THIOLLIERICRINID CRINOIDS FROM THE LOWER CRETACEOUS OF CRIMEA

by

Vladimir G. KLIKUSHIN

Geobios, n° 20, fasc. 5 p. 625-665, 18 fig., 2tabl., 1 pi. Lyon, octobre 1987 THIOLLIERICRINID CRINOIDS FROM THE LOWER CRETACEOUS OF CRIMEA

by

Vladimir G. KLIKUSHIN *

Abstract RÉSUMÉ

The family Thiolliericrinidae (Oxfordian - Hauteri- La famille Thiolliericrinidae (Oxfordien - Hauteri- vian) includes the stalked crinoids of the order Coma- vien) comprend les crinoïdes pédonculés de l’ordre tulida, inhabiting reef fades. Remains of 12 thiollieri- Comatulida, vivant dans des milieux récifaux. 12 crinid species (10 among them are new) belonging to espèces (dont 10 sont nouvelles) de Thiolliericrinidae six genera (Thiolliericrinus, Burdigalocrinus, Lorioli- sont décrites du Berriasien et du Yalanginien de la crinus, Conoideocrinus nov. gen., Umbocrinus nov. Crimée. Ces espèces appartiennent à six genres : gen., and Heberticrinus nov. gen.) from the Berria- Thiolliericrinus, Burdigalocrinus, Loriolicrinus, sian and Valanginian of Crimea are described. On the Conoideocrinus nov. gen., Umbocrinus nov. gen. et basis of an extensive material (hundreds of calyxes, Heberticrinus nov. gen. L’étude d’un matériel abon­ centr odor sals, brachials and columnals) the skeletal dant (des centaines de coupes dorsales, de pièces cen- morphology, the ontogenetic variations of the calyxes trodorsales et brachiales, de columnales) permet une and the phylogeny of the Thiolliericrinidae are discussion sur la morphologie du squelette, les varia­ reconstructed. The family taxonomy is defined more tions ontogénétiques des coupes dorsales et la phylo­ exactly regarding new morphological data. génie des Thiolliericrinidae. D’après les nouvelles données morphologiques la taxonomie de la famille est précisée.

KEY-WORDS : CRINOIDEA, ARTICULATA, COMATULIDA, THIOLLIERICRINIDAE, TAXONOMY, NOMENCLATURE, MOR­ PHOLOGY, ONTOGENY, PHYLOGENY, CRETACEOUS, USSR.

MOTS-CLÉS : CRINOÏDES, ARTICULATA, COMATULIDA, THIOLLIERICRINIDAE, TAXONOMIE, NOMENCLATURE, MOR­ PHOLOGIE, ONTOGENIE, PHYLOGENIE, CRÉTACÉ, URSS.

* Leningrad Mining Institute, 199026 Leningrad, USSR.

Geobios, n° 20, fasc. 5 p. 625-665, 18 fig., 2tabl., 1 pl. Lyon, octobre 1987 TABLE OF CONTENTS

1 - Introduction...... p. 626 5 - Reconstruction of the skeleton...... p. 652

2 - Localities and materials...... p. 626 Root formations ...... p. 652 Stem ...... p. 653 3 - Terminology...... p. 629 A rm s...... p. 653 4 - Taxonomie description...... p. 631 6 - Ontogenetic variations of the calyxes ...... p. 656 Genus Solonaerium E tallon emend. Ra s m u s s e n ...... p. 633 7 - Origin of the Thiolliericrinidae...... p. 657 Genus Thiolliericrinus E t a l l o n ...... p. 634 8 - Development of the Thiolliericrinidae...... p. 658 Genus Burdigalocrinus Ja e k e l ...... p. 639 Genus Loriolicrinus J a e k e l ...... p. 641 Acknowledgements...... p. 660 Genus Conoideocrinus nov. gen...... p. 644 R eferences...... p. 660 Genus Umbocrinus nov. gen...... p. 646 Genus Heberticrinus nov. gen...... p. 646

1 — INTRODUCTION

Sea-lilies of the Comatulida are freely mobile forms simularities with the Bourgueticrinida stems. Such which are widely distributed in the modern seas. combination of the features lead to complicated pro­ Calyxes of these crinoids, consisting of five radiais, blems for attempts to clear up the phylogenetical con­ five basais and one centro-dorsal plate bearing cirri, nections of the family and attracted many research are also found in the fossil record. workers to the study of this group. However, one peculiar exception exists in that large Nevertheless, many aspects remain in the nomencla­ group : the fossil family Thiolliericrinidae. The repre­ ture, the taxonomy and the phylogeny of the family sentatives of this family were stalked and led a fixed Thiolliericrinidae. The abundant new material at the life. The thiolliericrinid thecae were constructed like author’s disposal allows to propose a solution of these the true Comatulida, but their columns show problems.

2 — LOCALITIES AND MATERIALS

Remains of the Thiolliericrinidae were found in - 1 : the « yet-formed » bioherms. They are small some regions of the Crimea (fig. 1). The Belbek Val-'/ lenses (not more than one metre thick and up to 10 m ley is the most important among them according to long) enclosed into the strata of the microphytolithic the quantity of found specimens and to the diversity or bioclastic stratified limestones. The lenses are built of forms (table 1). In that locality, crinoids under dis­ with skeletons of colonial and solitary corals, calca­ cussion were found in the Upper Berriasian coral- reous algae and sponges. Oyster shells (Lopha, algal or coral-sponge bioherms. Three types of them Exogyra and others), brachiopod shells, echinoid spi­ can be distinguished : nes (Diplocidaris) are found in a great quantity. — 627 —

Fig. 1 — Localities of Thiolliericrinidae in the Crimea. Gisements à Thiolliericrinidae en Crimée.

Chornaya Basin (1 - Kuchky, 2 - Rodnoie), Belbek valley (3 - Belbek, Kuibyshevo, 4 - Karatlykh, 5 - Chuku), 6 - Bairakly, 7 - Tass- Kor Ravin (Mramornoie), 8 - Beshterek, 9 - Mezhgorie, 10 - Tanass (Alekseevka and Krasnossiolovka).

Thiolliericrinidae are found but extremely rarely in - 3 : «tower-like » bioherms, are the intermediate these bioherms ; type between the two previous ones. They show a dis­ coid shape with an average size (the height is not more - 2 :« barrier » reefs. They have large volumes (up than 7-8 m with the diameter up to 15-20 m) the fra­ to 15 m thick and several hundred metres long), built mework of these bodies is built by cloddy masses, of big nodules of the calcareous algae and, rarely, consisting of skeletons of colonial hexacorals, hydro- colonial corals. Here, in the reef rocks, lenses of the zoans and calcareous algae. The interstices between coarse-grained sandstones and « pockets » saturated bodies of the reef-forming organisms are filled with by the quartz pebble are frequent. One can note, loose calcareo-argillaceous masses. Therefore the cha­ among reef-dwellers (besides above mentioned oys­ racteristic tube-like forms appear in the process of ters), Prohinnites, Trichites, Heterodiceras and also weathering of these bioherms. Small oysters, brachio- nerineid gastropods and so on. The locality on the pods (Craniscus sp., Ismenia cf. perilustrus Sm ir ­ Chuku Mountain is the example of such « barrier » n o v a and others), calcareous sponges, bryozoans, reef. Rare columnals and calyxes of Heberticrinus small solitary corals, serpulids and others organisms heberti were found here ; also are among the reef-dwellers. Traces of boring — 628 —

Name of species L o cality No. on f i g . l Age Skeleton parts No. c o ll.

T R Cd Br С Ad

Thiolliericrinus belbekensis Belbek 3 В 2 8 2 1 11 -_ OK-85 Thiolliericrinus elongatus Belbek 3 В 2 1 ----- CX-98

Thiolliericrinus torosus Belbek 3 в 2 2 2 1 5 . -- CK-86

B urdigalocrinus maxiinus Rodnoie 2 в 2 1 _ 3 _ 4 CK-76,78 Belbek 3 в 2 1 3 1 - 19 - CK-81

Loriolicrinus asper Belbek 3 в 2 _ __ _ 68 CK-99 K aratlykh 4 в 2 3 1 5 12 27 - CK-82 Loriolicrinus laevis Rodnoie 2 в 2 4 _ 1 С К-80 Belbek 3 в 2 5 2 11 _ CK-90 B airakly b в 2 - - 1 -_ _ *CK-50 Tanass 10 в 1 -- 1 - -- *СК-18 Loriolicrinus perforatus Belbek 3 в 2 2 _ СХ-91 Karatlykh 4 в 2 1 ----- СК-95 Conoideocrinus conoideus Belbek 3 в 2 1 ----- СК-97

Umbocrinus urabonatus Rodnoie 2 в 2 1 СК-75 Belbek 3 в 2 27 _ 16 8 53 - СК-83,89 K aratlykh 4 в 2 4 - 3 - -- СК-94 Heberticrinus heberti Rodnoie 2 в 2 3 __ b 2 _ СК-61 Belbek 3 в 2 72 1 26 215 163 _ СК-51 Karatlykh 4 в 2 5 - 2 36 10 - СК-93 Chuku 5 в 2 6 - 1 1 9 - СХ-92 Tanass 10 в 1 - 1 1 --- * ск-зо Tanass 10 в 1 1 -- --- СК-69 Heberticrinus aff.heberti Belbek 3 в 2 3 - 5 - - - ск-эб

Heberticrinus remesi Beshterek 8 V 1 4 --- 9 - CK-5b,57

Heberti^* j-jius irregularis Rodnoie 2 в 2 5 ___ _ СК-74 Belbek 3 в 2 7 - 12 2 27 - СК-88 Tanass 10 в 1 2 - 1 - 1 - * СК-22,25 Tniolliericrinidae indet. Kuchki i в 2 __ 2 4 3 СК-64 Rodnoie 2 в 2 _- 1 5 39 СК-73 Belbek 3 в 2 - -- 269 297 15 СК-ЮО K aratlykh 4 в 2 - - - 45 11 3 СК-101 Ivlrarnornoie 7 в 1 __- 3 2 - СК-6ь Mezhgorie 9 в 2 _ --- 1 - * СК-37 Tanass 10 в 1 ~ - -- 1 - *СК-31

Tabl. 1 — Materials for 12 species of Thiolliericrinidae from the Lower Cretaceous deposits of the Crimea. The designations in the column « Age » : B1 - Lower Berriasian, B2 - Upper Berriasian, VI - Lower Valanginian ; in the column « Skeleton parts » : T - calyxes, R - radiais, Cd - centr odor sals, Br - brachials, С - columnals, Ad - root formations. The collections handed from Bogdanova & Loba- chiova (1977) are marked by asterisks in the column « Numbers of collections ».

Matériaux pour les 15 espèces de Thiolliericrinidae du Crétacé inférieur de la Crimée. Les indications dans la rubrique « Age » (âge) : B1 - Berriasien inférieur, B2 - Berriasien supérieur, VI - Valanginien inférieur ; dans la rubrique « Skeleton parts » (partie du squelette) : T - coupes dorsales, R - radiales, Cd - centrodorsales, Br - brachiales, С - columnales, Ad - formations racinales. Le matériel récolté par Bogdanova & Lobatchiova (1977) est signalé par des astérisques dans la rubrique « Number of collections » (numéros des collections).

organisms are frequent. The echinoderm remains are « Isocrinus » arzierensis (D e L o r i o l ), Apiocrinites common. Echinoid spines and test debris, starfish and the thecae of the true unstalked comatulids were plates and crinoid fragments were found in the found. « tower-like » bioherms. Columnals, thecae and bra­ chials of several Thiolliericrinidae species were collec­ The study of dozens « tower-like » bioherms shows ted here. Besides, rare stem fragments of that Thiolliericrinidae were distributed unevenly. — 629 —

They avoided settling on « yet-formed » bioherms, of Thiolliericrinidae indet. were found in this region the height of which was not more than 2 m. The (see « Kuchky » on fig. 1 and in table 1) in the orga- remains of the crinoids under discussion are almost nogenous limestones of the middle part of the Berria­ absent on the bioherms placed in the limestones sian stage. These limestones are slightly arenaceous having an arenaceous admixture or with then quart- and built by crinoidal debris (various species of the zous pebbles. Thiolliericrinidae avoided the bio­ genus Apiocrinites, rarely « Isocrinus » arzierensis herms, the main rock-forming component of which (De Loriol), calcareous sponges, oysters, corals was the calcareous algae. Under the same conditions, (solitary and colonial forms), brachiopods (Rioultina the thiolliericrinid remains are more frequent on bio­ robusta Sm irno va, Symphythyris arguenensis Mois- herms having many interstices filled with the argilla­ seev), serpulids etc. ceous mass. The finds of Thiolliericrinidae are accom­ panied, as a rule, by the finds of the bulb-like echi- Conditions of the existence of Thiolliericrinidae in noid spines, the spiny asterozoan plates and the dacty- the central and eastern Crimea are somewhat diffe­ late sponges. rent. For example, in the Tass-Kor Ravin (« Mramor- noie » - see table 1 and fig. 1), the Lower Berriasian The remains of the Thiolliericrinidae are found clays occur on the washed out surface of the marbled more often in the inner, more loose core of a bio- Tithonian limestones and contain small bioherms (the herm. However it does not mean that they are absent thickness is up to 1 m and the diameter is 2 to 3 m). on the external walls. Moreover, rare columnals and The main reef-building organisms are various calca­ thecae are also found outside the bioherm, in the sur­ reous sponges. Solitary and colonial corals and calca­ rounding rocks. But, in this case, they are often reous algae are scarce. The dwellers of these tiny reefs strongly rounded. However, the remains of one spe­ are small oysters, brachiopods, tubicolous worms and cies are found only in the bedded microphytolithic regular echinoids with large thorn-like spines. Cri­ limestones. This is Conoideocrinus conoideus. noids are rare. They же Apiocrinites sp., Eugeniacri- Heberticrinus heberti is a « background » species nites sp. and Isocrinus sp. (only columnals). The stem everywhere. But at almost every bioherm there is a joints of Thiolliericrinidae are found here. Occur­ characteristic species, distributed locally. Such is rence of crinoids in the Berriasian deposits of the Thiolliericrinus belbekensis in the « Belbek » locality Mramornoie was published by Lyssenko & Vakhrus­ (see fig. 1 and table 1), Loriolicrinus asper on the hev (1974, p. 149). Karatlykh Mountain, Heberticrinus irregularis in the In the villages Alekseevka and Krasnossiolovka Rodnoie environs etc. However frequently H. heberti (« Tanass » - see fig. 1 and table 1), the Thiolliericri­ is the species which only represents the family on a nidae are found in the small bioherms among the bioherm. Lower Berriasian clays and sandstones, which occur Occurrence of the Thiolliericrinidae in the Belbek on the washed out and extremely rough surface of the Valley was already published by Arendt (1974, p. 57), Tithonian limestones. These bioherms form an almost Arendt & Yanin (1964, p. 140). unbroken horizon, but reach the greatest develop- Towards the West of the Belbek River, in the Chor­ menl on the Tithonian substratum swells probably naya Basin (see « Rodnoie » in the table 1), the being the initial points of its forming. The bioherms remains of the late Berriasian thiolliericrinids were are built by the rolled debris of the basement rocks found in the paleofacies analogous to the described and by the calcareous alga pellets. Among reef- above. However, hydrozoans are the main reef- dwellers, there are rare corals, oysters, calcareous building organisms here (personal communication of sponges, sea-urchins and sea-lilies (Thiolliericrinidae I.Y. Bugrova), and rudists are usual among reef- indet. and Cyrtocrinida indet.). dwellers (Yanin 1985, p. 25). Besides, some columnals

3 — TERMINOLOGY

A special terminology, in regard to skeletal elements Ad - attachment (terminal disk) ; of the Thiolliericrinidae, is used in the following lAx - primaxillar (the axillar of the first brachial chapters. The principal designations are as below (fig. series) ; 2). IIAx - secondaxillar (the axillar of the second bra­ chial series) ; 630 — — 631 —

В - basal plate (the basal) ; SI - left second brachial series ; Be - centrodorsal basis ; Sm - marginal crenulation ; IIBr - secondbrachial (the plates of the second Sr - right secondbrachial series ; bra chial series) ; St - crenularium on the transversal (fulcral) ridge ; С - stem joint (columnal) ; The measurements used for the description of spe - Cd - centrodorsal plate (the centrodorsal) ; cies are (see fig. 2) : Co - opening of the axial canal ; Af - angle of the inclination of a radial facet ; Ct - cavity of the calyx ; Bf - width of a radial facet ; F - fossa on the centrodorsal ; Dc - diameter of a centrodorsal basis ; Fi - interradial furrow in the theca cavity ; Dt - diameter of a theca ; FI - lateral fossa ; He - height of a centrodorsal ; Fm - marginal fossa ; Hr - height of the external (free) radial face ; Fr - radial furrow in the theca cavity ; Ht - height of a theca ; R - radial plate (the radial) ; Lf - length of a radial facet ; Rf - radial facet (the upper face of the radial) ; Ri - intermediate ridge ; In some cases, to show the morphological features Rm - marginal ridge ; more clearly, the relative parameters are used : Rs - external (free) face of the radial ; Hr : Ht - relative height of the external radial face ; Rt - transversal (fulcral) ridge ; Bf : Lf - relative width of the radial facet. Sc - cirrus socket ;

4 — TAXONOMIC DESCRIPTION

Class Crinoidea brachial is axillary. Immovable junctions are absent in the arms. The branching of the arms is isotomous Subclass Articulata Miller, 1821 (sometimes slightly asymmetrical) and not less than Order COMATULIDA A.H. Clark , 1908 thrice-repeated. Secondbrachials and tertibrachials are closed, forming a massive tube. The centrodorsal Family Thiolliericrinidae A.H. Clark , 1908 with or without cirri. The short stem consists of seve­ ral columnals. The columnal articula are bifascial Type genus : (synartrial). The attachment disk has a shape of the cone-like knot. Thiolliericrinus Et a llo n , 1859.

Generic features : Exposure of the generic features Diagnosis : for the family meets a considerable difficulty. The The theca consists of five radiais, five basais (often fact is that the representatives of this group, just as invisible outside) and one centrodorsal. The first most reef-dwellers, have many variations concealing

Fig. 2 — Main elements of the Thiolliericrinidae skeleton. A - calyx without cirrus sockets on the centrodorsal (side view) ; В - calyx with fossae and cirrus sockets on the centrodorsal (side view) ; С - separate radial (view from above) ; D - calyx (view from below) ; E - calyx (view from above) ; F - relative partial recons­ truction of the skeleton ; G - cruciate columnal (the arrows mark the direction of the fulcral ridge) ; H - parallel columnal (the arrows as on fig. G) ; I - articulum of a columnal. For designations see the text. Éléments principaux du squelette des Thiolliericrinidae. A - coupe dorsale sans surface d’articulation des cirres sur la centrodorsale (vue latérale) ; В - coupe dorsale avec les fossettes et les surfaces d’articulation des cirres sur la centrodorsale (vue latérale) ; С - radiale isolée (vue en dessus) ; D - coupe dorsale (vue en dessous) ; E - coupe dorsale (vue en dessus) ; F - reconstruction partielle conventionnelle du squelette ; G - columnale à crêtes ful- crales croisées (les flèches marquent la direction de la crête fulcrale) ; H - columnale à crêtes fulcrales parallèles (les flèches comme la fig. G) ; I - facette articulaire d’un article. Explications dans le texte. — 632 — not only specific but also generic limits. However, one an oval centrodorsal can be found, lenghtening of can determine the main generic features, being guided which coincides with a direction of the fulcral ridge Dy phylogenetical considerations (see below), as fol­ on its basis. Sometimes the rounded-pentangular out­ lows : line is marked in large specimens. 1 - The degree of the development of cirrus sockets The stem facet can be greatly deepened on the lower on a centrodorsal, namely : a - they are functional centrodorsal face of H. heberti (this is typical for and are distributed regularly (Solonaerium) or irregu­ rather low calyxes), can be flat (in low calyxes) or can larly (Thiolliericrinus) ; b - there are no true sockets, project as an acute ridge (in high calyxes). The centro­ but structureless fossae scattered over the centro­ dorsal basis is mostly horizontal but it can be placed dorsal (Burdigalocrinus) or remained only on the obliquely with the calyx axis. Because of this, the cen­ suture between radiais and the centrodorsal (Umbo- trodorsal can be considerably higher on one side than crinus, partly Loriolicrinus) ; the calyx is deprived of on another. any traces of cirri (Conoideocrinus, Heberticrinus) ; The radial facets of H. heberti more often adjoin 2 - The degree of the development of radiais, directly the centrodorsal. But in some calyxes, one namely : radiais have a high external face and nearly can see a narrow strip of the free radial surface divi­ horizontal muscular facets (Solonaerium, Burdigalo­ ding the facet from the centrodorsal. The width of crinus, Conoideocrinus, Loriolicrinus, partly Thiol­ this strip can varied, though in a small degree, even on liericrinus) ; radiais are low, their facets are strongly different sides of the same specimen. inclined outside and bordered close to a centrodorsal (Heberticrinus, Umbocrinus, partly Thiolliericrinus) ; The size of basal plates is also subjected to a varia­ 3 - The degree of the development of basais, tion. Being concealed under the radiais, they never namely : a - basais are large (Solonaerium, Conoideo­ have an equal length. Some basais, even in one calyx, crinus, partly Thiolliericrinus and Burdigalocrinus) ; hardly reach a half of the distance from the centre to b - basais are small, hardly visible or not visible on the the outer edge while the others can appear upon the external calyx surface (Loriolicrinus, Umbocrinus external calyx surface. These variations are marked in and Heberticrinus). Heberticrinus heberti and Burdigalocrinus maximus. The basais of Umbocrinus umbonatus are quite visi­ Other features (size ratio of the radial ring and the ble, as a rule, on the calyx surface. However, the cases centro-dorsal, the stem facet construction etc.) are were marked, when some basais of individual speci­ present in the generic diagnoses given below besides mens are concealed under the radiais. the enumerated characteristics. These features, howe­ ver, have no importance. And what is more, the The sculpture of the external calyx surface is the above mentioned generic features, can neither separa­ relatively insignificantly variable feature. For exam­ tely, determine a genus in the family Thiolliericrini­ ple, the calyxes of Thiolliericrinus belbekensis are dae. However, they show quite clearly the differences covered with small granules and the radiais of T. toro- between the genera being presented as a table (table sus bear big knobs. However, a weak sculpture in the 2). form of small irregular granules is observed in some specimens of Umbocrinus umbonatus, for which the Intraspecific variation. The form and relative size smooth calyxes are typical. of the centrodorsal plate are the most variable featu­ res of Thiolliericrinidae. These variations are marked One must emphasize the difference between the mainly in cirriless forms. For example Heberticrinus intraspecific and the intrapopulational variations. heberti may have the centrodorsal height less than The specimens which belong to the same species are that of the radial ring, equal to it or several times relatively poorly differ in each of the local popula­ greater. Besides, the place of the greatest diameter is tion. At the same time, one can hardly found the situated in the base of the low centrodorsal or in the forms which would be united under the same specific upper part of high one, directly below the radial ring. name even the two bioherms are placed side by side. The centrodorsal diameter can be equal to that of radial ring among specimens with an identical size but Seven genera : Solonaerium Éta llo n , 1862 emend. the first diameter can exceed almost two times the Rasm ussen, 1978, Thiolliericrinus Éta llo n , 1859, second one. Heberticrinus heberti has a commonly Burdigalocrinus Jaekel, 1918, Loriolicrinus Jaekel, round centrodorsal outline. However, specimens with 1918, Conoideocrinus nov. gen., Heberticrinus nov. — 633 —

gen. and Umbocrinus nov. gen. Each of them is deter­ p. 327, fig. 443 g ; 1978, p. 360, fig. 432 g - sub Bala- mined by peculiarities of the calyx construction. nocrinus, lapsus calami). However, there are some species, belonging evidently to the family Thiolliericrinidae, for which only the Besides these above mentioned species, there is ano­ ther one which was placed by many authors in the columnals are known. These forms, mentioned family Thiolliericrinidae (De Loriol 1879, p. 195 ; usually under the generic name Apiocrinites or Bour- 1880, p. 11 ; 1889, p. 559 ; Carpenter 1884, p. 257 ; gueticrinus, are placed below. Their taxonomic posi­ Gislén 1924, p. 186 ; Roux 1978a, p. 226 ; 1978b, p. tion needs a more accurate definition. A 21 ; Rasmussen 1978c, p. 879) - namely Bourgueti­ crinus oôliticus. The available morphological and hlajue of genus 3c PВ Hr: Ht Bf :Lf Af° geological data (M’Coy 1848, p. 405 ; 1854, p. 53), in my opinion, testify against that interpretation of the Solonaerium + - + 0,24 0,54 38 species. Thiolliericrinus + + + С, 20 0,42 42 Upper Jurassic (Callovian) to Lower Cretaceous Burdigalocrinus - + + 0,35 0,39 40 (Hauterivian) ; Portugal, France, Switzerland, Ger­

Umbocrinus - + + 0,17 0,56 50 many, Rumania, Czechoslovakia, Crimea, Caucasus, Pamyrs. Loriolicrinus - + + 0,35 0,57 37

Oonoideocrinus - - + 0, bl 0,35 26

Heberticrinus - - + 0,12 0,47 42 Genus Solonaerium Étallo n, 1862 emend. Rasm ussen, 1978 Tabl. 2 — Main diagnostic features of the genera of the family Thiolliericrinidae. The designations (see fig. 2) : Sc - cir­ 1862 — Solonaerium Étallon (in Thurm ann & rus sockets (yes - no), F - fossae on the centrodorsal (yes - Étallon), p. 341 (errore typographico ex no), В - basais (visible on the calyx surface or not visi­ Solanocrinus). ble), Hr :Ht - average relative height of the free radial surface, Bf : Lf - average relative width of the radial 1978c — Solonaerium Étallon : Rasmussen, p. facet, Af° - average angle of the inclination of the radial facet (in degrees). 881. Caractères principaux de détermination des genres de la famille Thiolliericrinidae. Légende (voir fig. 2) : Sc Type species : zone d’insertion des cirres (présence - absence), F - fos­ ses sur la centr odor sale (présence - absence), В - basales Comatula sigillata Quenstedt, 1876. (visibles sur la surface de la coupe dorsale ou non visi­ bles), Hr : Ht - hauteur relative moyenne de la surface Diagnosis : libre de la radiale, Bf : Lf - largeur relative moyenne de la facette radiale, Af° - angle moyen de l’inclinaison de la facette radiale (en degrés). Radiais have a high free external surface. Basais are visible on the calyx surface. Cirrus sockets are of the same size in regular vertical rows. There are no fossae under radiais. Centrodorsal is high and conical. Stem facet is small, deepened, pentangular or rounded. Apiocrinites flexuosus G o l d f u s s (1826-1833, p. 186, pl. 57, fig. 4) (= Bourgueticrinus flexuosus One species. d’ORBlGNY, 1840, p. 96-98, pl. 17, fig. 13-15) - Kim- Solonaerium sigillatum (Quenstedt, 1876, p. 176, meridgian of Germany (see also : Quenstedt 1852, p. pi. 96, fig. 49,50 sub Comatula ( = Solanocrinites cos­ 612, pi. 53, fig. 17 ; 1876, p. 367, pi. 104, fig. 57, 58 ; tatus Go ldfuss, 1826-1833, p. 167 ex parte : pi. 50, Gislén 1924, p. 186). fig. 2 ; non pi. 50, fig. 7 a,b = type specimen for Solanocrinites costatus ; nec fig. 7 c,d = Solanocrini- Bourgueticrinus oosteri D e L o r io l (1879, p. 188, pi. 18, fig. 6,7) (= Bourgueticrinus flexuosus Oos- tes orbignyi ; necdum fig. 7 a,f = ? Solanocrinites costatus) (Solanocrinus costatus Étallon 1862, p. t e r , 1871, p. 130, 142, 149, pi. 19, fig. 1 and 23) - Lower Neocomian of Switzerland (see also : Rasmus­ 222 ; Solonaerium costatus Thurmann & Étallon sen 1961, p. 168). 1862, p. 341 ; Solanocrinus sigillatus Schliiter 1878, p .37 ; Comatula sigillata De Loriol 1879, p. 267 ; Bourgueticrinus sp. (Quenstedt 1876, p. 367, 431, Antedon sigillata Carpenter 1881a, p. 201, 202 ; pi. 106, fig. 43, 44) - Callovian of Germany. Solonaerium costatum Rasmussen 1978c, p. 881, fig. Bourgueticrinus sp. (Termier & Termier 1949, p. 57, 588-3). Kimmeridgian ; Nattheim (Württemberg), 86, pi. 6, fig. 8-14) - Upper Jurassic of Morocco (see Germany (fig. 3 A,B). also : Ubaghs 1953, p. 716, fig. 101 and Müller 1963, Upper Jurassic (Kimmeridgian) ; Germany. — 634 —

Fig. 3 — Calyxes and centrodorsals of Solonaerium and Thiolliericrinus (x 3). A - holotype of S. sigillatum (calyx from side and below, after Goldfuss 1826-1833, pi. 51, fig. 2 and Rasmussen 1978c, fig. 588-3). В - S. sigillatum (calyx from side and below, after Quenstedt 1876, pi. 96, fig. 49). С - holotype of T. arzierensis (centrodorsal from side and below, after De Loriol 1879, pi. 20, fig. 20 and Rasmussen 1961, pi. 35, fig. 13. D - lectotype of T. favieri (calyx from side, after De Loriol 1889, pi. 229, fig. 2). E - reconstruction by holotype of T. ribeiroi (calyx from side and below, after De Loriol 1880, pl. 1, fig. 3 ; 1891, pi. 29, fig. 16, 18 and Bather 1900, fig. 117-2).

Coupes dorsales et centrodorsales de Solonaerium et Thiolliericrinus. A - holotype de S. sigillatum (coupe dorsale vue de côté et en dessous, d’après Goldfuss et Rasmussen). В - S. sigillatum (coupe dorsale vue de côté et en dessous, d’après Quenstedt). С - holotype de T. arzierensis (centrodorsale vue de côté et en dessous, d’après De Loriol et Rasmussen). D - lectotype de T. favieri (coupe dorsale vue de côté, d’après De Loriol). E - reconstruction d’après l’holotype de T. ribeiroi (coupe dorsale vue de côté et en dessous, d’après De Loriol et Bather).

Genus Thiolliericrinus É t a l l o n , 1859 1889 — Thiolliericrinus Étallon : De Loriol, p. 431, 542.

1857 — Thiolliericrinus : Éta llo n , p. 301 (nomen 1900 — Thiolliericrinus Étallon : Bather p. 195. pro visorium). 1913 — Thiolliericrinus Étallon : A.H. Clark, p. 1859 — Thiolliericrinus Étallon : Étallon, p. 413, 234. 445. 1918 — Thiolliericrinus : Jaekel, p .71. 1879 — Thiolliericrinus Étallon : De Loriol, p. 1924 — Thiolliericrinus Étallon : Gislén, p. 186. 193. 1933 — Thiolliericrinus Étallon : Dacqué, p. 103. 1880 — Thiolliericrinus Étallon : De Loriol, p. 10. 1953 — Thiolliericrinus Étallon : Sieverts-Doreck 1881b — Thiolliericrinus Étallon : Carpenter, p. (in Ubaghs), p. 758. 377. 1961 — Thiolliericrinus Étallon : Rasmussen, p. 1883 — Thiolliericrinus Étallon : De Loriol, p. 65. 212. 1975 — Thiolliericrinus Étallon : Hess, p .69. (Thiolliericrinus ribeiroi : Carpenter 1881b, p. 377 ; 1978a — Thiolliericrinus Étallon : Roux, p. 234. De Loriol 1889, p. 557, 558, pi. 229, fig. 9 ; De Loriol 1891, p. 165 ex parte, pi. 29, fig. 16, 18-22, non fig. 17 1978b — Thiolliericrinus Étallon : Roux, p. A 20. = Burdigalocrinus lorioli ; Bather 1900, p. 195, fig. 1978c — Thiolliericrinus Étallon : Rasmussen, p. 117-2 ; Jaekel 1918, p. 71, fig. 63 ; Gislén 1924, p. 879. 187 ; Ubaghs 1953, p. 718, fig. 103 ; 1978, p. 72, fig. 52-1 ; Rasmussen 1978c, p. 880, fig. 588-2 g-h ; Roux Type species : 1978a, p. 234). Lusitanian ; Engenheiro (Ribatejo), Portugal (fig. 3 E). Humberticrinus favieri De Loriol ex Éta llo n, 1879.

Diagnosis : Thiolliericrinus belbekensis nov. sp. Radiais with either high or low free external sur­ fig. 4 A ,В ; 5 A-F face. Basais are visible or not on the calyx surface. The cirrus sockets have different size and are placed D e r iv a t io n o m i n is : irregularly. Some of the sockets are turned into struc­ tureless fossae. The centrodorsal is conical, high or The name of the species comes from the type loca­ low. The stem facet is small, flat or convex. lity. Six species : T. arzierensis De Loriol, T. favieri H o l o t y p e : (De Loriol ex Étallon), T. ribeiroi De l o r io l, T. belbekensis nov. sp., T. elongatus nov. sp., T. torosus CK-85-1 (fig. 5 A) ; Leningrad Mining Institute. nov. sp.

Upper Jurassic (Oxfordian) - Lower Cretaceous T y p e l o c a l it y : (Valanginian) ; Portugal, France, Switzerland, Cri­ mea. Kuibyshevo, Belbek Valley, Crimea. Thiolliericrinus arzierensis De Lo riol, 1889 (p. 560) T y p e l e v e l : (= Antedon valdensis De Loriol 1879, p. 266 ex parte : pi. 20, fig. 20, non fig. 19, 32, 33 = Solanocri- Upper Berriasian. nites valdensis) (Thiolliericrinus arzierensis : Gislén 1924, p. 187 ; Rasmussen 1961, p. 212, pi. 35, fig. 13 ; Hess 1975, p. 69, pi. 20, fig. 13). Valanginian ; D e s c r i p t i o n : Arzier (Vaud), Switzerland (fig. 3 C). The calyx is low, subcylindrical with the rounded = Thiolliericrinus favieri (De Loriol ex Étallon, pentangular outline. The basais are not visible on the 1879, p. 194, pi. 18, fig. 8-10 sub Thiolliericrinus external calyx surface. But small triangular plates are flexuosus (= Thiolliericrinus sp. É tallon, 1857, p. observed in some specimens in some interradia. 301 ; = Thiolliericrinus flexuosus : Étallon 1859, p. Basais are rod-like, narrowed towards the outer mar­ 445 ; Étallon 1862, p. 222, 237 ; De Loriol 1889, p. gin and risen abruptly inside the calyx. The external 553 ex parte, pi. 229, fig. 2,4,5,7,8, non fig. 3,6 = surface of the radiais is low, vertical and slightly con­ Loriolicrinus sp ; Kirk 1912, p. 75, pi. 5, fig. 4 ; Ras­ vex. It is covered with coarse granules. The radial mussen 1978c, p. 879, fig. 588-2b,c,d non fig. 588-2a facets are wide. The furrow-like depression is placed = Loriolicrinus sp. ; = Humberticrinus favieri under each radial facet on the external radial face. É tallon : De Loriol 1879, p. 195 ; 1889, p. 556 ; The calyx cavity is broad and shallow. 10 furrows lean 'Thiolliericrinus favieri : Gislén 1924, p. 187 ; Roux between the radiais and are fully pronounced on their 1978b, p. A 20 ; non Thiolliericrinus favieri : Ubaghs middle part. The inner radial surface is covered with 1953, p. 718, fig. 104 ; 1978, p. 72, fig. 52-2 = Thiol­ weak furrows. The basais are visible on the bottom of liericrinus sp. Bather 1900, p. 195, fig. 117-1). Kim- the calyx cavity only in eroded specimens, but are meridgian (Dicératien) . Valfin (Jura), France (fig. 3 commonly covered with a porous calcitic plug. The D). primaxillary has a strongly convex uneven external Thiolliericrinus ribeiroi De lo rio l, 1880 (pi. 11, surface. Its lower facet is muscular. The second axil­ fig. 1, fig. 3-11) (= Thiolliericrinus choffati De lary is very asymmetric with the muscular lower facet loriol, 1889, p. 547, 556, nomen provisorium) and with a strongly convex granulated external — 636 —

Fig. 4 A scheme of the disposition of the cirrus sockets and the fossae on centrodorsals of some Thiolliericrinus (x 4). The arrows mark the ends of the fulcral ridge. Schéma de la disposition des fossettes et des surfaces d’articulation des cirres sur les centrodorsales de quelques Thiolliericrinus. Les flèches marquent la direction de la crête fulcrale. A - T. belbekensis (holotype CK-85-1). В - T. belbekensis (CK-85-9). С - T. elongatus (holotype CK-98-1). D - T. torosus (holotype CK-86-1). — 637 —

surface. The height of the centrodorsal is about equal C o m p a r i s o n : to the height of the radial ring. 2 to 3 cirrus sockets T. belbekensis resembles T. favieri after its centro­ are under each the radiais. 11 to 12 sockets in all. dorsal construction (one row of sockets, roughly two Some of the sockets are replaced by structureless fos­ sae. The free surface of the centrodorsal between the under each of the radiais). The species under discus­ cirrus sockets is granulated like that of the external sion differs from T. favieri having relatively low radial surface. The lower centrodorsal face, in all exa­ radiais, a granulated external surface and fossae on its mined specimens, is poorly preserved. It is convex, centrodorsal. ovate-oblong and has a fulcral ridge on whose both sides two slit-like furrows are marked, surrounding in M a t e r i a l s : their turn by two marginal ridges. 8 calyxes, 2 radiais, 1 centrodorsal and 11 brachials (see table 1). D im e n s io n s : Ht = 5.5 - 8.4 mm ; Dt = 7.8-11.8 mm ; Dc = 5.3 D istribution : - 9.5 mm ; Af = 50° ; Hr : Ht = 0.10 - 0.18 ; Bf : Lf = 0.41 - 0.45. Berriasian ; Crimea.

Fig. 5 — Calyxes and brachials of Thiolliericrinus belbekensis (x 3). A - holotype CK-85-1 (calyx from side and below). В - CK-85-9 (centrodorsal with two basais from above). С - CK-85-10 (primaxillary from above and below). D - CK-85-13 (secondaxillary from above). E - CK-85-14 (secondaxillary from below). F - CK-85-17 (tertibrachial from below). Coupes dorsales et brachiales de Thiolliericrinus belbekensis. A - holotype CK-85-1 (coupe dorsale vue de côté et en dessous). В - CK-85-9 (centrodorsale avec deux basales, vue en dessus). С - CK-85-10 (primaxillaire, vue en dessus et en dessous). D - CK-85-13 (secundaxillaire, vue en dessus). E - CK-85-14 (secundaxillaire, vue en dessous). F - CK-85-17 (tertibrachiale, vue en dessous). — 638 —

Thiolliericrinus elongatus nov. sp. D e s c r i p t i o n : fig. 4 С ; 6 A The calyx is high, conical, narrowed downwards with the rounded-pentangular outline. Basais are visi­ ble on the external calyx surface as large triangular or D e r iv a t io n o m i n is : irregular diamond = shaped plates. The external The name of the species comes from « elongatus » radial surface is very high, sloping, flat and smooth. (elongated in Latin). The radial facets are narrow. The calyx cavity is broad and shallow. 10 furrows lead between the radiais also in their middle part are pronounced only H o l o t y p e : in the upper. The interradial furrows are wider and CK-98-1 (fig. 6 A) ; Leningrad Mining Institute. deeper. The inner radial surface is covered with bent furrows. The basais are invisible on the bottom of the calyx cavity. The centrodorsal is lower than the radial T y p e l o c a l it y : ring. 1 to 2 large, projected cirrus sockets are situated Kuibyshevo, Belbek Valley, Crimea. under each the radiais. 10 sockets in all. The centro­ dorsal surface is smooth. The lower face of the cen­ trodorsal is poorly preserved. It is convex, ovate- T y p e l e v e l : oblong and has a fulcral ridge on which both sides Upper Berriasian. there are two slit-like furrows surrounded by a weak marginal ridge. Fig. 6 — Skeletal parts of Thiolliericrinus (x 3). A - T. elongatus, holotype CK-98-1 (calyx from side and below). В - T. torosus, holotype CK-86-1 (calyx from side and below). С - T. toro­ sus, CK-86-6 (secon- daxillary from below). D - T. torosus, CK-86- 5 (radial from above). E - T. torosus ?, CK- 86-11 (cirral). в Éléments du squelette de Thiolliericrinus. A - T. elongatus, holotype CK-98-1 (coupe dorsale vue de côté et en dessous). В - T. torosus, holotype CK-86-1 (coupe dor­ sale vue de côté et en dessous). С - T. toro­ sus, CK-86-6 (seconde axillaire vue en des­ sous). D - T. torosus, CK-86-5 (radiale vue en dessus). E - T. torosus ?, CK-86-11 (cirrale).

D i m e n s io n s : genus, having a very high external radial surface and a narrow centrodorsal basis. Ht = 9.3 mm ; Dt = 12.4 mm ; De = 6.4 mm ; Af = 45° ; Hr : Ht = 0.37 - 0.39 ; Bf : Lf : = M a t e r i a l s : 0.36-0.37. One calyx (see table 1).

C o m p a r i s o n : D istribution : T. elongatus is similar with T. rïbeiroi but differs from this species, like from other species of that Berriasian ; Crimea. — 639 —

Thiolliericrinus t orosus nov. sp. M a t e r i a l s : figs. 4 D ; 6 B-E 2 calyxes, 1 centrodorsal, 2 radiais, 5 brachials, 4 cir- rals (see table 1). D e r iv a t io n o m i n i s : The name of the species comes from « torosus » D istribution : (pustulous in Latin). Berriasian ; Crimea.

H o l o t y p e : Genus Burdigalocrinus Ja e k e l , 1918 CK-86-1 (fig. 6 B) ; Leningrad Mining Institute.

1918 — Burdigalocrinus : Ja e k e l , p.71. Ty p e l o c a l it y : 1924 — Burdigalocrinus Ja e k e l : Gislén, p. 188. Kuibyshevo, Belbek Valley, Crimea. 1933 — Burdigalocrinus Ja e k e l : Dacqué, p. 103. 1953 — Burdigalocrinus Ja e k e l : Sieverts-Doreck Ty p e l e v e l : (in Ubaghs), p. 758. Upper Berriasian. 1961 — Burdigalocrinus Ja e k e l : Rasmussen, p. 213. T y p e l e v e l : 1978a —Burdigalocrinus Ja e k e l : Roux, p. 234. Upper Berriasian. 1978b —Burdigalocrinus Ja e k e l : Roux, p. A 20. 1978c —Burdigalocrinus Ja e k e l : Rasmussen, p. D e s c r ip t io n : 880. The calyx is low, conical, abruptly narrowed down­ wards with the rounded-pentangular outline. The T y p e s p e c ie s : basais are visible on the calyx surface as small convex Burdigalocrinus lorioli Ja e k e l , 1918. triangular plates. The external radial surface is very low, overlapping the centrodorsal, and covered with D i a g n o s i s : big irregular knobs. The calyx cavity is broad and The radiais have high free external surface. The shallow. 10 furrows go between the radiais and are basais are visible on the calyx surface. Cirrus sockets fully pronounced on their middle part. The inner are absent but there are structureless fossae placed, as radial surface is slightly striated. The basais are not a rule, regularly. The centrodorsal is high and conical. visible on the bottom of the calyx cavity. The pri- The stem facet is wide, elliptical and flat. maxillary has a muscular lower surface. Its external Two species : B. lorioli Ja e k e l , B. maximus nov. surface is tumid, cone-likely and covered with big sp. knobs. The height of the centrodorsal is about equal to the radial height, but the diameter of the centro­ Upper Jurassic (Oxfordian) - Lower Cretaceous dorsal is notably smaller than that of the radial ring. (Berriasian) ; Portugal, Crimea. Three cirrus sockets are placed close to each other Burdigalocrinus lorioli Ja e k e l , 1918 (p. 72, fig. 64 under each radial. Two large sockets are situated nea­ D,E) (= Thiolliericrinus ribeiroi D e L o r i o l , 1891, p. rer to the radial and a small one is placed below, bet­ 165 ex parte, pi. 29, fig. 17, non fig. 16, 18-22 = ween the large ones. 15 sockets in all. The lower cen­ Thiolliericrinus ribeiroi) (Burdigalocrinus lorioli : trodorsal face is convex, ovate = oblong and has a Gislén 1924, p. 190 ; Rasmussen 1961, p. 213 ; 1978c, fulcral ridge with two narrow interrupted depressions p. 881, fig. 588-4 ; Roux 1978b, p. A 20). Lusitanian ; on both sides, surrounded by a weak marginal ridge. Engenheiro (Ribatejo), Portugal (fig. 7 A).

D im e n s io n s : Burdigalocrinus maximus nov. sp. Ht = 7.2 mm ; Dt = 10.8 mm ; Dc = 6.0 mm ; Af = 57° ; Hr : Ht = 0.07 - 0.11 ; Bf : Lf = 0.44 - 0.55. fig. 7 B,C ; pl. 1, fig. 1-3

D e r iv a t io n o m i n is : Co m p a r is o n : The name of the species comes from « maximus » The centrodorsal of T. torosus is similar to that of (large in Latin). T. arzierensis but has a considerably greater number of the cirrus sockets (15 againts 6-8) being placed H o l o t y p e : regularly. CK-78-1 (fig. 7 B) ; Leningrad Mining Institute. — 640 —

Fig. 7 — Calyxes and centrodorsals of Burdigalocrinus (x 3). A - holotype of B. lorioli (centrodorsal from above, side and below, after De Loriol 1891, pi. 29, fig. 17). В - В. maximus, CK-78-1 (calyx from above and side). С - В. maximus, holotype CK-81-1 (calyx from above and side ; three radiais are lost). Coupes dorsales et centrodorsales de Burdigalocrinus. A - holotype of B. lorioli (centrodorsale vue en dessus, de côté et en dessous, d’après De Loriol). В - B. maximus, CK-78-1 (coupe dorsale vue en dessus et de côté). С - B. maximus, holotype CK-81-1 (coupe dorsale vue en dessus et de côté ; trois radiales sont absentes).

T y p e l o c a l it y : The radial and interradial furrows are weak. The inner radial surface is wrinkled. The basais are visible Rodnoie, Chornaya Basin, Crimea. on the bottom of the calyx cavity. The primaxillary is low with a muscular lower face. The external surface T y p e l e v e l : of the primaxillary is slightly convex and smooth. The Upper Berriasian. height of the centrodorsal is about equal to that of the radial ring but sometimes unequal on opposite sides of the calyx. 2-4 rounded or high-oval fossae are pla­ D e s c r i p t i o n : ced on the centrodorsal under each radial. The lower The calyx is high, broad-conical, rounded- centrodorsal face bears a wide-elliptical flat articular pentangular. The basais are visible on the external face. The fulcral ridge is pronounced. The lateral fos­ calyx surface as small convex triangular plates but sae are shallow and flat. The fulcral ridge is serrated sometimes not in all interradia. The external surface in some columnals. The marginal ridge is often twin­ of the radiais is high, convex or flattened and smooth. ned. The ridges are almost perpendicular to each Two weak fossae are on the upper part of the external other on two sides of one columnal. They are someti­ radial surface under the radial facet. The radial facets mes parallel or turned at an acute angle. are narrow. The calyx cavity is broad and shallow. — 641 —

Dimensions : Loriolicrinus sp. (= Thiolliericrinus flexuosus De Ht = 10.3 - 11.1 mm ; Dt = 13.5 - 16.3 mm ; De = L o r i o l , 1889, pi. 229, fig. 3,6). Kimmeridgian of 7.2 - 9.8 mm ; Af = 38 - 42° ; Hr : Ht = 0,31 - 0.44 ; France. Bf : Lf = 0.36 - 0.41. Solanocrinites jaegeri Q u e n s t e d t , 1852 (p. 601, pi. 51, fig. 33) (non Solanocrinites jaegeri Goldfuss 1826- Comparison : 1833, p. 168, pi. 50, fig. 9 = Chariocrinus ? jaegeri) (Solanocrinites jaegeri : Quenstedt 1858, p. 723, pi. В. maximus differs from В. lorioli by a massive 88, fig. 12 ; 1876, p. 177, pi. 96, fig. 51 . Schliiter large calyx and a low centrodorsal with the flat basis. 1878, p. 36,37 ; Carpenter 1881a, p. 215, pi. 11, fig. 22 ; De Loriol 1889, p. 559 ; Gislén 1924, p. 110 ; Materials : Thiolliericrinus ? jaegeri Carpenter 1881a, p. 209). Kimmeridgian of Germany. 2 calyxes, 2 centr odor sals, 3 radiais, 2 brachials, 23 Upper Jurassic (Oxfordian) - Lower Cretaceous columnals (see table 1). (Valanginian) ; Portugal, France, Germany, Crimea.

Distribution : Loriolicrinus insuetus (De L o r i o l , 1891, p. 167, pi. 29, fig. 23-29 sub Thiolliericrinus) (Loriolicrinus Berriasian ; Crimea. insuetus : Jaekel 1918, p. 72, fig. 65 ; Gislén 1924, p. 190, 207 . Roux 1978a, p. 234 ; 1978b, p. A 21 ; Ras­ mussen 1978c, p. 881, fig. 588-1). Lusitanian ; Serra das Figueiras (Algarve), Portugal (fig. 8 A). Genus Loriolicrinus Jaek el, 1918

1918 — Loriolicrinus : Ja ek el, p. 72. 1924 — Loriolocrinus Jaekel : Gislén, p. 190, 207. Loriolicrinus asper nov. sp. 1933 — Loriolocrinus Jaekel : Dacqué, p. 104. fig. 8 B-F ; pl. 1, fig. 4,5 1978a — Loriolocrinus Jaekel : Roux, p. 234. D e r iv a t io n o m i n is : 1978b — Loriolicrinus JAEKEL : Roux, p. A 21. The name of the species comes from « asper » 1978c — Loriolicrinus Ja e k e l : Rasmussen, p. 881. (rough, rugget in Latin).

Type species : H o l o t y p e : Thiolliericrinus insuetus De Lo riol, 1891. CK-82-1 (fig. 8 D) ; Leningrad Mining Institute.

Diagnosis : T y p e l o c a l it y : The radiais have a high free external surface. The Karatlykh Mountain, Belbek Valley, Crimea. basais are usually not visible but sometimes they are

on the calyx surface. There are no cirrus sockets on T y p e l e v e l : the centrodorsal but sometimes there are small fossae on the radial-centrodorsal suture. The centrodorsal is Upper Berriasian. low, cylindrical or conical. The stem facet is wide, elliptical, flat or concave. D e s c r i p t i o n : The calyx is low, broad-conical, rounded. The Four species : basais are not visible on the calyx surface. The exter­ nal radial surface is very high, convex and covered L. insuetus (De Loriol), L. asper nov. sp., L. lae- with big irregular knobs and ridges. The radial facets vis nov. sp., L. perforatus nov. sp. are wide. The calyx cavity is narrow and deep. The One can evidently attribute to the genus Loriolicri­ radial and interradial furrows are well pronounced. nus also two following species : The basais are not visible on the bottom of the calyx — 642 — cavity. The primaxillary is high with vertical, somew­ The centro-dorsal basis is wide and flat. In the middle hat convex and tubercular external surface. The of it, there is a narrow elliptical stem facet. The ful­ height of the centrodorsal is smaller than that of the cral ridge is sometimes hardly visibly serrated. The radial ring. The external centrodorsal surface has the marginal ridge, in most cases, bears big irregular same sculpture as the radiais. A slit-like depression is teeth. The columnals are low, parallel, rarely cruciate under each radial, on the radial-centrodorsal suture. or oblique. The external columnal surface is smooth or tubercular.

Fig. 8 — Calyxes, centrodorsals and brachials of Loriolicrinus (x 3). A - holotype of L. insuetus (calyx from side, after De Loriol 1891, pi. 29, fig. 23). В - L. asper, CK-82-5 (centrodorsal from above and below). С - L. asper, CK-82-10 (primaxillary from above, side and below). D - L. asper, holotype CK-82-1 (calyx from side). E - L. asper, CK-82-3 (young calyx from side). F - L. asper, CK-82-4 (centrodorsal from below). G - L. asper, CK-82-12 (tertibrachial from above). Coupes dorsales, centrodorsales et brachiales de Loriolicrinus. A - holotype de L. insuetus (coupe dorsale vue de côté, d’après De Loriol). B - L. asper, CK-82-5 (centrodorsale vue en dessus et en dessous). С - L. asper, CK-82-10 (primaxillaire vue en dessus, de côté et en dessous). D - L. asper, holotype CK-82-1 (coupe dorsale vue de côté). E - L. asper, CK-82-3 (coupe dorsale jeune vue de côté). F - L. asper, CK-82-4 (centrodorsale vue en dessous). G - L. asper, CK-82-12 (tertibrachiale vue en dessus). — 643 —

Dimensions : Loriolicrinus laevis nov. sp. Ht = 4.5 - 9.5 mm ; Dt = 5.9 - 15.7 mm ; De = 3.7 fig. 9 A-D ; pl. 1, fig. 6 - 10.2 mm ; A f = 41 - 42° ; Hr : Ht = 0.32 - 0.38 ; Bf : Lf = 0.48 - 0.66. D e r iv a t io n o m i n is : The name of the species comes from « laevis » Comparison : (smooth in Latin). L. asper differs from other species of the genus by very large convex radiais and by its coarse pustulate H o l o t y p e : sculpture. CK-80-1 (fig. 9 A) ; Leningrad Mining Institute.

Materials : T y p e l o c a l it y : 3 calixes, 5 centrodorsals, 1 radial ring, 12 bra­ chials, 95 columnals (see table 1). Rodnoie, Chornaya Basin, Crimea.

Distribution : T y p e l e v e l : Berriasian ; Crimea. Upper Berriasian.

Fig. 9 — Calyxes, centrodorsals and brachials of Loriolicrinus (x 3). A - L. laevis, holotype CK-80-1 (calyx from side and below). В - L. laevis, CK-18-1 (centrodorsal from above). С - L. laevis, CK-90-5 (young calyx from side). D - L. laevis, CK-90-6 (secondaxillary from below, side and above). E - L. perforatus, holotype CK-95-1 (calyx from side).

Coupes dorsales, centrodorsales et brachiales de Loriolicrinus. A - L. laevis, holotype CK-80-1 (coupe dorsale vue de côté et en dessous). B - L. laevis, CK-18-1 (centrodorsale vue en dessus). С - L. laevis, CK-90-5 (coupe dorsale jeune vue de côté). D - L. laevis, CK-90-6 (secundaxillaire vue en dessous, de côté et en dessus). E - L. perforatus, holotype CK-95-1 (coupe dorsale vue de côté). — 644 —

D e s c r i p t i o n : H o l o t y p e : The calyx is low, cylindrical or conical, smooth with CK-95-1 (fig. 9 E) ; Leningrad Mining Institute. rounded outline. The basais are not visible on the calyx surface. The external radial surface is low and T y p e l o c a l it y : convex. The radial facets are wide. The calyx cavity is narrow and shallow. The radial and interradial fur­ Karatlykh Mountain, Belbek Valley, Crimea. rows are well pronounced. The basais are visible on the bottom of the calyx cavity. The primaxillary is not T y p e l e v e l : high with the convex and smooth external surface. Upper Berriasian. The centrodorsal is low. Under each radial, on the radial-centrodorsal suture, there is one small, hardly D e s c r i p t i o n : visible fossa. The centrodorsal basis is broad-elliptical and concave. The stem facet is narrow and elliptical. The calyx is low, cylindrical or conical, smooth and Often there is the intermediate ridge besides the ful­ rounded. The basais are visible on the calyx surface. cral and marginal ones. The columnals are parallel or The external radial surface is high and convex. The rarely cruciate, and not high. radial facets are wide. The calyx cavity is narrow and shallow. The basais are not visible on the bottom of the calyx cavity. The centrodorsal is not high. One D im e n s i o n s : large fossa, turning downwards, is situated under Ht = 6.0 - 7.7 mm ; Dt = 10.0 - 11.7 mm ; Dc = each radial on the radial-centrodorsal suture. The 6.6 - 7.0 mm ; Af = 36° ; Hr : Ht = 0.25 - 0.32 ; Bf : centrodorsal basis is wide, round and flat. The stem Lf = 0.60 - 0.65. facet is narrow-elliptical.

D i m e n s i o n s : C o m p a r i s o n : ht = 6.2 - 9.2 mm ; Dt = 10.0 - 12.2 mm ; Dc = L. laevis differs from L. asper by its smooth calyx 7.0 - 8.0 mm ; Af = 36° ; Hr : Ht = 0.40 . Bf : Lf = surface ; from L. perforatus by the absence of the big 0.53. fossae under the radiais and by the absence of the basais on the external calyx surface ; from L. insuetus C o m p a r a i s o n : (with which the species under discussion has the grea­ test likeness) it differs by the radiais of small relative L. perforatus differs from other species of the genus height and by the concave centrodorsal basis. Loriolicrinus having a big fossa under each radial. This species has the great likeness with Solanocrinites jaegeri Q e n s t e d t (see above) as for the shape of the M a t e r i a l s : fossae on the calyx surface. But L. perforatus differs 9 calyxes, 3 centrodorsals, 2 brachials, 11 columnals from it by a smaller height of the centrodorsal and by (see table 1). relatively higher radiais.

M a t e r i a l s : D istribution : 3 calyxes (see table 1). Berriasian ; Crimea.

D istribution : Berriasian ; Crimea. Loriolicrinus perforatus nov. sp. fig. 9 E Genus Conoideocrinus nov. gen. Derivatio nominis : Type species : The name of the species comes from « perforatus » (holey, perforated in Latin). Conoideocrinus conoideus nov. gen., nov. sp. — 645 —

D ia g n o s is : T y p e l o c a l it y : The radiais have high external surface. The large Kuibyshevo, Belbek Valley, Crimea. basais are visible on the calyx surface. The cirrus soc­ kets and fossae are absent. The centrodorsal is small T y p e l e v e l : and conical. The stem facet is small, rounded and flat. Upper Berriasian.

D e s c r i p t i o n : Co m p a r is o n : The calyx is high-conical, smooth and rounded. The Conoideocrinus differs from other genera of the external radial surface is very high. The sutures bet­ family having high-conical calyx with large basais and ween the radiais, basais and the centrodorsal are with no trace of the cirrus socket or fossa. hardly visible. Radial facets are narrow. The calyx One type species. cavity is broad and deep. Radial and interradial fur­ Lower Cretaceous (Berriasian) ; Crimea. rows are weak. The height of the centrodorsal is smal­ ler than that of the radial ring. The centrodorsal basis is rounded and flat with a pronounced fulcral ridge.

Conoideocrinus conoideus nov. sp. D im e n s i o n s : fig. 10 A Ht = 7.9 mm ; Dt = 11.0 mm ; Dc = 6.2 mm ; Af = 26° ; Hr : Ht = 0.51 - 0.72 ; Bf : Lf = 0.33 - 0.38.

D e r iv a t io n o m i n i s : M a t e r i a l : The name of the species comes from « conoideus » (cone-shaped in Latin). One calyx (see table 1).

H o l o t y p e : D istribution : CK-97-1 (fig. 10 A) ; Leningrad Mining Institute. Berriasian ; Crimea.

Fig. 10 — Calyxes and brachials of Conoideocrinus and Umbocrinus (x 3). A - C. conoideus, holotype CK-97-1 (calyx from side). В - U. umbonatus, CK-89-4 (small calyx from side). С - U. umbonatus, CK-83-2 (calyx from side). D - U. umbonatus, CK-83-6 (odd secondbrachial from above - on the left, and from below - on the right). Е - U. umbonatus, CK-83-7 (even secondbrachial from above).

Coupes dorsales et brachiales de Conoideocrinus et Umbocrinus. A - C. conoideus, holotype CK-97-1 (coupe dorsale vue de côté). B - U. umbonatus, CK-89-4 (petite coupe dorsale vue de côté). С - U. umbonatus, CK-83-2 (coupe dorsale vue de côté). D - U. umbonatus, CK-83-6 (secundibrachiale impaire vue en dessus - à gauche, vue en dessous - à droite). E - U. umbonatus, CK-83-7 (secundibrachiale paire vue en dessus). — 646 —

Genus Umbocrinus nov. gen. The radial facets are wide. The calyx cavity is broad and shallow. The radial and interradial furrows are T y p e s p e c ie s : pronounced. The basais are visible or invisible on the bottom of the calyx cavity. The primaxillary is flatte­ Umbocrinus umbonatus nov. gen., nov. sp. ned with strongly convex, smooth or granulated exter­ nal surface. The height of the centrodorsal roughly D ia g n o s i s : equals that of the radial ring. One to three deep fossae are placed under each radial, on the radial = centro­ The radiais have low external surface. The basais dorsal suture. The centrodorsal basis is wide, rounded are visible on the calyx surface. A row of fossae is pla­ and concave. The stem facet is narrow-elliptical. The ced on the suture between the radiais and the centro­ columnals are low, parallel, rarely cruciate or obli­ dorsal. The centrodorsal is small and convex. The que. Their external surface is smooth or granulated. stem facet is rounded and concave. The fulcral ridge is serrated. Intermediate ridges are often developed on the columnal articula. C o m p a r i s o n :

Umbocrinus differs from Solonaerium and Thiollie­ D i m e n s i o n s : ricrinus lacking cirrus sockets ; from Burdigalocrinus Ht = 4.5 - 10.2 mm ; Dt = 7.0 - 21.0 mm ; Dc = by lower radiais and by the fossae being placed only 5.0 - 10.0 mm ; A f = 45 - 57° ; Hr : H t = 0.09 on the radial-centrodorsal suture ; from the similar -0,25 ; Bf : Lf = 0.48 - 0.64. genus Heberticrinus by the fossae on the calyx and the basais on the external calyx surface. M a t e r i a l s : One type species. Lower Cretaceous (Berriasian) ; Crimea. 32 calyxes, 21 centrodorsals, 8 brachials, 52 colum­ nals (see table 1).

D istribution : Umbocrinus umbonatus nov. sp. Berriasian ; Crimea. fig. 10 B-E ; pl. 1, fig. 7-11

D e r iv a t io n o m i n is : Genus Heberticrinus nov. gen. Th'ê name of the species comes from « umbonatus »

(convex in Latin). T y p e s p e c ie s :

Eugeniacrinus heberti D e L o r io l in P i c t e t , 1868. H o l o t y p e :

CK-89-1 (pl. 1, fig. 7) ; Leningrad Mining Institute. D i a g n o s i s : The radiais are with or without low external sur­ T y p e l o c a l it y : face. The basais are invisible on the calyx surface. Kuibyshevo, Belbek Valley, Crimea. Rarely they look like tiny triangular plates. The cen­ trodorsal is large, convex with no cirrus socket or fossa. The stem facet is concave, flat or rarely convex. T y p e l e v e l : Upper Berriasian. C o m p a r i s o n : Heberticrinus differs from all genera of the family D e s c r i p t i o n : lacking cirrus sockets or fossae on the centrodorsal. It The calyx is low, smooth and rounded. The basais differs from Conoideocrinus, which has no socket or are visible on the calyx surface but sometimes not in fossa, by the large massive centrodorsal, by the all interradia. The external radial surface is low, con­ absence of basais on the external calyx surface and by vex, smooth or rarely covered with small granules. the low radiais. — 647 —

В

Fig. 11 — Calyxes of Heberticrinus (x 3). A - holotype of H. algarbiensis (calyx from side and below, after De Loriol 1888, pi. 22, fig. 10). В - holotype of H. crassus (calyx from side and below, after Remes 1905, pi. 7, fig. 2). С - lectotype of H. remesi (calyx from side and below, after Remes 1905, pi. 7, fig. 6). D - H. remesi, CK-56-1 (calyx from side). E - H. aff. heberti, CK-96-2 (calyx from below). F - H. aff. heberti, CK-96-1 (calyx from side).

Coupes dorsales de Heberticrinus. A - holotype de H. algarbiensis (coupe dorsale vue de côté et en dessous, d’après De Loriol). B - holotype de H. crassus (coupe dorsale vue de côté et en dessous, d’après Remes). С - lectotype de H. remesi (coupe dorsale vue de côté et en dessous, d’après Remes). D - H. remesi, CK-56-1 (coupe dorsale vue de côté). E - H. aff. heberti, CK-96-2 (coupe dorsale vue en dessous). F - H. aff. heberti, CK-96-1 (coupe dorsale vue de côté). — 648 —

Fig. 12 — Calyxes and brachials of Heberticrinus heberti (x 3). A - CK-51-1 (calyx from above, from side and below). В - CK-51-2 (calyx from side and below). С - CK-51-5 (calyx from above and side). D - CK-51-7 (calyx from side and below). E - CK-51-10 (small calyx from side). F - CK-51-9 (small calyx from side). G - CK-69-1 (calyx from side). H - CK-51-97 (primaxillary from above and below). I - CK-30-1 (radial ring with basais from below). К - CK-51-70 (radial ring with basais from below). L - CK-51-133 (right even secondbrachial from above on the left, and from below on the right). M - CK-51-167 (secondaxillary from above). N - CK-51-206 (left odd secondbrachial from above on the left, and from below on the right). — 649 —

Five species : H. algarbiensis (D e Lo r io l ), H. crassus 1888 — Thiolliericrinus heberti De Lo r io l : De (Gisl é n ), H. heberti (D e Lo r io l in P ic t e t ), H. Loriol, p. 111. remesi (G is l é n ), H. irregularis nov. sp. One calyx found in the Hauterivian deposits of the settlement 1889 — Thiolliericrinus heberti De l o r io l : De Gunib in Daghestan (Caucasus) belongs to the genus Loriol, p. 545 (ex parte), pi. 228, fig. 1-4, 6- Heberticrinus. However, a poor preservation does not 13, pi. 229, fig. 1 (non pi. 228, fig. 5 = permit us to a specific determination. Thiolliericrinidae indet.). Upper Jurassic (Tithonian) - Lower Cretaceous 1897 — Thiolliericrinus heberti D e L o r i o l : (Hauterivian) ; Portugal, France, Czechoslovakia, Roman, p. 88, 331, 334. Rumania, Crimea, Caucasus. 1897 — Thiolliericrinus flexuosus Go l d f u s s sp. : Roman, p. 88, 331, 334. Heberticrinus algarbiensis (De Lo r io l , 1888, p. Ill, pi. 22, fig. 10, 11 sub Thiolliericrinus {Thiollieri­ 1905 — Thiolliericrinus heberti De Lo r io l ; Remes, crinus algarbiensis De Loriol 1889, p. 560 ; Burdiga- p.60 (ex parte), pi. 7, fig. 3 (non fig. 2 = locrinus algarbiensis : Gislén 1924, p. 189 ; Rasmus­ Heberticrinus crassus). sen 1961, p. 213, pi. 35, fig. 11, 12). Hauterivian ; 1912 — Thiolliericrinus heberti De l o r io l : Kirk, p. Alfandega (Algarve), Portugal (fig. 11 A). 75, pi. 7, fig. 3 (fig. ex De Loriol 1889, pi. Heberticrinus crassus (G i s l é n , 1924, p. 189 sub 228, fig. 2). emes Burdigalocrinus) (= Thiolliericrinus heberti R >, 1913 — Thiolliericrinus heberti (D e L o r io l ) : Jou- 1905, p.60 ex parte, pi. 7, fig. 2, non fig. 3 = Heberti­ kowsky & Favre, p. 386, pi. 15, fig. 1-6. crinus heberti. Tithonian . Stramberk, Czechoslova­ kia. 1924 — Burdigalocrinus heberti (De Lo r io l ) : Gis­ lén, p. 189.

1961 — Burdigalocrinus heberti (De Lo r io l ) : Ras­ mussen, p. 214. Heberticrinus heberti (D e Lo r io l in P i c t e t , 1868) 1978c— Thiolliericrinus heberti De Lo r io l : Ras­ figs. 11, E,F? ; 12 A-N ; pl. 1, fig . 12-16 mussen, p. 880, fig. 588-2e,f (fig. ex De L o rio l, 1889, pi. 228, fig. 7a et 2a).

1868 — Eugeniacrinus heberti : D e Lo r io l (in P ic ­ t e t ), p. 281, pi. 42, fig. 7,8. Le c t o t y p e : 1875 — Eugeniacrinus heberti (De Lo r io l ) : Pillet & F.J. Pictet (1868, pi. 42, fig. 8) ; Museum in Cham- Fromentel, p. 98, pi. 10, fig. 31-35. béry (Savoie, France). 1875 — Apiocrinus flexuosus (G o l d f u s s ) : Pillet &

Fromentel, p. 98, pi. 10, fig. 45-50. T y p e l o c a l it y : 1883 — Eugeniacrinus heberti D e L o r io l : D e Lémenc (Savoie), France. L o rio l, p. 158.

Coupes dorsales et brachiales de Heberticrinus heberti. A - CK-51-1 (coupe dorsale vue en dessus, de côté et en dessous). В - CK-51-2 (coupe dorsale vue de côté et en dessous). С - CK-51-5 (coupe dorsale vue en dessus et de côté). D - CK-51-7 (coupe dorsale vue de côté et en dessous). E - CK-51-10 (petite coupe dorsale vue de côté). F - CK-51-9 (petite coupe dorsale vue de côté). G - CK-69-1 (coupe dorsale vue de côté). H - CK-51-97 (primaxillaire vue en dessus et en dessous). I - CK-30-1 (cercle des radiales avec les basales, vue en dessous). К - CK-51-70 (cercle des radiales avec les basales, vue en dessous). L - CK-51-133 (secundibrachiale droite paire vue en dessus à gauche, vue en dessous à droite). M - CK-51-167 (secundaxillaire vue en dessus). N - CK-51-206 (secundibrachiale droite impaire vue en dessus à gauche, vue en dessous à droite). — 650 —

T y p e l e v e l : D istribution : Upper Tithonian. Tithonian and Berriasian ; France, Czechoslovakia, Rumania, Crimea. D ia g n o s is : The radial ring is narrowed upwards. Free radial surface is absent (i.e. radial facets are near the centro­ dorsal). The basais are not visible on the calyx sur­ face. The centrodorsal is low and wide. Its diameter is Heberticrinus remesi ( G i s l é n , 1924) considerably greater than that of the radial ring. The fig. 11 C,D centrodorsal basis is wide, rounded and concave. The stem facet is narrow-elliptical. 1905 — Thiolliericrinus flexuosus É tallon (Gold- f u s s ) : Remes, p. 61, pi. 7, fig. 4-6. D im e n s io n s : 1924 — Burdigalocrinus remesi : Gislén, p. 190. Ht = 3.0 - 9.6 mm ; Dt = 4.4 - 19.6 mm ; Dc = 3.5 - 15.0 mm ; A f = 47 - 52° ; Hr : H t = 0.00 ; Bf : Lf = 0.50- 0.63. L e c t o t y p e : Remes (1905, pi. 7, fig. 6) ; Charles University, Pra­ Re m a r k : gue (teste Zitt, in letteris 1.4.86). When De Loriol (in Pictet 1868, p. 281) was descri­ bing Eugeniacrinus heberti he indicated its occurrence T y p e l o c a l it y : in the « Brèche de Lémenc ». But he postulated the v Sequanian age of the species in his following publica­ Stramberk, Czechoslovakia. tions. However, this is inaccurate. Remains of the species under discussion were found in South-Eastern T y p e l e v e l : France (including Lémenc) together with the spines of Cidaris glandifera, the shells of the brachiopodPy- Upper Tithonian. gope janitor, the tests of the rudistid Heterodiceras luci and with the remains of the ammonite Paraula- D i a g n o s i s : cosphinctes transitorius. The cited faunistic assem­ blage demonstrates obviously the Late Tithonian age The radial ring is narrowed upwards. Free radial of H. heberti. This species was found in the same age surface is absent (i.e. radial facets are near the centro­ deposits in Czechoslovakia (RemeS 1905, p. 60) and dorsal). The basais are invisible on the calyx surface. Rumania (Patrulius 1956b, p. 198). But in the Crimea The centrodorsal is high, convex and narrowed down­ it is found in the reef massives directly above the beds wards. The centrodorsal basis is completely occupied with Dalmasiceras dalmasi (i.e. in the Upper Berria­ by a narrow-elliptical stem facet. sian). Note shall be taken that, according to Yanin’s opinion (in his letter of 1.3.86), the species Heterodi­ D i m e n s i o n s : ceras luci is characteristic for the Berriasian. One can assume therefore that a part of the French localities, H t - 9.3 - 12.9 mm ; Dt = 13.0 - 16.0 mm ; Dc = where remains of this rudistid and H. heberti were 9.0 - 10.4 mm ; A f = 33 - 46° ; Hr : Ht = 0.00 ; Bf : found, is Berriasian. Lf = 0.50 - 0.60. On some calyxes of H. heberti and H. irregularis affected by parasites, one can see the large holes (fig. M a t e r i a l s : 12 A, 13 A ; cf. Müller 1977, fig. 7). 4 calyxes, 9 columnals (see table 1).

M a t e r ia l s : D istribution : 87 calyxes, 32 centrodorsals, 2 radial rings, 258 bra­ chials, 184 columnals (see table 1). Tithonian, Czechoslovakia ; Valanginian, Crimea. — 651 —

Heberticrinus irregularis nov. sp. surface. The centrodorsal surface is smooth or very finely granulated. Some weak fossae are seen someti­ fig. 13 A-E mes on the radial-centrodorsal suture. The centrodor­ sal basis in projected as an acute ridge and completely De r iv a t io n o m i n i s : occupied by an elliptical stem facet being orientated The name of the species comes from « irregularis » obliquely to the calyx axis. In some cases, the basis is (irregular in Latin). wide, flat and rounded.

H o l o t y p e : D i m e n s i o n s : CK-88-1 (fig. 13 C) ; Leningrad Mining Institute. H t = 5.2 - 12.0 mm ; Dt = 6.9 - 18.5 mm ; Dc = 5.0 - 12.0 mm ; Af = 37 - 50° ; Hr : Ht = 0.06 -0.23 ; Bf : Lf = 0.36 - 0.50. Ty p e l o c a l it y :

Kuibyshevo, Belbek Valley, Crimea. C o m p a r i s o n : H. irregularis differs from other species of the Ty pe l e v e l : genus having an irregular centrodorsal and oblique Upper Berriasian. centrodorsal basis.

De s c r ip t io n : M a t e r i a l s : The calyx is high and rounded or rounded- 17 calyxes, 18 centrodorsals, 2 brachials, 28 colum- pentangular. The radial ring is high, cylindrical or nals (see table 1). narrowed downwards. Free radial surface is low or relatively high. Radial facets are narrow. The basais D istribution : are visible on the external calyx surface. The centro­ dorsal is high and convex with uneven external Berriasian ; Crimea.

В

Fig. 13 — Calyxes, controdorsal and columnal of Heberticrinus irregularis (x 3). A - CK-88-3 (calyx from side and below). В - CK-88-2 (calyx from side). С - holotype CK-88-1 (calyx from side). D - CK-88-22 (columnal and its articular face). E - CK-88-8 (centrodorsal from above). Coupes dorsales, centrodorsale et columnale de Heberticrinus irregularis. A - CK-88-3 (coupe dorsale vue de côté et en dessous). B - CK-88-2 (coupe dorsale vue de côté). С - holotype CK-88-1 (coupe dorsale vue de côté). D - CK-88-22 (columnale et sa facette articulaire). E - CK-88-8 (centrodorsale vue en dessus). — 652 —

5 — RECONSTRUCTION OF THE SKELETON

Ro o t f o r m a t io n s raised socket. Dactylate corallite served, in most cases, a substratum for the attachment crusts. One can mark two peculiarities of the attachment In some localities (see table 1), together with calyxes formations. First, a small diameter of the stem facets. and columnals, root formations of Thiolliericrinidae It is considerably smaller than the average diameter of were found (Ad - see fig. 2). One cannot determine columnals found in the same localities. This fact can the taxonomic position of these roots. It can only be mean that the Thiolliericrinidae stem is narrowed assume that they belong to Heberticrinus heberti or downwards from the calyx. Second circumstance : Umbocrinus umbonatus (the most common species). almost all stem facets on attachment formations have All attachment outgrowths are individual, i.e. only evident traces of the selective dissolution. The liga­ one articular facet for a stem is present on each. These ment fossae on both sides of the fulcral ridge are con­ formations are the monocrystalline calcitic crust siderably deepened. This dissolution is explained by (without root-like projections) bearing a nearly not the fact that the root was left on its place long after isolated stem socket (fig. 14 A,B). Sometimes, the the death of the animal, being exposed to the corro­ stem facet is raised as a distinct eminence (fig. 14 sion by sea water. C,D). Such eminences are formed, very likely, by an Described root formations are analogous to those accretion of the lower columnal with the attachment noted for other Thiolliericrinidae species (Étallon disc. It is characteristic that a rounded swelling, 1859, p. 445, 447 ; Patrulius 1956a, p. 193 ; Hess resembling a parasite settling, is situated near each 1975, p. 69, pi. 25, fig. 1 ; Rasmussen 1978c, p. 879).

В

Fig. 14 — Root formations of Thiolliericrinidae indet. (x 2). A - CK-100-3 : attachment to a coral limestone ; stem sockets is low. В - CK-100-4 : attachment to a coral ; stem socket is low. С - CK-100-2 : attachment to a shell fragment ; stem socket is high. D - CK-100-1 : attachment to a coral limestone ; stem socket is high.

Mode de fixation de quelques Thiolliericrinidae indét. A - CK-100-3 : fixation sur un calcaire corallien. В - CK-100-4 : fixation sur un polypier. С - CK-100-2 : fixation sur un fragment de coquille. D - CK-100-1 : fixation sur un calcaire corallien. St e m distal or proximal position of the columnal in a stem is probably the matter. Unfortunately, the lack of Étallon (1859, p. 445, 447) noted that the Thiollieri­ finds of stem fragments (i.e. some articulated colum­ crinus stem had consisted only of three or four colum­ nals) does not allow one to make the nature of a stem nals. Joukovsky & Favre (1913, p. 386) found 31 variation clear. calyxes and only 43 columnals of Heberticrinus heberti in one of the local exposures. Similar correla­ If the cirriless calyxes of Thiolliericrinidae are well tions are marked also in our materials. So, in the loca­ preserved they have also well preserved stem facet. lity « Belbek » (see table 1), there were found 129 However, available specimens of Thiolliericrinus calyxes and 638 columnals, belonging to several spe­ (with poverful cirri) have obviously abraded and des- cies. A comparison of the figures shows that the tructed centrodorsal basis (fig. 5 A, 6A,B) even when Thiolliericrinidae stem included roughly five colum­ the preservation as a whole is good. This circumstance nals. It coincides with Étallon’s data. can mean that the cirriless forms were attached to the Among Heberticrinus heberti columnals, cruciate stem (and, accordingly, to the substratum) during ones are the bulk, oblique ones are rare and parallel their whole life. On the contrary, the individuals ones are unique. The columnals of this species are low having cirri were detached from their stem and led a and disk-shaped. At the same time, the columnals free life though preserving a biletarel stem facet on with parallel ridges on both articula are prevalent in the centrodorsal (Kirk 1912, p. 75). The poor preser­ Burdigalocrinus maximus. The columnals of this spe­ vation of the centrodorsal basis in these forms is cies are, as a rule, relatively high. These circumstances explained, possibly, by their activity. The separation allow us to suppose that the short stem of some dis- of the stem during the life of an animal is confirmed coidal columnals was peculiar to the species with wide by the absence (among over 500 columnals) of a single flattened calyxes. But a relatively long stem, consis­ columnal belonging to Thiolliericrinus belbekensis, T. ting of high columnals, was typical for species with a elongatus or T. torosus. high conical calyx. The order of columnal combination in the stem is A r m s . unknown. However, one can assume, after the ana­ Thiolliericrinidae brachials are known till now only logy to Bourgueticrinida, that the columnals were joi­ after Rem e!s (1905, pi. 7, fig. 7,8). In our collection ned so that the fulcral ridge had an alternating « swin­ there are 580 brachials plates of several species of the ging » position. I doubt whether one can speak, in the family (see table 1). present case, of « spiral turning » as supposed by A rendt (1974, p. 75). In the species Loriolicrinus Only muscular articula are developed on all the bra­ asper, cruciate and parallel columnals are found in chials without exception, including the lower face of approximately equal numbers. One can think that axillaries. This leads to the conclusion that there were they were joined in a stem alternately. no immovable articulations in the arms of Thiollieri­ A powerful fulcral ridge risen above the articulum crinidae. plane and deep ligament fossae on both sides of them The axillaries of the first, second and third orders testify to the considerable flexibility of the stem arti­ can be determined with certain among available axil- culations in Thiolliericrinidae (Gislén 1924, p. 206). It lar plates. Consequently, arm branching was not less results in flexibility, as a rule, without any marked than threefold. increase of the separate columnal height as in Bour­ gueticrinida with an analogous articulum construc­ Not a plate has been found which should be placed tion. The high stem flexibility with the relatively low between the primaxillary and the radial. So, one can (strong) columnals agrees with the fact that Thiollieri­ conclude that Thiolliericrinidae have only one primi- crinidae inhabited very mobil waters on the coral reef. brachial (axillary) with muscular articulation directly joined to the radial. The external sculpture of columnals corresponds that of the centrodorsal, in most cases. The columnals The primaxillaries (IAx - see figs. 2 and 15) are the of Loriolicrinus asper are often knobby (pl. 1, fig. 5) largest plates in the crown, being greater than the and those of Umbocrinus umbonatus are covered radiais. The form of promaxillaries is various in diffe­ sometimes with small granules. At the same time, rent genera. The calyxes with high free radial surface smooth columnals are often found, belonging evi­ and with a small angle of the inclination of the radial dently to the ornamented calyxes. In this case, the facets have high axillaries with vertical external — 654 —

surface (Loriolicrinus, Burdigalocrinus). The calyxes SI Sr with low radiais whose facets are steeply inclined out­ side have wide and flattened primaxillaries hung over the centrodrosal and absolutely covering radial plates (Umbocrinus, Heberticrinus). Two upper primaxil- lary facets are of an identical size and inclined outside nearly parallel to the facets of the subjacent radiais. The lateral faces of all available axillar plates are flat­ tened. Consequently, thesé plates formed a closed llAx ring. The sculpture of the external surface of a pri- maxillary corresponds to that of the calyx. The pinnu- lar sockets are absent on primaxillaries. The brachials of most Articulata crinoids bear one IIBr3 pinnular socket placed on the upper inner margin of these plates. The fulcral ridges are not in parallels on lower and upper brachial faces. But these ridges can have only two positions swinging from one articulum to another. The available secondbrachials were divi­ l!Br2 ded into right and left, even and odd, accordingly (fig. 15). If one looks at the primaxillary from its external surface, the left odd (first) secondbrachial will be on its upper face to the left and right odd (first) secondbrachial to the right. The odd secondbrachials are cuneate with a thin margin at the inner side of a IIBM radius and with a thick one at the outer side where the pinnular socket is placed. Their external surface is very convex. The thick lateral side of the secondbra­ chial bears a wide flat face, i.e. these plates were close to the corresponding brachials of the adjacent radius. The even brachials are flattened and somewhat higher at the inner margin (where the pinnular socket is pla­ ced). They have convex external surface and flat late­ ral sides joined to corresponding secondbrachials of the adjacent radii. The number of secondbrachials in the arm branch was apparently odd and hardly more than seven or nine (according to the relatively great number of the available secondaxillaries). The exter­ nal secondbrachial surface has the sculpture analo­ gous to that of the calyx but feebly expressed. The secondaxillaries (IIAx - see figs. 2 and 15), according to their construction, are analogous to the

Fig. 15 — A scheme of the construction of Thiolliericrinidae bra­ Reconstitution de l’organisation d’un tronc brachial de Thiollieri­ chials (view from above, x 2). IAx - ring of five pri­ crinidae (vue en dessus, x 2). IAx - cercle des cinq primaxillaires, maxillaries, IIBrl - first secondbrachial, IIBr2 - second IIBrl - première secundibrachiale, IIBr2 - deuxième secundibrachiale, secondbrachial, IIBr3 - third secondbrachial, IIAx IIBr 3 - troisième secundibrachiale, IIAx - secundaxillaire, IIIBrl - -secondaxillar, IIIBrl - first tertibrachial, IIIBr2 -second première tertibrachiale, IIIBr2 - deuxième tertibrachiale, SI -série tertibrachial, SI - left arm series, Sr - right arm series. gauche d’un bras, Sr - série droite d’un bras. Le nombre des secundi- The number of secondbrachials is shown conditionally. brachiales reste hypothétique et n’est donné qu’à titre indicatif. — 655 —

Fig. 16 — Reconstruction of the skeleton of some Thiolliericrinidae (x 1) : Thiolliericrinus (below from the left ; a part of the arms is remo­ ved), Heberticrinus (in the centre) and Burdigalocrinus (from the right ; traces of parasites settlings are shown on the central radius). Reconstitution du squelette de quelques Thiolliericrinidae (x 1) : Thiolliericrinus (en bas à gauche ; une partie des bras est enlevée), Heberticrinus (au centre) et Burdigalocrinus (à droite ; les traces de l’action de parasites sont montrées sur le rayon central). primaxillaries but are asymmetrical. Their inner The crown of Thiolliericrinidae was uniform on the upper facet is considerably greater than the outer one. whole, differing only in details in different species The construction of tertibrachials is like that of the and even genera. It formed, in the closed position, a secondbrachials. Only the wedge-shaped form of continuous tube at least up to the level of the tertiaxil­ separate plates is more pronounced. Also the odd bra­ laries (fig. 16). By that, relatively very small pinnules chials have their external surface like an obtuse hori­ were enclosed into the crown and not visible from the zontal ridge. One failed to determine the number of outside. One can say that the crown skeleton was a brachials in the third arm series. The tertiaxillaries are test being made of closely adjusted and firmly united symmetrical having flattened lateral faces as before. brachial plates. The Thiolliericrinidae crown resem­ The axillaries of the second and third orders have no bled, in this respect, the crown of Triassic Encrinus pinnules. The tertibrachials have smooth external sur­ liliiformis L a m a r c k but was considerably greater. face without any dependence on the calyx ornamenta­ tion.

6 — ONTOGENETIC VARIATIONS OF THE CALYXES

According to available materials, the calyx onto­ geny could be reconstructed for three Thiolliericrini­ dae species : Loriolicrinus asper, Umbocrinus umbo- natus and Heberticrinus heberti. One fact attracts our attention : in all three forms, in spite of their belon­ ging to different genera, there are almost the same calyxes in young specimens, namely : high-conical or cylindrical with high free radial surface. However, subsequently the development of the calyxes procee­ ded by different ways in each of the three species. The radiais grew quicker than the centrodorsal in L. asper (fig. 17A). Therefore the centrodorsal of this species remained low and its upper surface is convex. The young forms of U. umbonatus (fig. 17B) have strongly pronounced though low basais and fully developed fossae under radiais. Then the centrodorsal and the radiais grew at an about equal speed. There­ fore the upper centrodorsal surface is flat. Simulta­ neously with the preserving of the basais on the exter­ nal calyx surface, the relative height of the free radial surface considerably decreased. The sutural pores did not disappear but even increased in number and size, so that they changed into irregular openings

Fig. 17 — Schematic sections of some Thiolliericrinidae calyxes showing the stages of the ontogenetic development (R - radial, В - basal, Cd, centrodorsal) : A - Loriolicrinus, В - Umbocrinus, С - Heberticrinus. Sections schématiques des coupes dorsales de quelques cd c Thiolliericrinidae, montrant les stages du développe­ ment ontogénétique. — 657 — occupying nearly the whole centrodorsal surface in the young specimens, disappeared in the adult ones. the oldest specimens. The stem facet, being flat in the The height of the radiais decreased. The diameter of young specimens, turned into concave in the adult the centrodorsal was roughly equal to that of the ones. radial ring in the young specimens. However, the cen­ The centrodorsal grew quicker than the radiais in trodorsal was considerably wider than the radiais in H. heberti (fig. 17C). Therefore, the upper centrodor­ the adult specimens. The stem facet was almost sal surface is very concave. The basais, being visible in always flat.

7 — ORIGIN OF THE THIOLLIERICRINIDAE

Many thiolliericrinids have the quite developed cirri and cirriless forms with quite large infrabasals, and with the mobil bifascial articulation of cirrals on their that they had many features common with encrinids. centrodorsals. Only Comatulida and Pentacrinida * Then, the development of pentacrinids followed a may have such cirrus construction among post- path of perfection of the cirral organs (with a moun­ paleozoic crinoids. So specialized cirral organs could ting of the five-rayed nodal symmetry and an appea­ not originated twice in the phylogeny of crinoids. rance of the cryptosymplectial sutures : Hagdorn Therefore, the origin of Thiolliericrinidae must be 1983, p. 357 ff.), transmutation of the calyx from consider in the close connection with the origin of dicyclic to cryptodicyclic and disappearance of the comatulids and pentacrinids in one phylogenetic morphological isolation of the calyx from the arms. sequence with these groups. Cited data show that origin and development of the Three hypothesis are possible in the comprehension pentacrinids followed without any connection with of the development of three given groups, namely : the considerably more late Thiolliericrinidae. Conse­ (1) stalked pentacrinids, (2) stalked thiolliericrinids or quently, only one of above mentioned hypothesis (3) stemless comatulids are initial. The embryological remains acceptable : the pentacrinids are the ancestor investigations showed that young stages of comatulids of the cirriferous Articulata. were stalked (Thomson 1865 ; A.H. Clark 1921). The By subsequent consideration of this hypothesis, two fine and peculiarly constructed perforation on the versions are possible : (1) Pentacrinida —» Thiollieri­ lower centrodorsal face of some (relatively primitive crinidae —> Comatulida (De Loriol 1880, p. 11 ; Kirk ?) comatulids (A.M. Clark 1973, fig. 1—3) are also 1912, p. 75 ; Jaekel 1918, p. 71 and others) or (2) Pen­ the illustration of the stalkness of comatulid ances­ tacrinida —> Comatulida —> Thiolliericrinidae (Ras­ tors. Available facts testify to the origin of Comatu­ mussen 1978b, p. 314 ; 1978c, p. 867, 879). lida from stalked forms and to the unacceptability of third (see above) hypothesis respectively. One can make out two stages in the skeleton deve­ lopment of the recent comatulids. Deltoid plates, Earliest pentacrinids known beginning with early large basais, minute infrabasals and columnals are Triassic (Holocrinidae) had a number of specific fea­ laid at early cystidean stage. Absence of cirri and the tures. One should note the following primitive signs : construction of the calyx basis permit to see, in this the distal attachement disc (replaced later into unatta­ stage, an analogy with the initial forms for articulate ched bulge), the infrabasals which can be seen on the crinoids (possibly, with Encrinidae). Further, the calyx surface, the narrow spot of the crown on the nodal plates on which cirri can developed subse­ radial = brachial suture. The newly-gained feature is quently are laid under the calyx in pentacrinoid stage. the only one : the nodals in proximal stem part with We can see here the transition from Encrinidae to short cirri (mostly not more than three in a whorl). Pentacrinida. The cirri develop on all the proximal The recent Proisocrinidae are the analogue for these nodals of pentacrinids. The cirri appear but only on ancient stages. Known finds of young pentacrinids one or two most proximal nodals of comatulids, (Carpentier 1884, pi. 30a, 35, 36) and encrinids (Hag- however some discoidal cirriless columnals which are dorn 1978, Abb. 21 ; 1982, Abb. 18, 19) also show homological to pentacrinid nodals are formed below that pentacrinids have been developed from attached them. Then, the detachment of the comatulid larva from the stem takes place. By that, the free animal had not carried yet that indivisible plate which should be called as a centrodorsal. This very important cir­

* Pentacrinida T o r t o n e s e (1938, p. 171, 177, 212) = Isocrinida cumstance shows that the comatulid centrodorsal was Sie v e r t s -D o r e c k , 1952. formed by some proximal nodal plates (Thomson 1865, p. 536). — 658 —

At the same time, all the Thiolliericrinidae without Thus, the hypothesis on the origin of the Thiollieri­ exception possessed of a centrodorsal. And what is crinidae from comatulids and that on the origin of the more, this plate assumes the relatively greater size in latter, in its turn, from pentacrinids seems to be the young specimens (fig. 8 E, 9 C). This indicates that most substantiated ones. Thiolliericrinidae originated from the forms having a The explanation of the origin of Thiolliericrinidae well developed large centrodorsal, and Comatulida - from the family Solanocrinitidae seems to be most from the forms with the centrodorsal consisting of convincing. Adutl individuals of this latter family several plates. have large centrodorsals where, under each radial, two vertical rows of the cirrus sockets are placed. Paracomatula Helvetica (H e s s , 1951, fig. 1—7) has the centrodorsal consisting of several nodal columnals Each row can contain up to five sockets and more. The basais are well visible on the calyx surface, the and crown already typical for comatulids, and serves radiais have high external surface and the centrodor­ a good example for the forms transitional from Pen- sal basis is flattened and smooth. In the enumeration tacrinida to Comatulida (see fig. 18). Carpenterocri- of these features one can see an evident coincidence nus mollis (C a r p e n t i e r , 1884, p. 338, pi. 33, fig. 7- with the calyx construction of Solonaerium, the first 10) can be an example of such transitional forms and most primitive genus of the family Thiolliericrini­ among recent crinoids. dae (fig. 18).

8 — DEVELOPMENT OF THE THIOLLIERICRINIDAE

During settling of the reefs, one of the groups of the proterogenetic evolution from the Solanocrinitacea » order Comatulida (possibly Solanocrinitidae) obtai­ (Rasmussen 1978b, p. 314). Thiolliericrinidae are evi­ ned there the most favourable conditions for its life dently not an unique example of the conservation of (pure water, an abundance of the light, heat, oxygen, larval features by adult animals adapted to the life in food). There was no necessity to change the place of the reef conditions. life, and therefore the moment of the larval stem tea­ The morphological variations of Thiolliericrinidae ring off began to be delayed. The individuals of Solo­ calyxes are presented on fig. 18, according to their naerium and Thiolliericrinus were free when they supposed phylogenetic correlation. became adult. After detaching from a stem these forms lived freely using their powerful centrodorsal Solonaerium seems to be the initial form with high cirri to attaching to the bottom (see fig. 16). There­ radiais, large basais and a regular disposition of the fore, it seems to me that the determination of Thiol­ cirrus sockets. Subsequently the disposition of the liericrinus as « a permanent larval form » (Carpenter sockets lost the regularity but the basais were still pre­ 1881b, p. 377) is quite good. served (Thiolliericrinus). The radiais can be high or low. Then the number of the cirrus sockets decreased, The sessile mode of life appeared so successful, at some sockets turned into structureless fossae. Later the time, that it began to be the cause of subsequent the sockets disappeared completely, but the fossae morphological adaptations of the skeleton. The cirri which had been scattered on the centrodorsal remai­ began to be lost as useless now (Gislén 1924, p. 206 ; ned (Burdigalocrinus). The full disappearance of the Rasmussen 1978a, p. 273), the stem became more cirri on the centrodorsal, with the preservation of powerful and more flexible. Here we witness the high radiais and large basais, is typical for Conoideo­ change from cirriferous forms to Burdigalocrinus and crinus. The centrodorsal fossae can remain on the Umbocrinus, having rudimentary cirrus fossae on the radial-centrodorsal suture. The gradual disapearance centrodorsal, and then to Conoideocrinus and Heber­ of these fossae can be observed simultaneously with ticrinus with no traces of cirri. The basais and partly the atrophy of the basais and the preservation of the the radiais were reducing (Bather 1900, p. 195 ; Kirk high radiais (Loriolicrinus). Otherways, fossae and basais could remain (and even decrease in size) with 1912, p. 78). The crown became more compact and the considerable reduction of the radial ring (Umbo­ more strongly constructed (see fig. 16). The animals crinus). The final disappearance of the fossae and the were attached during the whole life. By that, the basais, and the decrease of the relative size of radiais, attachment organs (the stem and the distal disc) were are marked in the genus Heberticrinus in which the similar to the larval structures of the « true » comatu­ principal morphological types can be determined lids. Just so « Thiolliericrinidae appear to be a according to the form and size of the centrodorsal. i. 8 A cee f h oii ad h dvlpet f an opooia tps f h Tilirciia clxs te otd arrows dotted (the calyxes Thiolliericrinidae the of types morphological main of development the and origin the of scheme A — 18 Fig. JURASSIC L. CRETACEOUS - B P - P céa e ’rgn e d dvlpeet e tps opooius rniax e cue drae ds Thiolliericrinidae. des dorsales coupes des principaux morphologiques types des développement du et l’origine de Schéma yblz te otne idpnet development). independent continued the symbolize Pentacrinus, Burdigalocrinus, Pc - Pc - С Paracomatula, Conoideocrinus, St - St Solanocrinites, - L Loriolicrinus. 69 — 659 — S - S Solonaerium, T - T Thiolliericrinus, U - U Umbocrinus, H - H Heberticrinus,

— 660 —

Acknowledgements

This article was written in the Historical Geology Chair of S.Y. Lobachiova. The information on the Czecho-Slovak the Leningrad Mining Institute. A part of the collections ser­ Thiolliericrinidae was received from Dr. J. Zitt (Mnicho- ving the materials for this work was handed to the author by vice, Czechoslovakia). The manuscript was reviewed by Dr. Dr. T.N. Bogdanova and Dr. S.V. Lobachiova (All-Union A.S. Tarakanov (All-Union Geological Institution, Lenin­ Geological Institution, Leningrad). The author consulted grad) and by Dr. S.N. Alekseev (All-Union Geological them on the Lower Cretaceous stratigraphy of the Crimea, Petroleum Institution, Leningrad). The translation of the Dr. B.T. Yanin (Moscow University) on the stratigraphical article into English was made by Mgr. Igor F. Naftul’eff and range of late Jurassic and early Cretaceous rudistids. The R.D. Kirpichnikov. The author thanks Prof. M. Roux for brachiopods from some localities were determined by Dr. his help in supervising the manuscript.

REFERENCES

ARENDT Y.A. (1974) - Sea-lilies Cyrtocrinids. Trudy ÉTALLON A. (1857) - Esquisse d’une description géologi- Paleontologicheskogo Instituta, Moscow, 144, 250 p. (in du Haut-Jura, et en particulier des environs de St- Russian). Claude. Ann. Sci. phys. natur. agricult. industr., Lyon, (3), 1 : 247-354. ARENDT Y.A. & YANIN B.T. (1964) - On Late Jurassic and Early Cretaceous crinoids of the Crimea. J. Paleon- ÉTALLON A. (1859) - Études paléontologiques sur le Haut- tol., Moscow, 3 : 140-142 (in Russian). Jura. Rayonnés du Corallien. Mém. Soc. Emul. Doubs, BATHER F.A. (1900) - The Echinoderma. In : Lankester Besançon, (3), 3 : 401-553. E.R., A Treatise on Zoology, pt. 3, London, 344 p. ÉTALLON A. (1862) - Études paléontologiques sur le Haut- CARPENTER P.H. (1881a) - On the genus Solanocrinus Jura. Monographie du Corallien. Mém. Soc. Emul. GOLDFUSS, and its relations to recent Comatulae. J. Doubs, Besançon, (3), 6 : 53-260. Linn. Soc. London, Zool., 15 : 187-217. GISLÉN T. (1924) - Echinoderm Studies. Zool. Bidrag CARPENTER P.H. (1881b) - Interesting new Crinoids. Uppsala, 9 : 330 p. Nature, London, 23 : p. 377. GOLDFUSS A. (1826-1833) - Petrefacta Germaniae, Tl.l CARPENTER P.H. (1884) - Report on the Crinoidea collec­ Düsseldorf, 252 p. ted during the Voyage of H.M.S. Challenger. Stalked Crinoids. Report on scientific results o f the Voyage o f HAGDORN H. (1978) - Muschel-Krinoiden Bioherme im H.M.S. Challenger, Zoology, London, 11, 442 p. Oberen Muschelkalk (mo 1 Anis) von Crailsheim und Schwabisch Hall (Südwestdeutschland). N. Jb. Geol. CLARK A.H. (1913) - Post-Paleozoic Crinoidea. In : Text­ Palaont. Abh., Stuttgart, 156, 1 : 31-86. book o f Paleontology edited by C.R. Eastman, adapted from the german of K.A. Zittel, vol. 1, London - New HAGDORN H. (1982) - Chelocrinus schlotheimi (QUENS- York : 225-242. TEDT) 1835 aus dem Oberen Muschelkalk (mo 1, Ani- sium) von Nordwestdeutschland. CLARK A.H. (1921) - A monograph of the existing crinoids Verôff. Naturk. Mus. 4 : 5-33. vol. 1. The Comatulids, pt. 2. Bull. U.S. Nat. Museum, Bielefeld, Washington, 82, 795 p. HAGDORN H. (1983) - Holocrinus doreckae n.sp. aus dem CLARK A.M. (1973) - Fossil and recent Comatulid Crinoids Oberen Muschelkalk und die Entwicklung von Soil with coelomic extensions penetrating the centrodorsal. bruchstellen im Stiel der Isocrinida. N. Jb. Geol. Bull. Brit. Mus. (Natur. Hist), Zool., London, 24, 9 : Palaont. Mh., Stuttgart, 6 : 345-368. 441-446. HESS H. (1951) - Ein neuer Crinoide aus mittleren Dogger DACQUÉ E. (1933) - Wirbellose des Jura. In : Gtirich G., der Nordschweiz (Paracomatula Helvetica n.gen. n.sp.). Leitfossilien, Berlin, 7, 1.1., 272 p. Eel. geol. Helv., Basel, 43, 2 : 208-216. — 661 —

HESS H. (1975) - Die fossilien Echinodermen des Schweizer OOSTER W.A. (1871) - Die organischen Reste der Pteropo- Juras. Verôff. Naturhist. Mus. Basel, 8, 130 p. denschicht einer Unterlage der Kreideformation in den JAEKEL O. (1918) - Phylogenie und System der Pelmato- Schweizer-Alpen. Protozoe Helvetica, Basel, 2 : 89-151. zoen. Palaeontol. Zeitsch., Stuttgart, 3, 1 : 1-128. ORBIGNY A. D’ (1840) - Histoire naturelle générale et particulière des Crinoïdes vivants et fossiles comprenant JOUKOVSKY E. & FAVRE J. (1913) - Monographie géolo­ la description zoologique et géologique de ces animaux. gique et paléontologique du Salève (Haute-Savoie, Paris, 98 p. France). Mém. Soc. Phys. Hist, natur. Genève, 37, 4 : 295-523. PATRULIUS D. (1956a) - Pozitia sistematica a formei Pseu- dosaccocoma strambergensis REMES, crinoid aberant al KIRK E. (1912) - The structure and relationships of certain malmului superior. Bul. stiint. Acad. Republ. Popul. Eleutherozoic Pelmatozoa. Proc. U.S. Nation. Mus., Romine, Bucuresti, 1, 3-4 : 187-195. Washington, 41 : 1-137. PATRULIUS D. (1956b) - Notà asupra unei faune lusita- LORIOL p. DE (1877-1879) - Monographie des Crinoïdes niene din regiunea vaii Casimcea (Pestera de la Gura fossiles de la Suisse. Mém. Soc. paléontol. Suisse, Dobrogei). Bull, stiint. Acad. Republ. Popul. Romine, Genève, 4-6, 300 p. Bucuresti, 1, 3-4 : 197-200. LORIOL P. DE (1880) - Description de quatre Echinodermes PICTET F.J. (1868) - Étude provisoire des fossiles de la nouveaux. Mém. Soc. paléontol. Suisse, Genève, 7 : 5- Porte-de-France, d’Aizy et de Lémenc. Mélanges paléon­ 15. tologiques, Genève, 1, lv.4 : 207-309. LORIOL P. DE (1882-1884) - Crinoïdes. In : Paléontologie PILLET L. & FROMENTEL E. (1875) - Description géolo­ française, Terrains oolitiques et jurassiques, Paris, 11, 1, gique et paléontologique de la colline de Lémenc sur 627 p. Chambéry. Mém. Acad. Savoie, Chambéry, (3), 4 : 135 LORIOL P. DE (1884-1889) - Crinoïdes. In : Paléontologie P- française, Terrains oolitiques et jurassiques, Paris, 11, 2, QUENSTEDT F.A. (1852) - Handbuch der Petrefacten- 580 p. kunde. Tübingen, 792 p. LORIOL p . DE (1888) - Description des Échinodermes. In : QUENSTEDT F.A. (1858) - Der Jura. Tübingen, 842 p. Recueil d’études paléontologiques sur la faune crétaci- que du Portugal, Lisbonne, 2, 2, 122 p. QUENSTEDT F.A. (1876) - Petrefactenkunde Deutschland, 4, Echinodermen, Leipzig, 742 p. LORIOL p . DE (1891) - Description de la faune jurassique du Portugal. Embranchement des échinodermes, Lis­ RASMUSSEN H.W. (1961) - A Monograph on the Creta­ bonne, 179 p. ceous Crinoidea. Biol. Skr. Dansk. Vid. Selsk, Koben- havn, 12, 1, 428 p. LYSSENKO N.I. & VAKRUSHEV B.A. (1974) - On mode of occurrences of Lower Cretaceous deposits on the nor­ RASMUSSEN H.W. (1978a) - Articulata. Postlarval onto­ thern slope of Chatyrdagh (Crimea). Izvestia ANSSSR, geny of fossil Crinoids. In : Treatise on Invertebrate Moscow, geol. ser., 4 : 148-150 (in Russian). Paleontology, Boulder, Colorado and Lawrence, pt. T. Echinodermata 2, 1 : 268-274. M’ COY F. (1848) - On some new Mesozoic radiata. Ann. Mag. Natur. Hist., London, (2), 2 : 397-420. RASMUSSEN H.W. (1978b) - Evolution of Articulate Crinoids. In : Treatise on Invertebrate Paleontology, M’COY F. (1854) - Contribution to British palaeontology. Boulder, Colorado and Lawrence, pt. T, Echinoder­ Cambridge, 272 p. mata, 2, 1 : 302-316. MÜLLER A.H. (1963) - Lehrbuch der Palaozoologie, Bd. 2. RASMUSSEN H.W. (1978c) - Articulata. In : Treatise on In­ Invertebraten, Th, 3, Jena, 698 p. vertebrate Paleontology, Boulder, Colorado and Law­ MÜLLER A.H. (1977) - Zur ichnologie der subherzynen rence, pt. T, Echinodermata 2, 3 : 813-1027. Oberkreide (Campan). Z. geol. Wiss., Berlin, 5, 7 : 881- REMES M. (1905) - Nachtràge zur Fauna von Stramberg 897. VI. Crinoiden-Asteriden- und Echinoiden-Reste aus dem MÜLLER A.H. (1978) - Lehrbuch der Palaozoologie, Bd. 2 Weissen Kalkstein von Stramberg. Beitr. Palaont. Geol. Invertebraten, Th. 3 (2 Aufl.), Jena, 748 p. Osterr. Ung., Wien und Leipzig, 18 : 59-63. ROMAN F. (1897) - Recherches stratigraphiques et paléon- tologiques dans le Bas-Languedoc. Ann. Univ. Lyon, 34, 345 p. ROUX M. (1978a) - Ontogénèse, variabilité et évolution morphofonctionnelle du pédoncule et du calice chez les Millericrinida (Échinodermes, Crinoïdes). Geobios, Lyon, 11, 2 : 213-241. ROUX M. (1978b) - Ontogénèse et évolution des Crinoïdes pédonculés depuis le Trias. Implications océanographi­ ques. Thèse Univ. ParisXI, n° 2082, 167 + 24+ 14 p., 51 fig. (inédit). SCHLÜTER C. (1878) - Ueber einige astylide Crinoiden. Zeit. deutsch. geol. Ges., Berlin, 30 : 27-66.

TERMIER H. & TERMIER G. (1949) - Hiérarchie et corré­ lation des caractères chez les Crinoïdes fossiles. Bull. Serv. Carte géol. Algérie, Paléontologie, Alger, (1), 10 : 1-90. THOMSON W. (1865) - On the Embryogeny of Antedon rosaceus, LlNCK (Comatula rosacea of LAMARCK). Phil, trans. Roy. Soc. London, 155 : 513-544.

THURMANN J. & ÉTALLON A. (1862) - Lethaea bruntru- tana. Nouv. mém. Soc. helv. sci. natur., Zürich, 19 : 144- 353.

TORTONESE E. (1938) - Gli Echinodermi del Museo di Torino, pt. 4. Oloturoidi e Crinoidi. Boll. Mus. Zool. anat. comp. Univ. Torino, (3), 46, 82 : 169-221. UBAGHS G. (1953) - Classe de Crinoïdes. In : J. PlVETEAU, Traité de Paléontologie, Paris, 3, 658-773. UBAGHS G. (1978) - Skeletal morphology of fossil Cri­ noids. In : Treatise on Invertebrate Paleontology, Boul­ der, Colorado and Lawrence, pt. T, Echinodermata 2, 1 : 58-216.

YANIN B.T. (1985) - Rudistids Matheronia from the Lower Valanginian of the Crimea. Paleontol. Zbornik, Lvov, 22 : 25-30 (in Russian).

Manuscrit déposé le 15.01.1987 Manuscrit définitif reçu le 02.06.1987 PLATE PLATE 1

Fig. 1-3 —Burdigalocrinus maximus nov. sp. 1. CK-81-3 : two radiais, view from above (x 5,0). Deux radiales, vue en dessus. 2. Same, view from below (x 5.0). Les mêmes, vue en dessous. 3. CK-81-4 : articular face of a columnal (x 3.1). Facette articulaire d’un article. Crimea, Belbek Valley : Berriasian.

Fig. 4,5 —Loriolicrinus asper nov. sp. 4. CK-99-1 : articular face of an oblique columnal (x 3.4). Facette articulaire d’un article oblique. 5. CK-99-4 : articular face of a columnal (x 5.0). Facette articulaire d’un article. Crimea, Belbek Valley ; Berriasian.

Fig. 6 — Loriolicrinus laevis nov. sp. 6. CK-90-9 : articular face of a columnal (x 4.7). Facette articulaire d’un article. Crimea, Belbek Valley : Berriasian.

Fig. 7-11 —Umbocrinus umbonatus nov. sp. 7. Holotype CK-89-1 : calyx, side view (x 6,9). Coupe dorsale, vue latérale. 8. CK-89-3 : calyx without two radiais (x 6.1). Coupe dorsale sans deux radiales. 9. CK-89-23 : centrodorsal, view from above (x 3.6). Centrodorsale, vue en dessus. 10. Same, view from below (x 3.6). La même, vue en dessous. 11. CK-89-24 : centrodorsal, view from above (x 3.8). Centrodorsale, vue en dessus. Crimea, Belbek Valley : Berriasian.

Fig. 12-16 —Heberticrinus heberti nov. sp. 12. CK-51-69 : centrodorsal, view from above (x 3.2). Centrodorsale, vue en dessus. 13. CK-51-71 : centrodorsal with basais, view from above (x 4,7). Centrodorsale avec les basales, vue en dessus. 14. CK-51-319 : articular face of an oblique columnal (x 5.2). Facette articulaire d’ui) article oblique. 15. CK-51-315 : articular face of a proximal cruciate columnal (x 4.1). Facette articulaire d’un article proximal à crê tes fulcrales croisées. 16. CK-51-313 : articular face of a distal cruciate columnal (x 3.4). Facette articulaire d’un article distal à crêtes fui craies croisées. Crimea, Belbek Valley : Berriasian. Geobios Pl. 1 n° 20, fasc. 5 V.G. Klikushin