are Typocidaris pistillum (quenstedt, 1852) arranged in horizontal rows : two rows of three to four (Pl. 1, Figs. 1-6) granules on each plate. The granules diminish in size towards the * perradial suture. . 1852 Cidaris pistillum, quenstedt, p. 578, pl. 49, At the ambitus, 16 ambulacral fig. 20. plates correspond in
. 1910 Cyathocidaris pistillum, Lambert & Thierry, height to one interambulacral plate. There are 5 inter- ambulacral p. 146. plates in a series. Primary tubercles are
. 1928 Cidaris Hagenowi, Ravn, p. 17, pl. 2, fig. 8. perforate and non crenulate, except on the adapical
. 1928 Cidaris Bolli, Ravn, p. 15, pl. 1, figs. 8, 20. side of tubercles above the ambitus. The areoles are 1939 cf. Cidaris Bolli, kongiel, p. 6, pl. 1, figs. 5-6. slightly conical and moderately sunken. Their distal
. 1972 Stereocidaris pistillum, Nestler, pp. 172-180, border has a steep slope, but is neither vertical, nor pl. 1-3. undercut. Scrobicules are surrounded by a full ring of . 1973 Stereocidaris pistillum, kutscher, pp. 108-109, small scrobicular tubercles, which are accompanied figs. 1, 2, 6. by a crescent-shaped élévation on the distal side. At . 1975 Stereocidaris pistillum, nestler, p. 82, fig. 131. the ambitus, a scrobicular (for full synonymy, cf. Nestler, 1972). ring consists of 15 tubercles. The uppermost scrobicule of each interambulacrum is Locus typicus: rudimentary. Extrascrobicular surfaces are covered by Rügen, German Democratie Republic. a very dense, fine granulation. These granulated sur¬ faces completely surround the scrobicules, but are best Stratum typicum: developed interradially. Adradially, a mere two or Schreibkreide, Lower Maastrichtian. three granules separate the scrobicular rings from the sutures. Interradially, the granules are arranged in Occurrences outside the Maastricht-area: irregular series radiating from the scrobicules. In our DDR: Lower Maastrichtian of Rügen (Nestler, specimens, there are no clearly visible furrows 1972); DK: Maastrichtian of Mpn, Sjaelland and Jyl- between these series. Horizontal and interradial sutu¬ land (Ravn, 1928), as Cidaris hagenowi and Cidaris res are depressed. Small, deep dépressions are present bolli\ PL: Maastrichtian of the Lublin-area (Kongiel, on horizontal adapical sutures. 1939), as Cidaris bolli. Diagnostic features: Specimens studied: 1. Interambulacral scrobicules widely separated. Hallembaye (C.P.L.-quarry) (Lg, B); 4.5 m over Loën 2. 5 or 6 interambulacral plates in a series. Horizon, Zevenwegen Beds, Gulpen Formation, 3. Uppermost interambulacral plate reduced. Upper Campanian: 1 specimen (coll. J. Jagt, n° 1180). 4. Sutures sunken, with deep dépressions on adapical Hallembaye (C.P.L.-quarry) (Lg, B); Lanaye Beds, horizontal sutures. 204 Joris F. GEYS
5. Ambulacral interporiferous areas well developed, . 1882 Cidaris sceptrifera, wright, pp. 54-57, pl. 5, figs. with two regular marginal vertical series of granu¬ 16-17; pl. 6, figs. 2-6; pl. 7, figs. 1-2; pl. 7a, figs. les; perradial granules smaller and arranged in irre- 1, 3. 1883 Cidaris sceptrifera, cotteau, -. 11. gular horizontal series, two on each plate. 1892 Stereocidaris sceptrifera, schlüter, pp. 110-118, 6. Interporous granules with a transverse groove. pl. 14, figs. 6-7; pl. 16, figs. 5-6. 7. Scrobicular tubercles distally accompanied by a . 1910 Stereocidaris sceptrifera, lambert & thierry, crescent shaped élévation. p. 152. 1911 Stereocidaris sceptrifera, lambert, p. 60, pl. 2, Discussion : fig. 31. The typical configuration of the interporiferous granu¬ 1925 Stereocidaris sceptrifera, lambert, p. 24. lation, as it occurs in T. pistillum, is known in very 1928 Stereocidaris sceptrifera, lambert & jeannet, few other species. This makes them easy to recognise p. 116. 1928 Cidaris and hard to confuse with other Cidaroida. sceptrifera, Ravn, p. 25. v 1935 Stereocidaris sceptrifera, Smiser, p. 24, pl. 2, fig. 2. Différences between the coronas of T. pistillum and 1939 Stereocidaris sceptrifera, Kongiel, p. 14, pl. 2, T. hagenowi (Desor, 1858) (Nestler, 1972, Pl. 4-5; fig. 3. Maastrichtian of occurs Rügen, D.D.R.), which in . 1968 Stereocidaris sceptrifera, Hynda, pp. 196-197, the same strata, are small and subtle. According to pl. 40, figs. 5-7. nestler (1972), the main différences are: a/ the lack 1970 Stereocidaris sceptrifera, blaszkiewicz, p. 159. of a transverse groove on the interporous granuliform . 1974 Stereocidaris sceptrifera, savchinskaya, pp. 309- partition in T. hagenowi, b/ the absence of crescent- 310, pl. 94, fig. 7. like élévations near the scrobicular tubercles in T. 1979 Typocidaris sceptrifera, fournier, p. 9. hagenowi, d the periphery of the areoles, which is . 1980 Stereocidaris sceptrifera, fischer, p. 266, pl. 131, fig. 3. vertical to undercut in T. hagenowi, but less steep in non 1882 Cidaris sceptrifera, wright, pl. 6, figs. la-lh. T. pistillum, and d/ the areoles of T. hagenowi, which non 1928 Stereocidaris sceptrifera, mortensen, pp. 226-227. are so over deep that only the mamelon thrusts out non 1966 Stereocidaris sceptrifera, Fell, p. U327, figs. 242- the rim, while the entire upper part of the boss does 3a, b. so in T. pistillum. Ambulacral plates of T. danica Ravn, 1928 (Pl. 2; Locus typicus: Danian of Denmark) are similar to those of T. pistil¬ Lewes, Sussex, England. lum and of T. hagenowi. They also show one marginal tubercle and two horizontal rows of smaller, perradial Stratum typicum: tubercles. Yet, T. danica has more interambulacral "Upper Chalk". plates in each series (7) than both other species. In how much this différence is sufficiënt to justify the Occurrences outside the Benelux-countries : distinction of T. danica as a separate species, remains F: Turonian of Sarthe dept. (Cotteau & Triger, to be etablished. Examination of Ravn's type speci¬ 1860); Coniacian to Maastrichtian of Aude dept. mens would be necessary. (lambert, 1911), Charente, Charente-Maritime, Eure, Eure-et-Loire, Loir-et-Cher, Marne, Oise, Typocidaris sceptrifera (KONIG in mantell, 1822) Seine-et-Oise, Seine-Maritime, Somme and Yonne (Pl. 1, Figs. 7-9) depts. (Cotteau, 1862). GB: Santonian of Kent, Sus¬ sex and Wiltshire (Wright, 1882). D: Upper Turo¬ * . 1822 Cidaris sceptrifera, MANTELL, p. 194, pl. 17, fig. nian of Salzgitter (Schlüter, 1892). DDR: Lower 12(r). Maastrichtian of Rügen (Roemer, 1841). DK: Conia¬ . 1840 Cidaris sceptrifera, agassiz, p. 10. cian and Santonian of Bornholm (Ravn, 1928). E: . 1841 Cidarites sceptrifera, roemer, p. 28. Campanian of Tremp-area (Lambert, 1925). SU: . 1846 Cidaris sceptrifera, agassiz & desor, p. 328. Coniacian of the to . 1848 Cidaris sceptrifera, Bronn, p. 300. Don-Basin; Turonian Campanian of the 1850 Cidaris sceptrifera, SORIGNET, p. 6. Volhynian-Podolian Plateau, Ukranian S.S.R. 1850 Cidaris sceptrifera, DlXON, p. 338. (Hynda, 1968; Savchinskaya, 1974). 1854 Cidaris sceptrifera, MORRIS, p. 74.
. 1855 Cidaris sceptrifera, DESOR, p. 13, pl. 5, fig. 28(r). Figured specimens in the K.B.I.N.-collections: 1859 Cidaris sceptrifera, coquand, p. 1013. IST 10250, figured herein, Pl. 1, Fig. 8. IST 10251, . 1860 Cidaris sceptrifera, cotteau & Triger, p. 253, figured herein, Pl. 1, Fig. 7. IST 10252, figured herein, pl. 42, figs. 1-8 (r + c). Pl. 1, Fig. 9. (Specimens IST 9093 and IST 9094 are . Cidaris sceptrifera, COTTEAU, pp. 251-256, pl. 1056, figs. 57-58. incomplete and isolated radioles, figured by Smiser
. (1935), pl. 2, fig. 2 a-b). v 1874 Cidaris sceptrifera, COTTEAU, p. 641. 1875 Cidaris sceptrifera, Quenstedt, p. 175, pl. 68, fig. 4. Specimens studied: 1878 Cidaris sceptrifera, COTTEAU, pp. 430-434, pl. 78, Maastricht (NLb, NL); Maastricht Formation, Maas¬ figs. 2-5. trichtian: 16 fragments, among which IST 10250, Typocidaris in the type-Maastrichtian 205
10251 and 10252. Valkenburg (NLb, NL); Maastricht named, based on radioles. A corrélation between Formation, Maastrichtian: 22 fragments. Sibbe (NLb, these radioles and coronal parts has subsequently been NL); Maastricht Formation, Maastrichtian: 2 frag¬ made by Forbes in Dixon (1850), who figured two ments. Vroenhoven (BLb, B); Maastricht Formation, nearly complete coronas, in connection with radioles. Maastrichtian: 1 fragment. A clear description of coronal plates was first given by Cotteau & Triger (1860). Hence, we use the Description: name T. sceptrifera in the meaning of the latter Interambulacral primary tubercles are perforate, non authors. crenulate. The diameter of the scrobicule is 3.5 times Specimen n° V85 of Agassiz' collection, was included larger than the mamelon's. Only the mamelon protru- in the synonymy of T. sceptrifera by desor (1855), des over the scrobicular ring. Some scrobicules are by Cotteau & Triger (1860) and by Cotteau slightly elliptical. Scrobicules are deeply sunken. Their (1862). This must be an error. According to Lambert distal borders are steep, but never overhanging. Scro¬ & Jeannet (1928), specimen n° V85 belongs to Pseu- bicular rings are complete and raised. They consist of docidaris galeotti (desor, 1855) from the Cenoma- 14 coarse tubercles, alternating with smaller, radially nian of Mexico ! elongated granules. Scrobicules are not confluent, but Some authors have confused T. sceptrifera with Stereo- scrobicular cidaris cretosa rings of adjacent plates touch each other (Mantell, 1835). As early as 1840, adorally and adapically. Only in the vicinity of the Morris considered them as synonymous. Yet, Man- apical System, a narrow row of very small granules tell's interprétation of both species was mainly based can occur along the horizontal sutures. The uppermost on earlier figures, without proper description or expia- scrobicule in each ambulacrum is reduced and its nation. The situation was clarified by Cotteau tubercle rudimentary. Adradial extrascrobicular sur¬ (1862), who re-established, with full motivation, the faces are exceedingly narrow and limited to an irre- existence of both species. In spite of cotteau's argu¬ gular row of small granules. Adorally, the scrobicules mentations, the synonymy of these species persisted are in the moderately wide and covered by an irregular, works of subséquent British authors, such as dense, fine granulation. The interradial suture is Wright (1882), who figured a specimen of S. cretosa, depressed in a conspicuous furrow. On the adapical under the name T. sceptrifera (Pl. 6, Figs. la-lb). The half of the corona, horizontal sutures are marked by same error was made by Mortensen (1928) (Fig. 76) distinct grooves. and by Fell (1966) (Figs. 242-3a, b), who reproduced Ambulacra are sinuous and moderately narrow. Pori- Wright's figure. S. cretosa is easily distinguished ferous zones are strongly depressed. Pores are not from T. sceptrifera by its strongly reduced adapical conjugate, but separated by a granuliform interporous scrobicules, and its much wider interporiferous zones partition. Neural grooves can be seen. The axes of and interradial extrascrobicular surfaces. S. cretosa is the pore-pairs are almost horizontal. Interporiferous the type-species of the genus Stereocidaris. areas are covered by a dense granulation. These gra¬ nules are arranged in four vertical series. A fifth and Typocidaris serrata (desor, 1858) a sixth series can occur in the vicinity of the ambitus. (PI. 2, Figs. 1-4) The outermost granules are slightly larger than the * others. There is no . 1858 horizontal regularity in their arran¬ Cidaris serrata, desor, p. 450 (pro parte). gement. The ratio ambulacral to interambulacral . 1862 Cidaris serrata, cotteau, pp. 306-308, pl. 1074, plates, at the ambitus, is 20 to 1. figs. 1-11. . 1867 Cidaris hirudo, Wright, pp. 64-67, pl. 10, figs. 2-3 Discussion: (pro parte) (non sorignet, 1850). 1887 Cidaris serrata, Gauthier, pp. 251-252 Typocidaris sceptrifera has been (radioles). compared to T. sub- . 1892 Cidaris serrata, schlüter, vesiculosa p. 83. (d'Orbigny, 1850) (Cotteau, 1862, 1897 Cidaris serrata, Lambert, p. 142, pl. 2, fig. 9. Pl. 1059, 1060, 1061; 1910 Upper Cretaceous of Western Typocidaris serrata, Lambert & Thierry, p. 152. 1911 Europe). Yet, T. sceptrifera is easily recognised by its Typocidaris serrata, Lambert, pp. 34, 42, 46, 69.
v . 1935 strikingly coarse, raised scrobicular rings. Moreover, Typocidaris serrata, Smiser, p. 23, pl. 1, fig. 12c its interradial extrascrobicular surfaces are narrower (non fig. 12b). 1939 and its granulation is coarser than in T. subvesiculosa. Typocidaris serrata, KONGIEL, pp. 12-13, pl. 1, figs. 21-25. Very coarse scrobicular rings are also a characteristic 1968 Stereocidaris serrata, hynda, feature in T. forchhammeri (agassiz & desor, p. 197, pl. 40, fig. 10. 1970 Typocidaris serrata, Blaszkiewicz, 1846) (Cotteau, Pl. p. 159. 1863, 1078, 1079; Danian of v ? 1984 Stereocidaris cf. serrata, Naidin, p. 26. Western Europe). The latter species shows only two vertical series of granules in its interporiferous sur¬ Locus typicus: faces, while T. sceptrifera has four or six. The scrobi¬ Meudon, Seine-et-Oise, France. cules of T. forchhammeri are shallow, while those of T. sceptrifera are deep. Stratum typicum: The species under discussion has been described and "Sénonien supérieur". 206 Joris f. geys
Occurrences outside the Maastricht-area: 2. Interradial and horizontal sutures depressed. F: "Sénonien" of Seine-et-Oise (COTTEAU, 1862), 3. Scrobicular rings with 15 small tubercles. Marne (Gauthier, 1887); Campanian of Charente 4. Interporiferous areas moderately narrow, with six (Geys, unpublished). PL: Maastrichtian of the Lublin vertical series of granules, of which the perradial area (Kongiel, 1939). SU: Upper Maastrichtian of ones are very small. the Lvov-area, Ukrainia (Hynda, 1968); Maastrich¬ tian of Mangyshlak-peninsula, Kazachstan (naidin, Discussion : e. a., 1984). Typocidaris serrata has few characteristic features and therefore, the species is difficult to recognise. Confu¬ Figured specimens in the K.B.I.N. -collections: sion with some other species is easy. Some features IST 9086, figured by Smiser (1935), pl. 1, fig. 12c. permit however, to distinguish them. Areoles are IST 9087 and IST 9088, resp. a coronal fragment and much deeper in T. hagenowi (Desor, 1858) (Nest- a radiole, figured by Smiser (1935; pl. 1, fig. 12a-b) ler, 1972, Pl. 4-5; Maastrichtian of Rügen, D.D.R.) and 9083, IST 9084 and IST 9085, figured and des- and in T. sceptrifera (koenig in mantell, 1822). cribed by Lambert (1897, 1910), are unconvincing. Moreover, interporiferous zones of T. hagenowi have a very different granulation. Scrobicular rings are Specimens studied: coarser and ambulacra are narrower in T. forchham- Sibbe (NLb, NL); Maastricht Fm., Maastrichtian: meri (Agassiz & Desor, 1846). Interradial extra¬ 1 interambulacrum with parts of adjacent ambulacra scrobicular surfaces are wider and/or differently struc- (IST 9086). Kanne (Albert Canal) (BLb, B); lst Bryo- tured in T. subvesiculosa (Orbigny, 1850) (Cot- zoa-bed, Meerssen Beds, Maastricht Fm., Maastrich¬ teau, 1862, pl. 1059-1061; Upper Cretaceous of Western tian : 1 interambulacrum with parts of adjacent ambula¬ Europe), in T. hirudo (Sorignet, 1850) cra (coll. J. Jagt, n° 2283). Vroenhoven (Albert Canal) (Cotteau, 1862, pl. 1054; "Sénonien" of Western (BLb, B): Geulhem Beds, Houthem Fm., Danian: 1 Europe), in "Cidaris" faujasi Desor, 1856 (Cot- specimen (coll. J. Jagt n° 494). Maastricht (St. Pieters¬ teau, 1863, pl. 1077; "Sénonien" of France) and in berg) (NLb, NL); Maastricht Fm., Maastrichtian: 2 T. serrifera (forbes, 1850) (cotteau, 1862, pl. fragments (coll. NHMM n° MM-7262). Kunrade (NLb, 1071; "Sénonien" of France). NL); Kunrade Fm., Maastrichtian: 1/5 corona (coll. The holotype of T. serrata has never been figured. M. Van Birgelen). Hallembaye (CPL-quarry) (Lg, B); The first proper description and useful figures of the Lanaye Beds, Gulpen Fm., Lower Maastrichtian: species were published by cotteau (1862). hence, 3 impressions in flint (coll. L. Indeherberge). the name had to be used herein, sensu Cotteau. For reasons explained above, part of the specimens Description: figured as Cidaris hirudo, by Wright (1867) (pl. 10, Ambulacra are sinuous and moderately narrow. Pori- figs. 2-3), problably belong to T. serrata. T. serrata has been mentioned from the Lower Maas¬ ferous zones are depressed. The pores are not conju- gate. The axis of the pore-pairs is subhorizontal. trichtian of Rügen (d.d.R.) by desor (1858) and Interporiferous areas are moderately narrow and show by schlüter (1892). These records are based on six vertical series of granules. Adradial granules are incomplete radioles and therefore highly question- able. more important in size than perradial ones. At the ambitus, the ratio of ambulacral to interambulacral plates is 15 to 1. Interambulacral plates are 5 in a series. Primary tuber- Typocidaris ubaghsi nov. sp. cles are perforate, non crenulate. The areoles are (Pl. 2, Figs. 5-9) conical and moderately shallow, so that the bosses 1928 Stereocidaris cf. Merceyi, Krenckel, p. 18, fig. 2, figs. protrude highly over the rim of the scrobicular rings. 21-23. Scrobicular rings are complete and consist of 15 small scrobicular tubercles, which are only slightly larger than surrounding granules. In each ambulacrum, the Locus typicus: uppermost scrobicule is reduced. Extrascrobicular sur¬ Heure-le-Romain, prov. Liège, Belgium. faces are densely granulated. Adradially, these sur¬ Stratum faces are interrupted by the scrobicular rings, which typicum: touch the adradial sutures. Interradial extrascrobicu¬ Lower Gulpen Formation, Upper Campanian. lar surfaces are well developed only on the adapical side. Horizontal and interradial sutures are depressed Holotype: and visible as faint grooves on the corona. K.B.I.N., n° IST 10253.
Diagnostic features: Occurrence outside the Benelux: 1. Interradial extrascrobicular area relatively narrow DDR: Lower Maastrichtian of Rügen (H. KRENCKEL, and sinuous. 1928). Typocidaris in the type-Maastrichtian 207
Dimensions : than the former, for specimens of similar size. Still D = 42-48 mm; h = 29 mm; ds = 13-23 mm; dp = more important: diagnostic features for Stereocidari- 10-17 mm; h/D = 0,60-0,69; ds/D = 0,31-0,48; dp/D nae, such as the high shape of interambulacral plates = 0,24-0,35. and suturai grooves, are not sufficiently prominent in C. perlata. Hence, the relationship between T. Description : ubaghsi and C. perlata is more than questionable: the The peristome is not sunken; gill slits are absent. latter species probably does not belong to the Stereo- Ambulacra are sinuous and wide. Poriferous zones cidarinae at ail. are depressed. The pores are not conjugate, but a The ambulacra of Stereocidaris s.s. cretosa (Mantell, neural groove is visible near the adorai suture of each 1835) (Cotteau, 1862, pl. 1067; "Sénonien" of plate. A granule is present between the pores of each Western Europe) are also very similar to those of T. axes pair. The of the pore-pairs are slightly oblique. ubaghsi. However, the granules of S. cretosa are The interporiferous areas are densely granulated. arranged in 6 to 8 regular, vertical series. Those of Adradially, there is a single regular, vertical series of T. ubaghsi are arranged in horizontal rows, without granules, each corresponding with a pore-pair. Perra- any vertical regularity. The same lack of vertical regu- dially, the remaining parts of each interporiferous area larity is displayed in the ambulacral granules of Stereo¬ are covered with a very dense, irregular, fine granula¬ cidaris s.s. merceyi (cotteau, 1862) (pl. 1068; Lower tion. The latter granules are not "Sénonien" of arranged in vertical France). Yet, both species are clearly series, but in horizontal rows of 5 to 7. There are different from T. ubaghsi, in having strongly reduced, about 80 plates in each ambulacral series. At the ambi- rudimentary adapical interambulacral scrobicules. For tus, 23 ambulacral plates correspond in that height to one reason, T. ubaghsi does not belong to the genus interambulacral plate. Stereocidaris s.s. Interambulacral plates are five in a series. Primary tubercles are perforate, non crenulate. The areoles Phylogenetical considérations: are slightly conical and deeply sunken. The distal The closest relative known, to the species under dis¬ border of the areoles is vertical, or undercut. The cussion is probably Typocidaris hagenowi (desor, areoles are surrounded by 19 small, scrobicular tuber¬ 1858), from the Lower Maastrichtian of cles. Rûgen, The uppermost scrobicule of each interambula- D.D.R. Its ambulacra have the same structure as crum is rudimentary. Extrascrobicular surfaces are those of T. ubaghsi: vertical series of well developed covered by a very dense, fine granulation. Interradial, primary tubercles on the adradial sides of the inter¬ as well as adorai extrascrobicular areas are well deve- poriferous areas. Perradilally, smaller granules occur, loped and wide. Adradially, these granulated surfaces arranged in two horizontal rows on each are much plate. T. narrower. The granules are arranged in hagenowi differs from T. ubaghsi, because the vertical more or less regular rows, radiating from the scrobicu- regularity in the arrangement of perradial granules les and separated by faint furrows, running from each did not completely disappear in the former. These scrobicule to the scrobicules on both adjacent plates granules are less numerous and less of the other series. tightly packed in Horizontal, as well as interradial T. hagenowi than in T. ubaghsi. sutures are clearly visible and A depressed. plausible interrelationship is illustrated in the cladogram (Fig. 1). From this figure, one can read Diagnostic features: that the similarity between the ambulacra of S. mer¬ 1. Widely separated interambulacral scrobicules. ceyi and T. ubaghsi is to 2. 5 or 6 probably due convergence. interambulacral plates in a series. During its évolution, T. ubaghsi and its ancestors 3. Uppermost interambulacral plate reduced. can have passed the 4. Distinct suturai following thresholds: a/ dif- grooves. ferentiation from a 5. Ambulacral Typocidaroid ancestor, by the interporiferous areas wide, with two acquisition of multiple horizontal rows of regular vertical series of perradial granules, separated by a granules on each ambulacral wide plate (threshold 4), perradial belt of small granules, arranged in b/ differentiation from the stem-T. hagenowi by the irregular horizontal rows. loss of vertical regularity in perradial granules (thres¬ hold 5). Affinities: The fragments from the Lower The new Maastrichtian of species shows a superficial similarity to "Ci- Rügen (D.D.R.), which were daris" s. 1. provisionally identified perlata (sorignet, 1850) (G. cotteau, as Stereocidaris cf. merceyi by Krenckel (1928), pro¬ 1862, pl. 1062, 1063), from the "Senonian" of France. bably to Both belong T. ubaghsi. Krenckel's description species have wide perradial fits the new interporiferous species very well. I disagree with Nestler areas, a covered with dense granulation. But on closer (1972) who believed Krenckel's specimens to belong examination, very important différences corne to to light. Temnocidaris baylei The latter The ambulacra of T. Cotteau, 1863. are wider and more ubaghsi species can be identified sinuous than in C. easily by its numerous coro- perlata. The latter nal species has consi- dépressions, which are absent in the derably more plates in each interambulacral fragments series described by Krenckel. 208 Joris f. geys
Doubtful species Folx-les-Caves (B) and Ciply (B). The same radioles were attributed to Temnocidaris danica, by lambert By far the great majority of the Cidaroid remains in (1897). Later, this author changed his opinion (lam¬ Cretaceous strata consists of isolated interambulacral bert, 1911): radioles from the Ciply Phosphatic plates and radioles. The identification of both kinds Chalk were reinterpreted as belonging to D. faujasi. of fossils is particularly difficult, especially when their Isolated interambulacral plates were attributed to the préservation is not optimal. Unfortunately, this is very same species (coll. KBIN, n° IST 9078), the main often the case in the material I studied. argument was their occurrence is the same beds as the radioles. In a large number of species, interambulacral plates Repeating Lambert's statements, offer little or no diagnostic features, which allow an without due criticism, radioles and fragments of D. unequivocal identification. The interprétation of iso¬ faujasi were also mentioned by smiser (1935) from lated interambulacral plates, not in connection with the Ciply Phosphatic Chalk (coll. KBIN, n° IST 9079, neighbouring plates, or with parts of an ambulacrum, 9080, 9081). Radioles referred to D. is very often an almost impossible task. Establishing faujasi, are very similar to a the presence of a given species on such poor evidence, large number of other Cidaroid spines. The corona is is obviously risky. Still less justified is the introduction still unknown, and therefore the species remains doubtful. More and better of new species, merely based on a few such fragments, preserved fossils are although this was common practice during the 19th needed, before the existence of D. faujasi can be established. century. Hence, names created for isolated coronal c. "Stereocidaris" plates must be approached with caution. The safest pseudohirudo (cotteau, 1862) was erected for isolated radioles from the Campanian way to deal with them should be to consider them all invalid. of Meudon (F). Similar radioles have been reported Analogous considérations can be made for taxa based by Lambert (1911) from the Nouvelles Chalk at Har- on isolated radioles. Radioles, in anatomical associa¬ mignies (Hainaut, B) (coll. KBIN, n° IST 9052). This statement has been tion with the corona, are only known in a very restrict- repeated uncritically by Smiser ed number of species. To my opinion, only such spe¬ (1935). d. cies can have identifiable radioles. When giving new "Typocidaris" arenata lambert, 1911 was esta¬ blished for a names to isolated radioles, the risk of creating syno- single, isolated interambulacral plate from the Lower Gulpen Chalk (Campanian) at Heure- nyms is real. These radioles can belong to species le-Romain which are already known, but of which no specimen (Liège, B) (coll. KBIN, n° 9092). The plate is not unlike those of T. pistillum (Quenstedt, was ever found with the spines attached. The situation 1852), which occurs in strata of the same age. No data being is still more complex because differently shaped radio¬ available on the ambulacra of T. arenata, the specimen les do not necessary belong to different species: the described by same animal can have several types of spines, covering lambert is in fact unidentifiable. e. "Balanocidaris" schlueteri Lambert, 1911 has its test. Inversely, some echinoids have radioles which been introduced for a single, isolated radiole from the can hardly be distinguished from each other. As a rule, isolated Cidaroid radioles can sometimes be Ciply Phosphatic Chalk (Maastrichtian) at Ciply (Hai¬ naut, B) (coll. KBIN, n° a identified up to genus-level, but rarely more accurate- IST 9068). It is club-shaped radiole, but its state of préservation does not permit ly. Only very few species can be recognised, by their to radioles alone. For these reasons, the naming of iso¬ distinguish the presence or absence of a perforation in its acetabulum. Hence, the radiole could belong to lated radioles should be banned. Existing names, as as merely based on such material, must be considered as Tylocidaris, well to Balanocidaris. f. "Cidaris" ciplyensis Lambert, 1911 is a name used referring to morphospecies, not to taxonomie units. for some small, isolated radioles from the Ciply Phos¬ From the preceeding, one can conclude that certain statements concerning the presence of species in the phatic Chalk (Maastrichtian) at Ciply (Hainaut, B) Maastrichtian of Belgium and The Netherlands, (coll. KBIN, n° IST 9059, 9060). These radioles were described as C. montainvillensis Lambert, 1897 should be approached with caution. These "doubtful (Dano-Montian of the species" are shortly discussed below. Paris-Basin, France). The holo- a. "Dorocidaris" venulosoides (schlüter, 1897) has type of this taxon later showed to be a spine of Typo¬ cidaris been reported from the Ciply Phosphatic Chalk at forchhammeri (agassiz & Desor, 1846), so that the Ciply radioles were in need of a new name Ciply, by Lambert (1911) (coll. KBIN n° IST 9082), who described a few isolated interambulacral plates. (Lambert, 1911). Perhaps misinterpreting Lam¬ bert's words, Smiser (1935) united C. ciplyensis with b. "Dorocidaris" faujasi (Desor, 1855) has been C. forchhammeri. syntypes, established as a "species", based on isolated radioles Both figured by lam¬ bert from Maastricht. Without acceptable arguments, a (1897) are badly corroded fragments of radioles, showing no details of the corona from the Cotentin peninsula was attributed to base and lacking the top. They are in reality far less well preserved than the same species, by cotteau (1865). Similar radio¬ shown on Lambert's figures. In my opinion, both syntypes les were reported by cotteau (1865, 1874) from the are unidentifiable and C. ciplyensis should be ignored. Maastrichtian of Maastricht (NL), Valkenburg (NL), Typocidaris in the type-Maastrichtian 209
g. "Balanocidaris" hardouini Desor, 1855 has been j. "Cidaris" perornata forbes, 1850 is a rare species introduced for some isolated, small, club-shaped from the Upper Chalk of Sussex, England. Whereas radioles from the Ciply Tuffaceous Chalk (Danian), at least one well preserved specimen, from England, at Ciply (Hainaut, B). Similar spines have been found with radioles attached, has been described by forbes by Binkhorst (1859) in Maastrichtian or Danian (1850), the fossils from continental Europe, assigned strata in the Maastricht area (Limburg, NL), by cot- to the same species merely consist of isolated radioles teau (1874) and by Lambert (1897) in the Malogne and small fragments. Smiser (1935) reported the spe¬ Beds (Maastrichtian or Danian) at Ciply and by cies from the Maastrichtian at Maastricht (NL), but smiser (1935) in the Maastrichtian or Danian at his statement is based merely on some isolated spines Geulhem (Limburg, NL). The occurrence of this spe¬ (coll. KBIN, n° IST 9077). These spines are slender, cies in the Maastrichtian has not been convincingly thorny and similar to those attributed to "C. " spinosis- demonstrated. Meijer (1959) as well as Geys & sima Agassiz, 1846, "C." leptacantha Agassiz, 1846 Marquet (1983) consider it a typical Danian species, or indeed, "C." perornatal Because of these uncer- belonging to the genus Tylocidaris. Radioles of Tylo- tainties, I consider the presence of the latter species cidaris are indeed common in deposits of Danian âge in the Upper Cretaceous of the Low Countries not (Brotzen, 1959). The specimens figured by Smiser demonstrated. (1935) (coll. KB1N, n° IST 9072, 9073, 9074, 9075 and k. Typocidaris forchhammeri (agassiz & desor, 9076) are somewhat eroded and show almost no 1846) was first described from the Danian at Faxe definite generic characteristics. Yet, they probably (DK). It was subsequently reported from the Dano- belonging to Tylocidaris. Montian at Vigny (F) and from Mons (B). Unfortuna- h. "Cidaris" filamentosa Agassiz, 1846. A few iso¬ tely, a lot of nomenclatorial confusion exists for the lated radioles from Maastricht (NL) were assigned to Belgian specimens. The collections of the K.B.I.N. this species by Smiser (1935) (coll. KBIN, n° 9061 posses a large number of coronal fragments, which and 9062). The species has been named by Agassiz are to be classified in this species. They are labelled & Desor (1846) after a few isolated spines of "Maastrichtian - Voort colliery, Limburg, Belgium". unknown origin. The corona, corresponding to these Knowing that Danian strata in Limburg were often radioles is completely unknown. The similarity misinterpreted as Upper Maastrichtian, and taking between the radioles described by smiser (1935) and into account the known stratigraphical distribution of some, which were attributed to "C." forchhammeri the species elsewhere, thcre is no unequivocal évi¬ desor, 1846, is striking. Therefore I consider the dence for the presence of T. forchhammeri in the presence of "C" filamentosa in the Low Countries Belgian Upper Cretaceous. Most probably, the spe¬ unsufficiently demonstrated. cies is confined to Danian and Montian deposits. i. "Cidaris" subvesiculosa d'orblgny, 1850 is a well known and common species in the "Senonian" of Western Europe. It has been reported by Bosquet Acknowledgements (1957) and by Mourlon (1880), from the Maastrich¬ tian of Limburg (NL), in a list, without discussion. I would like to express my sincere gratitude to Dr. X. MisONNE, smiser director of the K.B.I.N. at to P. (1935) figured two coronal fragments and one Brussels, Dr. sartenaer, head of the radiole (coll. KBIN, n° IST 9089, 9090, 9091), which department of Palaeontology and to Dr. L. Van De Poel, head of the section for Invertebrates from he attributed to this Secundary and Tertiary (IST), species. Both fragments are badly for permission and facilities to study the collections in their care. I corroded and too small to be properly identified. The also wish to thank Drs. A.W.F. meijer (Natuurhistorisch Museum, radiole is similar to those, figured by cotteau (1864) Maastricht, the Netherlands) for the opportunity to examine some as Cidaris subvesiculosa, but also to those of C. ser- specimens from the collections of his institution, as well as Mr. J. Jagt rata, and to some of C. sceptriferal Hence, also this (Venlo, The Netherlands), Mr. L. indeherberghe (Zon¬ hoven, Belgium) and Mr. M. Van birgelen (Heerlen, The radiole is hardly identifiable. The presence of C. sub¬ Netherlands), for the loan of specimens from their private collections. vesiculosa in the Maastrichtian of the Low Countries I particularly want to thank Dr. A.V. Dhondt (K.B.I.N.. Brussels) has still to be demonstrated. for critically reading the manuscript and for help in many ways. 210 Joris F. GEYS
References agassiz, l., 1840. Catalogus systematicus ectyporum echi- d'Orbigny, A., 1850. Prodrome de Paléontologie stratigra- nodermatum fossilium musei Neocomensis. Petitpierre, Neu- phique universelle des animaux mollusques et rayonnés, châtel, 20 pp. faisant suite au Cours élémentaires de Paléontologie et de Géologie stratigraphique. 2ième volume. Masson, Paris, Agassiz, L. & Desor, E., 1846. Catalogue raisonné des 427 pp. familles, des genres et des espèces de la Classe des Echino- dermes I. Annales des Sciences Naturelles 3, Zoologie 6: Fell, H.B. & Pawson, D.L., 1966. Echinacea. In moore, 305-374. R.C. (editor), Treatise on Invertebrate Paleontology, Part U, Echinodermata 3. Geological Society of America, New Binkhorst Van Den Binkhorst, J.T., 1859. Esquisse géo¬ York and Lawrence, pp. U367-U437. logique et paléontologique des couches crétacées du Lim- Fischer, bourg et plus spécialement de la craie tuffeau. Van Osch- J.-C., 1980. 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Grenoble, fossiles observés dans la formation crétacé du sud-ouest da 96 pp., 6 pl. la France. Bulletin de la Société Géologique de France (2) gravesen, P., 1979. Remarks on the regular echinoids in 16: pp. 945-1023. the Upper Maastrichtian and Lower Danian of Denmark. In cotteau, G., 1857-1878. Etude sur les échinides fossiles du Birkelund, T. & bromley, R. (editors), Cretaceous- département de l'Yonne. Baillière, Paris, 518 pp., 84 pl. Tertiary boundary Events. Symposium I. The Maastrichtian and Danian of Denmark. Kpbenhavn, pp. 72-73. cotteau, G., 1861-1867. Paléontologie française. Descrip¬ Gray, J.E., 1825. An or tion des animaux invertébrés commencée par Alcide d'Orbi- attempt to divide the Echinida, Sea Eggs, into natural families. Annals of Philosophy (N.S.) gny. Terrain Crétacé 7, Echinides réguliers. Masson, Paris, 10: 423-431. 892 pp., pl. 1007-1204. Hynda, V.A., 1968. Typ Echinodermata, Pidtip Eleuthero- cotteau, g., 1874. Notes sur les échinides crétacés de la zoa. 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Plate 1
Fig. 1. Typocidaris pistillum (QuENSTEDT, 1852) CPL-quarry, Hallembaye (Lg, B); Zevenwegen Chalk, Gulpen Fm., Upper Campanian; coll. J. Jagt, Venlo (n° 1180); adapical view; x 2.
Fig. 2. The same; latéral view; X 2.
Fig. 3. The same; adorai view; x 2.
Fig. 4. The same; adapical part of ambulacrum; x 8.
Fig. 5. The same; ambital part of ambulacrum, showing perradial granulation; x 8. Fig. 6. Typocidaris pistillum (Quenstedt, 1852) CPL-quarry, Hallembaye (Lg, B); Lanaye Chalk, Gulpen Fm., Lower Maastrichtian; coll. L. Indeherberghe, Zonhoven; fragment; x 6. Fig. 7. Typocidaris sceptrifera (Konig in Mantell, 1822) Maastricht (NLb, NL); Maastricht Fm., Maastrichtian; coll. KBIN (n° IST 10251); fragment; x 5. Fig. 8. Typocidaris sceptrifera (Konig in Mantell, 1822) Maastricht (NLb, NL); Maastricht Fm., Maastrichtian; coll. KBIN (n° IST 10250); fragment; x 4. Fig. 9. Typocidaris sceptrifera (Konig in Mantell, 1822) Maastricht (NLb, NL); Maastricht Fm., Maastrichtian; coll. KBIN (n° IST 10252); fragment; x 5. 214 Joris F, GEYS Typocidaris in the type-Maastrichtian 215
Plate 2
1. Fig. Typocidaris serrata (Desor, 1858) Sibbe (NLb, NL); Maastrichtian; coll. KB1N (n" IST 9086); latéral view; x 4. Fig. 2. The same; adapical view; x 4.
Fig. 3. Typocidaris serrata (Desor, 1858) Albert Canal section, Kanne (BLb, B); Meerssen Chalk, Maastricht Fm., Maastrich¬ tian; coll. J. Jagt, Venlo (n° 2283); latéral view; x 4.
Fig. 4. The same; detail of ambulacrum; x 11.
Fig. 5. Typocidaris ubaghsi nov. sp. Heure-le-Romain (Lg, B); Lower Gulpen Fm., Upper Campanian; coll. KBIN (n° IST 10253); adapical view; x 1,2.
Fig. 6. The same; adorai view; x 1,2.
Fig. 7. The same; latéral view; x 1,2.
Fig. 8. The same; detail of interambulacrum; x 4.
Fig. 9. The same; detail of ambulacrum; X 4.