thiolliericrinid crinoids from the lower cretaceous of crimea

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

THIOLLIERICRINID CRINOIDS FROM THE LOWER CRETACEOUS OF CRIMEAbyVladim ir G. KLIKUSHIN *AbstractThe family Thiolliericrinidae (Oxfordian - Hauterivian)includes the stalked crinoids of the order Comatulida,inhabiting reef fades. Remains of 12 thiolliericrinidspecies (10 among them are new) belonging tosix genera (Thiolliericrinus, Burdigalocrinus, Loriolicrinus,Conoideocrinus nov. gen., Umbocrinus nov.gen., and Heberticrinus nov. gen.) from the Berriasianand Valanginian of Crimea are described. On thebasis of an extensive material (hundreds of calyxes,centr odor sals, brachials and columnals) the skeletalmorphology, the ontogenetic variations of the calyxesand the phylogeny of the Thiolliericrinidae arereconstructed. The family taxonomy is defined moreexactly regarding new morphological data.RÉSUMÉLa famille Thiolliericrinidae (Oxfordien - Hauterivien)comprend les crinoïdes pédonculés de l’ordreComatulida, vivant dans des milieux récifaux. 12espèces (dont 10 sont nouvelles) de Thiolliericrinidaesont décrites du Berriasien et du Yalanginien de laCrimée. Ces espèces appartiennent à six genres :Thiolliericrinus, Burdigalocrinus, Loriolicrinus,Conoideocrinus nov. gen., Umbocrinus nov. gen. etHeberticrinus nov. gen. L’étude d’un matériel abondant(des centaines de coupes dorsales, de pièces centrodorsaleset brachiales, de columnales) permet unediscussion sur la morphologie du squelette, les variationsontogénétiques des coupes dorsales et la phylogéniedes Thiolliericrinidae. D ’après les nouvellesdonnées morphologiques la taxonomie de la familleest précisée.KEY-WORDS : CRINOIDEA, ARTICULATA, COMATULIDA, THIOLLIERICRINIDAE, TAXONOMY, NOMENCLATURE, MOR­PHOLOGY, ONTOGENY, PHYLOGENY, CRETACEOUS, USSR.MOTS-CLÉS : CRINOÏDES, ARTICULATA, COMATULIDA, THIOLLIERICRINIDAE, TAXONOMIE, NOMENCLATURE, MOR­PHOLOGIE, ONTOGENIE, PHYLOGENIE, CRÉTACÉ, URSS.* Leningrad Mining Institute, 199026 Leningrad, USSR.Geobios, n° 20, fasc. 5 p. 625-665, 18 fig., 2 tab l., 1 pl. Lyon, octobre 1987

TABLE OF CONTENTS1 - Introduction........................................................p. 6262 - Localities and materials.....................................p. 6263 - Term inology........................................................p. 6294 - Taxonomie description....................................p. 631Genus Solonaerium E t a llo n emend.Ra s m u s s e n ...........................................................p. 633Genus Thiolliericrinus E t a l l o n .....................p. 634Genus Burdigalocrinus Ja e k e l .......................p. 639Genus Loriolicrinus J a e k e l .............................p. 641Genus Conoideocrinus nov. gen.......................p. 644Genus Umbocrinus nov. gen.............................p. 646Genus Heberticrinus nov. gen...........................p. 6465 - Reconstruction of the skeleton..................... ..p. 652Root formations ............................................... ..p. 652Stem ......................................................................p. 653A rm s .................................................................... ..p. 6536 - Ontogenetic variations of the calyxes ......... p. 6567 - Origin of the Thiolliericrinidae..................... p. 6578 - Development of the Thiolliericrinidae......... p. 658Acknowledgements.............................................. p. 660R eferences.............................................................. p. 6601 — INTRODUCTIONSea-lilies of the Comatulida are freely mobile formswhich are widely distributed in the modern seas.Calyxes of these crinoids, consisting of five radiais,five basais and one centro-dorsal plate bearing cirri,are also found in the fossil record.However, one peculiar exception exists in that largegroup : the fossil family Thiolliericrinidae. The representativesof this family were stalked and led a fixedlife. The thiolliericrinid thecae were constructed likethe true Comatulida, but their columns showsimularities with the Bourgueticrinida stems. Suchcombination of the features lead to complicated problemsfor attempts to clear up the phylogenetical connectionsof the family and attracted many researchworkers to the study of this group.Nevertheless, many aspects remain in the nomenclature,the taxonomy and the phylogeny of the familyThiolliericrinidae. The abundant new material at theauthor’s disposal allows to propose a solution of theseproblems.2 — LOCALITIES AND MATERIALSRemains of the Thiolliericrinidae were found insome regions of the Crimea (fig. 1). The Belbek Val-'/ley is the most important among them according tothe quantity of found specimens and to the diversityof forms (table 1). In that locality, crinoids under discussionwere found in the Upper Berriasian coralalgalor coral-sponge bioherms. Three types of themcan be distinguished :- 1 : the « yet-formed » bioherms. They are smalllenses (not more than one metre thick and up to 10 mlong) enclosed into the strata of the microphytolithicor bioclastic stratified limestones. The lenses are builtwith skeletons of colonial and solitary corals, calcareousalgae and sponges. Oyster shells (Lopha,Exogyra and others), brachiopod shells, echinoid spines(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 inthese bioherms ;- 2 :« barrier » reefs. They have large volumes (upto 15 m thick and several hundred metres long), builtof big nodules of the calcareous algae and, rarely,colonial corals. Here, in the reef rocks, lenses of thecoarse-grained sandstones and « pockets » saturatedby the quartz pebble are frequent. One can note,among reef-dwellers (besides above mentioned oysters),Prohinnites, Trichites, Heterodiceras and alsonerineid gastropods and so on. The locality on theChuku Mountain is the example of such « barrier »reef. Rare columnals and calyxes of Heberticrinusheberti were found here ;- 3 : «tower-like » bioherms, are the intermediatetype between the two previous ones. They show a discoidshape with an average size (the height is not morethan 7-8 m with the diameter up to 15-20 m) the frameworkof these bodies is built by cloddy masses,consisting of skeletons of colonial hexacorals, hydrozoansand calcareous algae. The interstices betweenbodies of the reef-forming organisms are filled withloose calcareo-argillaceous masses. Therefore the characteristictube-like forms appear in the process ofweathering of these bioherms. Small oysters, brachiopods(Craniscus sp., Ismenia cf. perilustrus Sm ir ­n o v a and others), calcareous sponges, bryozoans,small solitary corals, serpulids and others organismsalso are among the reef-dwellers. Traces of boring

— 628 —Name of s p ecies L o cality No. on f i g . l Age S keleton p a rts No. c o ll.T R Cd Br С AdT h io llie r ic r in u s belbekensis Belbek 3 В 2 8 2 1 11 - _ OK-85T h io llie r ic r in u s elongatus Belbek 3 В 2 1 - - - - - CX-98T h io llie r ic r in u s torosus Belbek 3 в 2 2 2 1 5 . - - CK-86B urdigalocrinus maxiinus Rodnoie 2 в 2 1 _ 3 _ 4 CK-76,78Belbek 3 в 2 1 3 1 - 19 - CK-81L o rio lic rin u s asper Belbek 3 в 2 _ _ _ _ 68 CK-99K aratlykh 4 в 2 3 1 5 12 27 - CK-82L o rio lic rin u s la e v is Rodnoie 2 в 2 4 _ 1 С К-80Belbek 3 в 2 5 2 11 _ CK-90B airakly b в 2 - - 1 - _ _ *CK-50Tanass 10 в 1 - - 1 - - - *СК-18L o rio lic rin u s p e rfo ra tu s Belbek 3 в 2 2 _СХ-91Karatlykh 4 в 2 1 - - - - - СК-95Conoideocrinus conoideus Belbek 3 в 2 1 - - - - - СК-97Umbocrinus urabonatus Rodnoie 2 в 2 1 СК-75Belbek 3 в 2 27 _ 16 8 53 - СК-83,89K aratlykh 4 в 2 4 - 3 - - - СК-94H eb erticrin u s h e b e r ti Rodnoie 2 в 2 3 _ _ b 2 _ СК-61Belbek 3 в 2 72 1 26 215 163 _ СК-51Karatlykh 4 в 2 5 - 2 36 10 - СК-93Chuku 5 в 2 6 - 1 1 9 - СХ-92Tanass 10 в 1 - 1 1 - - - * ск-зоTanass 10 в 1 1 - - - - - СК-69H e b erticrin u s a f f .h e b e r t i Belbek 3 в 2 3 - 5 - - - ск-эбH eb erticrin u s rem esi B eshterek 8 V 1 4 - - - 9 - CK-5b,57H eberti^* j-jius i r r e g u l a r is Rodnoie 2 в 2 5 _ _ _ _СК-74Belbek 3 в 2 7 - 12 2 27 - СК-88Tanass 10 в 1 2 - 1 - 1 - * СК-22,25T n io llie ric rin id a e in d et. Kuchki i в 2 _ _ 2 4 3 СК-64Rodnoie 2 в 2 _ - 1 5 39 СК-73Belbek 3 в 2 - - - 269 297 15 СК-ЮОK aratlykh 4 в 2 - - - 45 11 3 СК-101Ivlrarnornoie 7 в 1 _ _ - 3 2 - СК-6ьMezhgorie 9 в 2 _ - - - 1 -* СК-37Tanass 10 в 1 ~ - - - 1 - *СК-31Tabl. 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 & Lobachiova(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 » (partiedu squelette) : T - coupes dorsales, R - radiales, Cd - centrodorsales, Br - brachiales, С - columnales, Ad - formations racinales. Lematériel 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 arecommon. Echinoid spines and test debris, starfishplates and crinoid fragments were found in the« tower-like » bioherms. Columnals, thecae and brachialsof several Thiolliericrinidae species were collectedhere. B esides, ra re stem frag m en ts o f« Isocrinus » arzierensis (D e L o r i o l ), Apiocrinitesand the thecae of the true unstalked comatulids werefound.The study of dozens « tower-like » bioherms showsthat Thiolliericrinidae were distributed unevenly.

— 629 —They avoided settling on « yet-formed » bioherms,the height of which was not more than 2 m. Theremains of the crinoids under discussion are almostabsent on the bioherms placed in the limestoneshaving an arenaceous admixture or with then quartzouspebbles. Thiolliericrinidae avoided the bioherms,the main rock-forming component of whichwas the calcareous algae. Under the same conditions,the thiolliericrinid remains are more frequent on biohermshaving many interstices filled with the argillaceousmass. The finds of Thiolliericrinidae are accompanied,as a rule, by the finds of the bulb-like echinoidspines, the spiny asterozoan plates and the dactylatesponges.The remains of the Thiolliericrinidae are foundmore often in the inner, more loose core of a bioherm.However it does not mean that they are absenton the external walls. Moreover, rare columnals andthecae are also found outside the bioherm, in the surroundingrocks. But, in this case, they are oftenstrongly rounded. However, the remains of one speciesare found only in the bedded microphytolithiclimestones. This is Conoideocrinus conoideus.Heberticrinus heberti is a « background » specieseverywhere. But at almost every bioherm there is acharacteristic species, distributed locally. Such isThiolliericrinus belbekensis in the « Belbek » locality(see fig. 1 and table 1), Loriolicrinus asper on theKaratlykh M ountain, Heberticrinus irregularis in theRodnoie environs etc. However frequently H. hebertiis the species which only represents the family on abioherm.Occurrence of the Thiolliericrinidae in the BelbekValley was already published by Arendt (1974, p. 57),Arendt & Yanin (1964, p. 140).Towards the West of the Belbek River, in the ChornayaBasin (see « Rodnoie » in the table 1), theremains of the late Berriasian thiolliericrinids werefound in the paleofacies analogous to the describedabove. However, hydrozoans are the main reefbuildingorganisms here (personal communication ofI.Y. Bugrova), and rudists are usual among reefdwellers(Yanin 1985, p. 25). Besides, some columnalsof Thiolliericrinidae indet. were found in this region(see « Kuchky » on fig. 1 and in table 1) in the organogenouslimestones of the middle part of the Berriasianstage. These limestones are slightly arenaceousand built by crinoidal debris (various species of thegenus Apiocrinites, rarely « Isocrinus » arzierensis(De Loriol), calcareous sponges, oysters, corals(solitary and colonial forms), brachiopods (Rioultinarobusta Sm irno va, Symphythyris arguenensis Moisseev),serpulids etc.Conditions of the existence of Thiolliericrinidae inthe central and eastern Crimea are somewhat different.For example, in the Tass-Kor Ravin (« Mramornoie» - see table 1 and fig. 1), the Lower Berriasianclays occur on the washed out surface of the marbledTithonian limestones and contain small bioherms (thethickness is up to 1 m and the diameter is 2 to 3 m).The main reef-building organisms are various calcareoussponges. Solitary and colonial corals and calcareousalgae are scarce. The dwellers of these tiny reefsare small oysters, brachiopods, tubicolous worms andregular echinoids with large thorn-like spines. Crinoidsare rare. They же Apiocrinites sp., Eugeniacrinitessp. and Isocrinus sp. (only columnals). The stemjoints of Thiolliericrinidae are found here. Occurrenceof crinoids in the Berriasian deposits of theM ramornoie was published by Lyssenko & Vakhrushev(1974, p. 149).In the villages Alekseevka and Krasnossiolovka(« Tanass » - see fig. 1 and table 1), the Thiolliericrinidaeare found in the small bioherms among theLower Berriasian clays and sandstones, which occuron the washed out and extremely rough surface of theTithonian limestones. These bioherms form an almostunbroken horizon, but reach the greatest developmenlon the Tithonian substratum swells probablybeing the initial points of its forming. The biohermsare built by the rolled debris of the basement rocksand by the calcareous alga pellets. Among reefdwellers,there are rare corals, oysters, calcareoussponges, sea-urchins and sea-lilies (Thiolliericrinidaeindet. and Cyrtocrinida indet.).3 — TERMINOLOGYA special terminology, in regard to skeletal elementsof the Thiolliericrinidae, is used in the followingchapters. The principal designations are as below (fig.2).Ad - attachment (terminal disk) ;lAx - primaxillar (the axillar of the first brachialseries) ;IIAx - secondaxillar (the axillar of the second brachialseries) ;

630 —

— 631 —В - basal plate (the basal) ;Be - centrodorsal basis ;IIBr - secondbrachial (the plates of the secondbra chial series) ;С - stem joint (columnal) ;Cd - centrodorsal plate (the centrodorsal) ;Co - opening of the axial canal ;Ct - cavity of the calyx ;F - fossa on the centrodorsal ;Fi - interradial furrow in the theca cavity ;FI - lateral fossa ;Fm - marginal fossa ;Fr - radial furrow in the theca cavity ;R - radial plate (the radial) ;Rf - radial facet (the upper face of the radial) ;Ri - intermediate ridge ;Rm - marginal ridge ;Rs - external (free) face of the radial ;Rt - transversal (fulcral) ridge ;Sc - cirrus socket ;SI - left second brachial series ;Sm - marginal crenulation ;Sr - right secondbrachial series ;St - crenularium on the transversal (fulcral) ridge ;The measurements used for the description of spe -cies are (see fig. 2) :A f - angle of the inclination of a radial facet ;Bf - width of a radial facet ;Dc - diameter of a centrodorsal basis ;Dt - diameter of a theca ;He - height of a centrodorsal ;Hr - height of the external (free) radial face ;Ht - height of a theca ;Lf - length of a radial facet ;In some cases, to show the morphological featuresmore clearly, the relative parameters are used :Hr : H t - relative height of the external radial face ;Bf : Lf - relative width of the radial facet.4 — TAXONOMIC DESCRIPTIONClass CrinoideaSubclass Articulata Miller, 1821Order COM ATULIDA A .H . Clark, 1908Family Thiolliericrinidae A .H . Cla r k, 1908Type genus :Thiolliericrinus Et a llo n, 1859.Diagnosis :The theca consists of five radiais, five basais (ofteninvisible outside) and one centrodorsal. The firstbrachial is axillary. Immovable junctions are absentin the arms. The branching of the arms is isotomous(sometimes slightly asymmetrical) and not less thanthrice-repeated. Secondbrachials and tertibrachialsare closed, forming a massive tube. The centrodorsalwith or without cirri. The short stem consists of severalcolumnals. The columnal articula are bifascial(synartrial). The attachment disk has a shape of thecone-like knot.Generic features : Exposure of the generic featuresfor the family meets a considerable difficulty. Thefact is that the representatives of this group, just asmost reef-dwellers, have many variations concealingFig. 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 (sideview) ; С - separate radial (view from above) ; D - calyx (view from below) ; E - calyx (view from above) ; F - relative partial reconstructionof the skeleton ; G - cruciate columnal (the arrows mark the direction of the fulcral ridge) ; H - parallel columnal (thearrows 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 lessurfaces d’articulation des cirres sur la centrodorsale (vue latérale) ; С - radiale isolée (vue en dessus) ; D - coupe dorsale (vue endessous) ; E - coupe dorsale (vue en dessus) ; F - reconstruction partielle conventionnelle du squelette ; G - columnale à crêtes fulcralescroisées (les flèches marquent la direction de la crête fulcrale) ; H - columnale à crêtes fulcrales parallèles (les flèches commela fig. G) ; I - facette articulaire d’un article.Explications dans le texte.

— 632 —not only specific but also generic limits. However, onecan determine the main generic features, being guidedDy phylogenetical considerations (see below), as follows:1 - The degree of the development of cirrus socketson a centrodorsal, namely : a - they are functionaland are distributed regularly (Solonaerium) or irregularly(Thiolliericrinus) ; b - there are no true sockets,but structureless fossae scattered over the centrodorsal(Burdigalocrinus) or remained only on thesuture between radiais and the centrodorsal (Umbocrinus,partly Loriolicrinus) ; the calyx is deprived ofany traces of cirri (Conoideocrinus, Heberticrinus) ;2 - The degree of the development of radiais,namely : radiais have a high external face and nearlyhorizontal muscular facets (Solonaerium, Burdigalocrinus,Conoideocrinus, Loriolicrinus, partly Thiolliericrinus); radiais are low, their facets are stronglyinclined outside and bordered close to a centrodorsal(Heberticrinus, Umbocrinus, partly Thiolliericrinus) ;3 - The degree of the development of basais,namely : a - basais are large (Solonaerium, Conoideocrinus,partly Thiolliericrinus and Burdigalocrinus) ;b - basais are small, hardly visible or not visible on theexternal calyx surface (Loriolicrinus, Umbocrinusand Heberticrinus).Other features (size ratio of the radial ring and thecentro-dorsal, the stem facet construction etc.) arepresent in the generic diagnoses given below besidesthe enumerated characteristics. These features, however,have no importance. And what is more, theabove mentioned generic features, can neither separately,determine a genus in the family Thiolliericrinidae.However, they show quite clearly the differencesbetween the genera being presented as a table (table2).Intraspecific variation. The form and relative sizeof the centrodorsal plate are the most variable featuresof Thiolliericrinidae. These variations are markedmainly in cirriless forms. For example Heberticrinusheberti may have the centrodorsal height less thanthat of the radial ring, equal to it or several timesgreater. Besides, the place of the greatest diameter issituated in the base of the low centrodorsal or in theupper part of high one, directly below the radial ring.The centrodorsal diameter can be equal to that ofradial ring among specimens with an identical size butthe first diameter can exceed almost two times thesecond one. Heberticrinus heberti has a commonlyround centrodorsal outline. However, specimens withan oval centrodorsal can be found, lenghtening ofwhich coincides with a direction of the fulcral ridgeon its basis. Sometimes the rounded-pentangular outlineis marked in large specimens.The stem facet can be greatly deepened on the lowercentrodorsal face of H. heberti (this is typical forrather low calyxes), can be flat (in low calyxes) or canproject as an acute ridge (in high calyxes). The centrodorsalbasis is mostly horizontal but it can be placedobliquely with the calyx axis. Because of this, the centrodorsalcan be considerably higher on one side thanon another.The radial facets of H. heberti more often adjoindirectly the centrodorsal. But in some calyxes, onecan see a narrow strip of the free radial surface dividingthe facet from the centrodorsal. The width ofthis strip can varied, though in a small degree, even ondifferent sides of the same specimen.The size of basal plates is also subjected to a variation.Being concealed under the radiais, they neverhave an equal length. Some basais, even in one calyx,hardly reach a half of the distance from the centre tothe outer edge while the others can appear upon theexternal calyx surface. These variations are marked inHeberticrinus heberti and Burdigalocrinus maximus.The basais of Umbocrinus umbonatus are quite visible,as a rule, on the calyx surface. However, the caseswere marked, when some basais of individual specimensare concealed under the radiais.The sculpture of the external calyx surface is therelatively insignificantly variable feature. For example,the calyxes of Thiolliericrinus belbekensis arecovered with small granules and the radiais of T. torosusbear big knobs. However, a weak sculpture in theform of small irregular granules is observed in somespecimens of Umbocrinus umbonatus, for which thesmooth calyxes are typical.One must emphasize the difference between theintraspecific and the intrapopulational variations.The specimens which belong to the same species arerelatively poorly differ in each of the local population.At the same time, one can hardly found theforms which would be united under the same specificname even the two bioherms are placed side by side.Seven genera : Solonaerium Éta llo n, 1862 emend.Rasm ussen, 1978, Thiolliericrinus Éta llo n, 1859,Burdigalocrinus Jaekel, 1918, Loriolicrinus Jaekel,1918, Conoideocrinus nov. gen., Heberticrinus nov.

— 633 —gen. and Umbocrinus nov. gen. Each of them is determinedby peculiarities of the calyx construction.However, there are some species, belonging evidentlyto the family Thiolliericrinidae, for which only thecolumnals are known. These forms, mentionedusually under the generic name Apiocrinites or Bourgueticrinus,are placed below. Their taxonomic positionneeds a more accurate definition.hlajue of genus 3c P В Hr: Ht Bf :Lf Af°Solonaerium + - + 0,24 0,54 38T h io llie ric rin u s + + + С, 20 0,42 42Burdigalocrinus - + + 0,35 0,39 40Umbocrinus - + + 0,17 0,56 50L o rio lic rin u s - + + 0,35 0,57 37Oonoideocrinus - - + 0, bl 0,35 26H eberticrinus - - + 0,12 0,47 42Tabl. 2 — Main diagnostic features of the genera of the familyThiolliericrinidae. The designations (see fig. 2) : Sc - cirrussockets (yes - no), F - fossae on the centrodorsal (yes -no), В - basais (visible on the calyx surface or not visible),Hr :Ht - average relative height of the free radialsurface, Bf : Lf - average relative width of the radialfacet, Af° - average angle of the inclination of the radialfacet (in degrees).Caractères principaux de détermination des genres de lafamille Thiolliericrinidae. Légende (voir fig. 2) : Sczone d’insertion des cirres (présence - absence), F - fossessur la centr odor sale (présence - absence), В - basales(visibles sur la surface de la coupe dorsale ou non visibles),Hr : Ht - hauteur relative moyenne de la surfacelibre de la radiale, Bf : Lf - largeur relative moyenne dela facette radiale, Af° - angle moyen de l’inclinaison dela facette radiale (en degrés).Apiocrinites flexuosus G o l d f u s s (1826-1833, p.186, pl. 57, fig. 4) (= Bourgueticrinus flexuosusd ’ORBlGNY, 1840, p. 96-98, pl. 17, fig. 13-15) - Kimmeridgianof Germany (see also : Quenstedt 1852, p.612, pi. 53, fig. 17 ; 1876, p. 367, pi. 104, fig. 57, 58 ;Gislén 1924, p. 186).Bourgueticrinus oosteri D e L o r i o l (1879, p. 188,pi. 18, fig. 6,7) (= Bourgueticrinus flexuosus Ooste r , 1871, p. 130, 142, 149, pi. 19, fig. 1 and 23) -Lower Neocomian of Switzerland (see also : Rasmussen1961, p. 168).Bourgueticrinus sp. (Quenstedt 1876, p. 367, 431,pi. 106, fig. 43, 44) - Callovian of Germany.Bourgueticrinus sp. (Termier & Termier 1949, p. 57,86, pi. 6, fig. 8-14) - Upper Jurassic of Morocco (seealso : Ubaghs 1953, p. 716, fig. 101 and Müller 1963,p. 327, fig. 443 g ; 1978, p. 360, fig. 432 g - sub Balanocrinus,lapsus calami).Besides these above mentioned species, there is anotherone which was placed by many authors in thefamily Thiolliericrinidae (De Loriol 1879, p. 195 ;1880, p. 11 ; 1889, p. 559 ; Carpenter 1884, p. 257 ;Gislén 1924, p. 186 ; Roux 1978a, p. 226 ; 1978b, p.A 21 ; Rasmussen 1978c, p. 879) - namely Bourgueticrinusoôliticus. The available morphological andgeological data (M’Coy 1848, p. 405 ; 1854, p. 53), inmy opinion, testify against that interpretation of thespecies.Upper Jurassic (Callovian) to Lower Cretaceous(Hauterivian) ; Portugal, France, Switzerland, Germany,Rumania, Czechoslovakia, Crimea, Caucasus,Pamyrs.Genus Solonaerium Étallo n, 1862 emend.Rasm ussen, 19781862 — Solonaerium Étallon (in Thurm ann &Étallon), p. 341 (errore typographico exSolanocrinus).1978c — Solonaerium Étallon : Rasmussen, p.881.Type species :Comatula sigillata Quenstedt, 1876.Diagnosis :Radiais have a high free external surface. Basais arevisible on the calyx surface. Cirrus sockets are of thesame size in regular vertical rows. There are no fossaeunder radiais. Centrodorsal is high and conical. Stemfacet is small, deepened, pentangular or rounded.One species.Solonaerium sigillatum (Quenstedt, 1876, p. 176,pi. 96, fig. 49,50 sub Comatula ( = Solanocrinites costatusGo ldfuss, 1826-1833, p. 167 ex parte : pi. 50,fig. 2 ; non pi. 50, fig. 7 a,b = type specimen forSolanocrinites costatus ; nec fig. 7 c,d = Solanocrinitesorbignyi ; necdum fig. 7 a,f = ? Solanocrinitescostatus) (Solanocrinus costatus Étallon 1862, p.222 ; Solonaerium costatus Thurmann & Étallon1862, p. 341 ; Solanocrinus sigillatus Schliiter 1878,p .37 ; Comatula sigillata De Loriol 1879, p. 267 ;A ntedon sigillata Carpenter 1881a, p. 201, 202 ;Solonaerium costatum Rasmussen 1978c, p. 881, fig.588-3). Kimmeridgian ; Nattheim (Württemberg),Germany (fig. 3 A,B).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, 18and 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 , 18591857 — Thiolliericrinus : Ét a llo n, p. 301 (nomenpro visorium).1859 — Thiolliericrinus Étallon : Étallon, p. 413,445.1879 — Thiolliericrinus Étallon : De Loriol, p.193.1880 — Thiolliericrinus Étallon : De Loriol, p. 10.1881b — Thiolliericrinus Étallon : Carpenter, p.377.1883 — Thiolliericrinus Étallon : De Loriol, p. 65.1889 — Thiolliericrinus Étallon : De Loriol, p.431, 542.1900 — Thiolliericrinus Étallon : Bather p. 195.1913 — Thiolliericrinus Étallon : A.H. Clark, p.234.1918 — Thiolliericrinus : Jaekel, p .71.1924 — Thiolliericrinus Étallon : Gislén, p. 186.1933 — Thiolliericrinus Étallon : Dacqué, p. 103.1953 — Thiolliericrinus Étallon : Sieverts-Doreck(in Ubaghs), p. 758.1961 — Thiolliericrinus Étallon : Rasmussen, p.212.

1975 — Thiolliericrinus Étallon : Hess, p .69.1978a — Thiolliericrinus Étallon : Roux, p. 234.1978b — Thiolliericrinus Étallon : Roux, p. A 20.1978c — Thiolliericrinus Étallon : Rasmussen, p.879.Type species :Humberticrinus favieri De Loriol ex Étallo n,1879.Diagnosis :Radiais with either high or low free external surface.Basais are visible or not on the calyx surface.The cirrus sockets have different size and are placedirregularly. Some of the sockets are turned into structurelessfossae. The centrodorsal is conical, high orlow. The stem facet is small, flat or convex.Six species : T. arzierensis De Loriol, T. favieri(De Loriol ex Étallon), T. ribeiroi De l o r io l, T.belbekensis nov. sp., T. elongatus nov. sp., T. torosusnov. sp.Upper Jurassic (Oxfordian) - Lower Cretaceous(Valanginian) ; Portugal, France, Switzerland, Crimea.Thiolliericrinus arzierensis De Lo rio l, 1889 (p. 560)(= Antedon valdensis De Loriol 1879, p. 266 exparte : pi. 20, fig. 20, non fig. 19, 32, 33 = Solanocrinitesvaldensis) (Thiolliericrinus arzierensis : Gislén1924, p. 187 ; Rasmussen 1961, p. 212, pi. 35, fig.13 ; Hess 1975, p. 69, pi. 20, fig. 13). Valanginian ;Arzier (Vaud), Switzerland (fig. 3 C).Thiolliericrinus favieri (De L o r io l ex É t a llo n ,1879, p. 194, pi. 18, fig. 8-10 sub Thiolliericrinusflexuosus (= Thiolliericrinus sp. É t a llo n , 1857, p.301 ; = Thiolliericrinus flexuosus : Étallon 1859, p.445 ; Étallon 1862, p. 222, 237 ; De Loriol 1889, p.553 ex parte, pi. 229, fig. 2,4,5,7,8, non fig. 3,6 =Loriolicrinus sp ; Kirk 1912, p. 75, pi. 5, fig. 4 ; Rasmussen1978c, p. 879, fig. 588-2b,c,d non fig. 588-2a= Loriolicrinus sp. ; = Humberticrinus favieriÉ ta llo n : De Loriol 1879, p. 195 ; 1889, p. 556 ;'Thiolliericrinus favieri : Gislén 1924, p. 187 ; Roux1978b, p. A 20 ; non Thiolliericrinus favieri : Ubaghs1953, p. 718, fig. 104 ; 1978, p. 72, fig. 52-2 = Thiolliericrinussp. Bather 1900, p. 195, fig. 117-1). Kimmeridgian(Dicératien) . Valfin (Jura), France (fig. 3D).Thiolliericrinus ribeiroi De lo r io l, 1880 (pi. 11,fig. 1, fig. 3-11) (= Thiolliericrinus choffati Deloriol, 1889, p. 547, 556, nomen provisorium)(Thiolliericrinus ribeiroi : Carpenter 1881b, p. 377 ;De Loriol 1889, p. 557, 558, pi. 229, fig. 9 ; De Loriol1891, p. 165 ex parte, pi. 29, fig. 16, 18-22, non fig. 17= Burdigalocrinus lorioli ; Bather 1900, p. 195, fig.117-2 ; Jaekel 1918, p. 71, fig. 63 ; Gislén 1924, p.187 ; Ubaghs 1953, p. 718, fig. 103 ; 1978, p. 72, fig.52-1 ; Rasmussen 1978c, p. 880, fig. 588-2 g-h ; Roux1978a, p. 234). Lusitanian ; Engenheiro (Ribatejo),Portugal (fig. 3 E).Thiolliericrinus belbekensis nov. sp.D e r iv a t io n o m i n i s :fig. 4 A ,В ; 5 A-FThe name of the species comes from the type locality.H o l o t y p e :CK-85-1 (fig. 5 A) ; Leningrad Mining Institute.T y p e l o c a l it y :Kuibyshevo, Belbek Valley, Crimea.T y p e l e v e l :Upper Berriasian.D e s c r i p t i o n :The calyx is low, subcylindrical with the rounded =pentangular outline. The basais are not visible on theexternal calyx surface. But small triangular plates areobserved in some specimens in some interradia.Basais are rod-like, narrowed towards the outer marginand risen abruptly inside the calyx. The externalsurface of the radiais is low, vertical and slightly convex.It is covered with coarse granules. The radialfacets are wide. The furrow-like depression is placedunder each radial facet on the external radial face.The calyx cavity is broad and shallow. 10 furrows leanbetween the radiais and are fully pronounced on theirmiddle part. The inner radial surface is covered withweak furrows. The basais are visible on the bottom ofthe calyx cavity only in eroded specimens, but arecommonly covered with a porous calcitic plug. Theprimaxillary has a strongly convex uneven externalsurface. Its lower facet is muscular. The second axillaryis very asymmetric with the muscular lower facetand with a strongly convex granulated external

— 636 —Fig. 4A scheme of the disposition of the cirrus sockets and the fossae on centrodorsals of some Thiolliericrinus (x 4). The arrows mark theends of the fulcral ridge.Schéma de la disposition des fossettes et des surfaces d’articulation des cirres sur les centrodorsales de quelques Thiolliericrinus. Lesflèches 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 (holotypeCK-86-1).

— 637 —surface. The height of the centrodorsal is about equalto the height of the radial ring. 2 to 3 cirrus socketsare under each the radiais. 11 to 12 sockets in all.Some of the sockets are replaced by structureless fossae.The free surface of the centrodorsal between thecirrus sockets is granulated like that of the externalradial surface. The lower centrodorsal face, in all examinedspecimens, is poorly preserved. It is convex,ovate-oblong and has a fulcral ridge on whose bothsides two slit-like furrows are marked, surrounding intheir turn by two marginal ridges.D i m e n s i o n s :Ht = 5.5 - 8.4 mm ; Dt = 7.8-11.8 mm ; Dc = 5.3- 9.5 mm ; A f = 50° ; H r : H t = 0.10 - 0.18 ; Bf : Lf= 0.41 - 0.45.C o m p a r i s o n :T. belbekensis resembles T. favieri after its centrodorsalconstruction (one row of sockets, roughly twounder each of the radiais). The species under discussiondiffers from T. favieri having relatively lowradiais, a granulated external surface and fossae on itscentrodorsal.M a t e r i a l s :8 calyxes, 2 radiais, 1 centrodorsal and 11 brachials(see table 1).D i s t r i b u t i o n :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 r iv a t io n o m i n i s :fig. 4 С ; 6 AThe name of the species comes from « elongatus »(elongated in Latin).H o l o t y p e :CK-98-1 (fig. 6 A) ; Leningrad Mining Institute.T y p e l o c a l it y :Kuibyshevo, Belbek Valley, Crimea.T y p e l e v e l :Upper Berriasian.D e s c r i p t i o n :The calyx is high, conical, narrowed downwardswith the rounded-pentangular outline. Basais are visibleon the external calyx surface as large triangular orirregular diamond = shaped plates. The externalradial surface is very high, sloping, flat and smooth.The radial facets are narrow. The calyx cavity isbroad and shallow. 10 furrows lead between theradiais also in their middle part are pronounced onlyin the upper. The interradial furrows are wider anddeeper. The inner radial surface is covered with bentfurrows. The basais are invisible on the bottom of thecalyx cavity. The centrodorsal is lower than the radialring. 1 to 2 large, projected cirrus sockets are situatedunder each the radiais. 10 sockets in all. The centrodorsalsurface is smooth. The lower face of the centrodorsalis poorly preserved. It is convex, ovateoblongand has a fulcral ridge on which both sidesthere are two slit-like furrows surrounded by a weakmarginal ridge.вFig. 6 — Sk eletal parts ofThiolliericrinus (x 3).A - T. elongatus,h o lo ty p e CK -98-1(calyx from side andbelow). В - T. torosus,h o lo ty p e C K -86-1(calyx from side andbelow). С - T. torosus,CK-86-6 (secondaxillaryfrom below).D - T. torosus, CK-86-5 (radial from above).E - T. torosus ?, CK-86-11 (cirral).Éléments du squelettede Thiolliericrinus.A - T. elongatus,h o lo ty p e C K -98-1(coupe dorsale vue decôté et en dessous). В -T. torosus, holotypeCK-86-1 (coupe dorsalevue de côté et endessous). С - T. torosus,CK-86-6 (secondeaxillaire vue en dessous).D - T. torosus,CK-86-5 (radiale vueen dessus). E - T.torosus ?, CK-86-11(cirrale).D i m e n s i o n s :Ht = 9.3 mm ; Dt = 12.4 mm ; De = 6.4 mm ; A f= 45° ; H r : H t = 0.37 - 0.39 ; Bf : Lf : =0.36-0.37.C o m p a r i s o n :T. elongatus is similar with T. rïbeiroi but differsfrom this species, like from other species of thatgenus, having a very high external radial surface and anarrow centrodorsal basis.M a t e r i a l s :One calyx (see table 1).D is t r i b u t i o n :Berriasian ; Crimea.

— 639 —Thiolliericrinus t orosus nov. sp.D e r iv a t io n o m i n i s :figs. 4 D ; 6 B-EThe name of the species comes from « torosus »(pustulous in Latin).H o l o t y p e :CK-86-1 (fig. 6 B) ; Leningrad Mining Institute.Ty p e l o c a l it y :Kuibyshevo, Belbek Valley, Crimea.Ty p e l e v e l :Upper Berriasian.T y p e l e v e l :Upper Berriasian.D e s c r i p t i o n :The calyx is low, conical, abruptly narrowed downwardswith the rounded-pentangular outline. Thebasais are visible on the calyx surface as small convextriangular plates. The external radial surface is verylow, overlapping the centrodorsal, and covered withbig irregular knobs. The calyx cavity is broad andshallow. 10 furrows go between the radiais and arefully pronounced on their middle part. The innerradial surface is slightly striated. The basais are notvisible on the bottom of the calyx cavity. The primaxillaryhas a muscular lower surface. Its externalsurface is tumid, cone-likely and covered with bigknobs. The height of the centrodorsal is about equalto the radial height, but the diameter of the centrodorsalis notably smaller than that of the radial ring.Three cirrus sockets are placed close to each otherunder each radial. Two large sockets are situated nearerto the radial and a small one is placed below, betweenthe large ones. 15 sockets in all. The lower centrodorsalface is convex, ovate = oblong and has afulcral ridge with two narrow interrupted depressionson both sides, surrounded by a weak marginal ridge.D im e n s io n s :Ht = 7.2 mm ; Dt = 10.8 mm ; Dc = 6.0 mm ; A f= 57° ; Hr : H t = 0.07 - 0.11 ; Bf : Lf = 0.44 - 0.55.Co m p a r is o n :The centrodorsal of T. torosus is similar to that ofT. arzierensis but has a considerably greater numberof the cirrus sockets (15 againts 6-8) being placedregularly.M a t e r i a l s :2 calyxes, 1 centrodorsal, 2 radiais, 5 brachials, 4 cirrals(see table 1).D i s t r i b u t i o n :Berriasian ; Crimea.Genus Burdigalocrinus Ja e k e l , 19181918 — Burdigalocrinus : Ja e k e l , p.71.1924 — Burdigalocrinus Ja e k e l : Gislén, p. 188.1933 — Burdigalocrinus Ja e k e l : Dacqué, p. 103.1953 — Burdigalocrinus Ja e k e l : Sieverts-Doreck(in Ubaghs), p. 758.1961 — Burdigalocrinus Ja e k e l : Rasmussen, p.213.1978a —Burdigalocrinus Ja e k e l : Roux, p. 234.1978b —Burdigalocrinus Ja e k e l : Roux, p. A 20.1978c —Burdigalocrinus Ja e k e l : Rasmussen, p.880.T y p e s p e c ie s :Burdigalocrinus lorioli Ja e k e l , 1918.D i a g n o s i s :The radiais have high free external surface. Thebasais are visible on the calyx surface. Cirrus socketsare absent but there are structureless fossae placed, asa rule, regularly. The centrodorsal is high and conical.The stem facet is wide, elliptical and flat.Two species : B. lorioli Ja e k e l , B. maximus nov.sp.Upper Jurassic (Oxfordian) - Lower Cretaceous(Berriasian) ; Portugal, Crimea.Burdigalocrinus lorioli Ja e k e l , 1918 (p. 72, fig. 64D,E) (= Thiolliericrinus ribeiroi D e L o r i o l , 1891, p.165 ex parte, pi. 29, fig. 17, non fig. 16, 18-22 =Thiolliericrinus ribeiroi) (Burdigalocrinus lorioli :Gislén 1924, p. 190 ; Rasmussen 1961, p. 213 ; 1978c,p. 881, fig. 588-4 ; Roux 1978b, p. A 20). Lusitanian ;Engenheiro (Ribatejo), Portugal (fig. 7 A).Burdigalocrinus maximus nov. sp.fig. 7 B,C ; pl. 1, fig. 1-3D e r iv a t io n o m i n i s :The name of the species comes from « maximus »(large in Latin).H o l o t y p e :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 :Rodnoie, Chornaya Basin, Crimea.T y p e l e v e l :Upper Berriasian.D e s c r i p t i o n :The calyx is high, broad-conical, roundedpentangular.The basais are visible on the externalcalyx surface as small convex triangular plates butsometimes not in all interradia. The external surfaceof the radiais is high, convex or flattened and smooth.Two weak fossae are on the upper part of the externalradial surface under the radial facet. The radial facetsare narrow. The calyx cavity is broad and shallow.The radial and interradial furrows are weak. Theinner radial surface is wrinkled. The basais are visibleon the bottom of the calyx cavity. The primaxillary islow with a muscular lower face. The external surfaceof the primaxillary is slightly convex and smooth. Theheight of the centrodorsal is about equal to that of theradial ring but sometimes unequal on opposite sidesof the calyx. 2-4 rounded or high-oval fossae are placedon the centrodorsal under each radial. The lowercentrodorsal face bears a wide-elliptical flat articularface. The fulcral ridge is pronounced. The lateral fossaeare shallow and flat. The fulcral ridge is serratedin some columnals. The marginal ridge is often twinned.The ridges are almost perpendicular to eachother on two sides of one columnal. They are sometimesparallel or turned at an acute angle.

— 641 —Dimensions :Ht = 10.3 - 11.1 mm ; Dt = 13.5 - 16.3 mm ; De =7.2 - 9.8 mm ; A f = 38 - 42° ; Hr : H t = 0,31 - 0.44 ;Bf : Lf = 0.36 - 0.41.Comparison :В. maximus differs from В. lorioli by a massivelarge calyx and a low centrodorsal with the flat basis.Materials :2 calyxes, 2 centr odor sals, 3 radiais, 2 brachials, 23columnals (see table 1).Distribution :Berriasian ; Crimea.Genus Loriolicrinus Jaek el, 1918Loriolicrinus sp. (= Thiolliericrinus flexuosus DeL o r i o l , 1889, pi. 229, fig. 3,6). Kimmeridgian ofFrance.Solanocrinites jaegeri Q u e n s t e d t , 1852 (p. 601, pi.51, fig. 33) (non Solanocrinites jaegeri Goldfuss 1826-1833, p. 168, pi. 50, fig. 9 = Chariocrinus ? jaegeri)(Solanocrinites jaegeri : Quenstedt 1858, p. 723, pi.88, fig. 12 ; 1876, p. 177, pi. 96, fig. 51 . Schliiter1878, p. 36,37 ; Carpenter 1881a, p. 215, pi. 11, fig.22 ; De Loriol 1889, p. 559 ; Gislén 1924, p. 110 ;Thiolliericrinus ? jaegeri Carpenter 1881a, p. 209).Kimmeridgian of Germany.Upper Jurassic (Oxfordian) - Lower Cretaceous(Valanginian) ; Portugal, France, Germany, Crimea.Loriolicrinus insuetus (De L o r i o l , 1891, p. 167, pi.29, fig. 23-29 sub Thiolliericrinus) (Loriolicrinusinsuetus : Jaekel 1918, p. 72, fig. 65 ; Gislén 1924, p.190, 207 . Roux 1978a, p. 234 ; 1978b, p. A 21 ; Rasmussen1978c, p. 881, fig. 588-1). Lusitanian ; Serradas Figueiras (Algarve), Portugal (fig. 8 A).1918 — Loriolicrinus : Ja ek el, p. 72.1924 — Loriolocrinus Jaekel : Gislén, p. 190, 207.1933 — Loriolocrinus Jaekel : Dacqué, p. 104.1978a — Loriolocrinus Jaekel : Roux, p. 234.1978b — Loriolicrinus JAEKEL : Roux, p. A 21.1978c — Loriolicrinus Ja e k e l : Rasmussen, p. 881.Type species :Thiolliericrinus insuetus De Lo rio l, 1891.D e r iv a t io n o m i n i s :Loriolicrinus asper nov. sp.fig. 8 B-F ; pl. 1, fig. 4,5The name of the species comes from « asper »(rough, rugget in Latin).H o l o t y p e :CK-82-1 (fig. 8 D) ; Leningrad Mining Institute.Diagnosis :The radiais have a high free external surface. Thebasais are usually not visible but sometimes they areon the calyx surface. There are no cirrus sockets onthe centrodorsal but sometimes there are small fossaeon the radial-centrodorsal suture. The centrodorsal islow, cylindrical or conical. The stem facet is wide,elliptical, flat or concave.Four species :L. insuetus (De Loriol), L. asper nov. sp., L. laevisnov. sp., L. perforatus nov. sp.One can evidently attribute to the genus Loriolicrinusalso two following species :T y p e l o c a l it y :Karatlykh M ountain, Belbek Valley, Crimea.T y p e l e v e l :Upper Berriasian.D e s c r i p t i o n :The calyx is low, broad-conical, rounded. Thebasais are not visible on the calyx surface. The externalradial surface is very high, convex and coveredwith big irregular knobs and ridges. The radial facetsare wide. The calyx cavity is narrow and deep. Theradial and interradial furrows are well pronounced.The basais are not visible on the bottom of the calyx

— 642 —cavity. The primaxillary is high with vertical, somewhatconvex and tubercular external surface. Theheight of the centrodorsal is smaller than that of theradial ring. The external centrodorsal surface has thesame sculpture as the radiais. A slit-like depression isunder each radial, on the radial-centrodorsal suture.The centro-dorsal basis is wide and flat. In the middleof it, there is a narrow elliptical stem facet. The fulcralridge is sometimes hardly visibly serrated. Themarginal ridge, in most cases, bears big irregularteeth. The columnals are low, parallel, rarely cruciateor oblique. The external columnal surface is smoothor 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 :Ht = 4.5 - 9.5 mm ; Dt = 5.9 - 15.7 mm ; De = 3.7- 10.2 mm ; A f = 41 - 42° ; H r : H t = 0.32 - 0.38 ;Bf : Lf = 0.48 - 0.66.Comparison :L. asper differs from other species of the genus byvery large convex radiais and by its coarse pustulatesculpture.Materials :3 calixes, 5 centrodorsals, 1 radial ring, 12 brachials,95 columnals (see table 1).D e r iv a t io n o m i n i s :Loriolicrinus laevis nov. sp.fig. 9 A-D ; pl. 1, fig. 6The name of the species comes from « laevis »(smooth in Latin).H o l o t y p e :CK-80-1 (fig. 9 A) ; Leningrad Mining Institute.T y p e l o c a l it y :Rodnoie, Chornaya Basin, Crimea.Distribution :Berriasian ; Crimea.T y p e l e v e l :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 :The calyx is low, cylindrical or conical, smooth withrounded outline. The basais are not visible on thecalyx surface. The external radial surface is low andconvex. The radial facets are wide. The calyx cavity isnarrow and shallow. The radial and interradial furrowsare well pronounced. The basais are visible onthe bottom of the calyx cavity. The primaxillary is nothigh with the convex and smooth external surface.The centrodorsal is low. Under each radial, on theradial-centrodorsal suture, there is one small, hardlyvisible fossa. The centrodorsal basis is broad-ellipticaland concave. The stem facet is narrow and elliptical.Often there is the intermediate ridge besides the fulcraland marginal ones. The columnals are parallel orrarely cruciate, and not high.D i m e n s i o n s :Ht = 6.0 - 7.7 mm ; Dt = 10.0 - 11.7 mm ; Dc =6.6 - 7.0 mm ; A f = 36° ; Hr : H t = 0.25 - 0.32 ; Bf :Lf = 0.60 - 0.65.C o m p a r i s o n :L. laevis differs from L. asper by its smooth calyxsurface ; from L. perforatus by the absence of the bigfossae under the radiais and by the absence of thebasais on the external calyx surface ; from L. insuetus(with which the species under discussion has the greatestlikeness) it differs by the radiais of small relativeheight and by the concave centrodorsal basis.M a t e r i a l s :9 calyxes, 3 centrodorsals, 2 brachials, 11 columnals(see table 1).D i s t r i b u t io n :Berriasian ; Crimea.H o l o t y p e :CK-95-1 (fig. 9 E) ; Leningrad Mining Institute.T y p e l o c a l i t y :Karatlykh M ountain, Belbek Valley, Crimea.T y p e l e v e l :Upper Berriasian.D e s c r i p t i o n :The calyx is low, cylindrical or conical, smooth androunded. The basais are visible on the calyx surface.The external radial surface is high and convex. Theradial facets are wide. The calyx cavity is narrow andshallow. The basais are not visible on the bottom ofthe calyx cavity. The centrodorsal is not high. Onelarge fossa, turning downwards, is situated undereach radial on the radial-centrodorsal suture. Thecentrodorsal basis is wide, round and flat. The stemfacet is narrow-elliptical.D i m e n s i o n s :ht = 6.2 - 9.2 mm ; Dt = 10.0 - 12.2 mm ; Dc =7.0 - 8.0 mm ; A f = 36° ; H r : H t = 0.40 . Bf : Lf =0.53.C o m p a r a i s o n :L. perforatus differs from other species of the genusLoriolicrinus having a big fossa under each radial.This species has the great likeness with Solanocrinitesjaegeri Q e n s t e d t (see above) as for the shape of thefossae on the calyx surface. But L. perforatus differsfrom it by a smaller height of the centrodorsal and byrelatively higher radiais.M a t e r i a l s :3 calyxes (see table 1).D i s t r i b u t i o n :Loriolicrinus perforatus nov. sp.Derivatio nominis :fig. 9 EThe name of the species comes from « perforatus »(holey, perforated in Latin).Berriasian ; Crimea.Genus Conoideocrinus nov. gen.Type species :Conoideocrinus conoideus nov. gen., nov. sp.

— 645 —D ia g n o s is :The radiais have high external surface. The largebasais are visible on the calyx surface. The cirrus socketsand fossae are absent. The centrodorsal is smalland conical. The stem facet is small, rounded andflat.T y p e l o c a l it y :Kuibyshevo, Belbek Valley, Crimea.T y p e l e v e l :Upper Berriasian.Co m p a r i s o n :Conoideocrinus differs from other genera of thefamily having high-conical calyx with large basais andwith no trace of the cirrus socket or fossa.One type species.Lower Cretaceous (Berriasian) ; Crimea.D e s c r i p t i o n :The calyx is high-conical, smooth and rounded. Theexternal radial surface is very high. The sutures betweenthe radiais, basais and the centrodorsal arehardly visible. Radial facets are narrow. The calyxcavity is broad and deep. Radial and interradial furrowsare weak. The height of the centrodorsal is smallerthan that of the radial ring. The centrodorsal basisis rounded and flat with a pronounced fulcral ridge.Conoideocrinus conoideus nov. sp.D e r iv a t io n o m i n i s :fig. 10 AThe name of the species comes from « conoideus »(cone-shaped in Latin).D i m e n s i o n s :H t = 7.9 mm ; Dt = 11.0 mm ; Dc = 6.2 mm ; Af= 26° ; Hr : H t = 0.51 - 0.72 ; Bf : Lf = 0.33 - 0.38.M a t e r i a l :One calyx (see table 1).H o l o t y p e :CK-97-1 (fig. 10 A) ; Leningrad Mining Institute.D i s t r i b u t i o n :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 —T y p e s p e c ie s :Genus Umbocrinus nov. gen.Umbocrinus umbonatus nov. gen., nov. sp.D i a g n o s i s :The radiais have low external surface. The basaisare visible on the calyx surface. A row of fossae is placedon the suture between the radiais and the centrodorsal.The centrodorsal is small and convex. Thestem facet is rounded and concave.C o m p a r i s o n :Umbocrinus differs from Solonaerium and Thiolliericrinuslacking cirrus sockets ; from Burdigalocrinusby lower radiais and by the fossae being placed onlyon the radial-centrodorsal suture ; from the similargenus Heberticrinus by the fossae on the calyx and thebasais on the external calyx surface.One type species.Lower Cretaceous (Berriasian) ; Crimea.The radial facets are wide. The calyx cavity is broadand shallow. The radial and interradial furrows arepronounced. The basais are visible or invisible on thebottom of the calyx cavity. The primaxillary is flattenedwith strongly convex, smooth or granulated externalsurface. The height of the centrodorsal roughlyequals that of the radial ring. One to three deep fossaeare placed under each radial, on the radial = centrodorsalsuture. The centrodorsal basis is wide, roundedand concave. The stem facet is narrow-elliptical. Thecolumnals are low, parallel, rarely cruciate or oblique.Their external surface is smooth or granulated.The fulcral ridge is serrated. Intermediate ridges areoften developed on the columnal articula.D i m e n s i o n s :Ht = 4.5 - 10.2 mm ; Dt = 7.0 - 21.0 mm ; Dc =5.0 - 10.0 mm ; A f = 45 - 57° ; Hr : H t = 0.09-0,25 ; Bf : Lf = 0.48 - 0.64.M a t e r i a l s :32 calyxes, 21 centrodorsals, 8 brachials, 52 columnals(see table 1).Umbocrinus umbonatus nov. sp.fig. 10 B-E ; pl. 1, fig. 7-11D i s t r i b u t i o n :Berriasian ; Crimea.D e r iv a t io n o m i n i s :Th'ê name of the species comes from « umbonatus »(convex in Latin).H o l o t y p e :CK-89-1 (pl. 1, fig. 7) ; Leningrad Mining Institute.T y p e l o c a l it y :Kuibyshevo, Belbek Valley, Crimea.T y p e l e v e l :Upper Berriasian.D e s c r i p t i o n :The calyx is low, smooth and rounded. The basaisare visible on the calyx surface but sometimes not inall interradia. The external radial surface is low, convex,smooth or rarely covered with small granules.T y p e s p e c ie s :Genus Heberticrinus nov. gen.Eugeniacrinus heberti D e L o r i o l in P i c t e t , 1868.D i a g n o s i s :The radiais are with or without low external surface.The basais are invisible on the calyx surface.Rarely they look like tiny triangular plates. The centrodorsalis large, convex with no cirrus socket orfossa. The stem facet is concave, flat or rarely convex.C o m p a r i s o n :Heberticrinus differs from all genera of the familylacking cirrus sockets or fossae on the centrodorsal. Itdiffers from Conoideocrinus, which has no socket orfossa, by the large massive centrodorsal, by theabsence of basais on the external calyx surface and bythe low radiais.

— 647 —ВFig. 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(Gisl é n), H. heberti (D e Lo r io l in P ic t e t ), H.remesi (G is l é n ), H . irregularis nov. sp. One calyxfound in the Hauterivian deposits of the settlementGunib in Daghestan (Caucasus) belongs to the genusHeberticrinus. However, a poor preservation does notpermit us to a specific determination.Upper Jurassic (Tithonian) - Lower Cretaceous(Hauterivian) ; Portugal, France, Czechoslovakia,Rumania, Crimea, Caucasus.Heberticrinus algarbiensis (De Lo r io l , 1888, p.Ill, pi. 22, fig. 10, 11 sub Thiolliericrinus {Thiolliericrinusalgarbiensis De Loriol 1889, p. 560 ; Burdigalocrinusalgarbiensis : Gislén 1924, p. 189 ; Rasmussen1961, p. 213, pi. 35, fig. 11, 12). Hauterivian ;Alfandega (Algarve), Portugal (fig. 11 A).Heberticrinus crassus (G i s l é n , 1924, p. 189 subBurdigalocrinus) (= Thiolliericrinus heberti Rem es>,1905, p.60 ex parte, pi. 7, fig. 2, non fig. 3 = Heberticrinusheberti. Tithonian . Stramberk, Czechoslovakia.Heberticrinus heberti (D e L o r io l in P i c t e t , 1868)figs. 11, E,F? ; 12 A-N ; pl. 1, fig . 12-161868 — Eugeniacrinus heberti : D e Lo r io l (in P ic ­t e t ), p. 281, pi. 42, fig. 7,8.1875 — Eugeniacrinus heberti (De Lo r io l ) : Pillet &Fromentel, p. 98, pi. 10, fig. 31-35.1875 — Apiocrinus flexuosus (G o l d f u s s ) : Pillet &Fromentel, p. 98, pi. 10, fig. 45-50.1883 — Eugeniacrinus heberti D e L o r io l : D eL o rio l, p. 158.1888 — Thiolliericrinus heberti De Lo r io l : DeLoriol, p. 111.1889 — Thiolliericrinus heberti De l o r io l : DeLoriol, p. 545 (ex parte), pi. 228, fig. 1-4, 6-13, pi. 229, fig. 1 (non pi. 228, fig. 5 =Thiolliericrinidae indet.).1897 — T h io lliericrin u s h eb erti D e L o r i o l :Roman, p. 88, 331, 334.1897 — Thiolliericrinus flexuosus Go l d f u s s sp. :Roman, p. 88, 331, 334.1905 — Thiolliericrinus heberti De Lo r io l ; Remes,p .60 (ex parte), pi. 7, fig. 3 (non fig. 2 =Heberticrinus crassus).1912 — Thiolliericrinus heberti De l o r io l : Kirk, p.75, pi. 7, fig. 3 (fig. ex De Loriol 1889, pi.228, fig. 2).1913 — Thiolliericrinus heberti (D e L o r io l ) : Joukowsky& Favre, p. 386, pi. 15, fig. 1-6.1924 — Burdigalocrinus heberti (De Lo r io l ) : Gislén,p. 189.1961 — Burdigalocrinus heberti (De Lo r io l ) : Rasmussen,p. 214.1978c— Thiolliericrinus heberti De Lo r io l : Rasmussen,p. 880, fig. 588-2e,f (fig. ex DeL o rio l, 1889, pi. 228, fig. 7a et 2a).Le c t o t y p e :F .J. Pictet (1868, pi. 42, fig. 8) ; Museum in Chambéry(Savoie, France).T y p e l o c a l it y :Lémenc (Savoie), France.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 :Upper Tithonian.D ia g n o s is :The radial ring is narrowed upwards. Free radialsurface is absent (i.e. radial facets are near the centrodorsal).The basais are not visible on the calyx surface.The centrodorsal is low and wide. Its diameter isconsiderably greater than that of the radial ring. Thecentrodorsal basis is wide, rounded and concave. Thestem facet is narrow-elliptical.D im e n s io n s :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.Re m a r k :When De Loriol (in Pictet 1868, p. 281) was describingEugeniacrinus heberti he indicated its occurrencein the « Brèche de Lémenc ». But he postulated theSequanian age of the species in his following publications.However, this is inaccurate. Remains of thespecies under discussion were found in South-EasternFrance (including Lémenc) together with the spines ofCidaris glandifera, the shells of the brachiopodPygopejanitor, the tests of the rudistid Heterodicerasluci and with the remains of the ammonite Paraulacosphinctestransitorius. The cited faunistic assemblagedemonstrates obviously the Late Tithonian ageof H. heberti. This species was found in the same agedeposits in Czechoslovakia (RemeS 1905, p. 60) andRumania (Patrulius 1956b, p. 198). But in the Crimeait is found in the reef massives directly above the bedswith Dalmasiceras dalmasi (i.e. in the Upper Berriasian).Note shall be taken that, according to Yanin’sopinion (in his letter of 1.3.86), the species Heterodicerasluci is characteristic for the Berriasian. One canassume therefore that a part of the French localities,where remains of this rudistid and H. heberti werefound, is Berriasian.On some calyxes of H. heberti and H. irregularisaffected by parasites, one can see the large holes (fig.12 A, 13 A ; cf. Müller 1977, fig. 7).M a t e r ia l s :87 calyxes, 32 centrodorsals, 2 radial rings, 258 brachials,184 columnals (see table 1).D is t r ib u t io n :Tithonian and Berriasian ; France, Czechoslovakia,Rumania, Crimea.Heberticrinus remesi ( G i s l é n , 1924)fig. 11 C,D1905 — Thiolliericrinus flexuosus É t a l l o n ( G o ld -f u s s ) : Remes, p. 61, pi. 7, fig. 4-6.1924 — Burdigalocrinus remesi : Gislén, p. 190.L e c t o t y p e :Remes (1905, pi. 7, fig. 6) ; Charles University, Prague(teste Zitt, in letteris 1.4.86).T y p e l o c a l i t y :vStramberk, Czechoslovakia.T y p e l e v e l :Upper Tithonian.D i a g n o s i s :The radial ring is narrowed upwards. Free radialsurface is absent (i.e. radial facets are near the centrodorsal).The basais are invisible on the calyx surface.The centrodorsal is high, convex and narrowed downwards.The centrodorsal basis is completely occupiedby a narrow-elliptical stem facet.D i m e n s i o n s :H t - 9.3 - 12.9 mm ; Dt = 13.0 - 16.0 mm ; Dc =9.0 - 10.4 mm ; A f = 33 - 46° ; H r : H t = 0.00 ; Bf :L f = 0.50 - 0.60.M a t e r i a l s :4 calyxes, 9 columnals (see table 1).D i s t r i b u t i o n :Tithonian, Czechoslovakia ; Valanginian, Crimea.

— 651 —Heberticrinus irregularis nov. sp.De r iv a t io n o m i n i s :fig. 13 A-EThe name of the species comes from « irregularis »(irregular in Latin).H o l o t y p e :CK-88-1 (fig. 13 C) ; Leningrad Mining Institute.Ty p e l o c a l it y :Kuibyshevo, Belbek Valley, Crimea.Ty pe l e v e l :Upper Berriasian.surface. The centrodorsal surface is smooth or veryfinely granulated. Some weak fossae are seen sometimeson the radial-centrodorsal suture. The centrodorsalbasis in projected as an acute ridge and completelyoccupied by an elliptical stem facet being orientatedobliquely to the calyx axis. In some cases, the basis iswide, flat and rounded.D i m e n s i o n s :H t = 5.2 - 12.0 mm ; Dt = 6.9 - 18.5 mm ; Dc =5.0 - 12.0 mm ; A f = 37 - 50° ; Hr : Ht = 0.06-0.23 ; Bf : Lf = 0.36 - 0.50.C o m p a r i s o n :H. irregularis differs from other species of thegenus having an irregular centrodorsal and obliquecentrodorsal basis.De s c r i p t i o n :The calyx is high and rounded or roundedpentangular.The radial ring is high, cylindrical ornarrowed downwards. Free radial surface is low orrelatively high. Radial facets are narrow. The basaisare visible on the external calyx surface. The centrodorsalis high and convex with uneven externalM a t e r i a l s :17 calyxes, 18 centrodorsals, 2 brachials, 28 columnals(see table 1).D i s t r i b u t i o n :Berriasian ; Crimea.ВFig. 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 dorsalevue de côté). D - CK-88-22 (columnale et sa facette articulaire). E - CK-88-8 (centrodorsale vue en dessus).

— 652 —5 — RECONSTRUCTION OF THE SKELETONRo o t fo r m a t io n sIn some localities (see table 1), together with calyxesand columnals, root formations of Thiolliericrinidaewere found (Ad - see fig. 2). One cannot determinethe taxonomic position of these roots. It can only beassume that they belong to Heberticrinus heberti orUmbocrinus umbonatus (the most common species).All attachment outgrowths are individual, i.e. onlyone articular facet for a stem is present on each. Theseformations are the monocrystalline calcitic crust(without root-like projections) bearing a nearly notisolated stem socket (fig. 14 A,B). Sometimes, thestem facet is raised as a distinct eminence (fig. 14C,D). Such eminences are formed, very likely, by anaccretion of the lower columnal with the attachmentdisc. It is characteristic that a rounded swelling,resembling a parasite settling, is situated near eachraised socket. Dactylate corallite served, in mostcases, a substratum for the attachment crusts.One can m ark two peculiarities of the attachmentformations. First, a small diameter of the stem facets.It is considerably smaller than the average diameter ofcolumnals found in the same localities. This fact canmean that the Thiolliericrinidae stem is narroweddownwards from the calyx. Second circumstance :almost all stem facets on attachment formations haveevident traces of the selective dissolution. The ligamentfossae on both sides of the fulcral ridge are considerablydeepened. This dissolution is explained bythe fact that the root was left on its place long afterthe death of the animal, being exposed to the corrosionby sea water.Described root formations are analogous to thosenoted for other Thiolliericrinidae species (Étallon1859, p. 445, 447 ; Patrulius 1956a, p. 193 ; Hess1975, p. 69, pi. 25, fig. 1 ; Rasmussen 1978c, p. 879).ВFig. 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 mÉtallon (1859, p. 445, 447) noted that the Thiolliericrinusstem had consisted only of three or four columnals.Joukovsky & Favre (1913, p. 386) found 31calyxes and only 43 columnals of Heberticrinusheberti in one of the local exposures. Similar correlationsare marked also in our materials. So, in the locality« Belbek » (see table 1), there were found 129calyxes and 638 columnals, belonging to several species.A comparison of the figures shows that theThiolliericrinidae stem included roughly five columnals.It coincides with Étallon’s data.Among Heberticrinus heberti columnals, cruciateones are the bulk, oblique ones are rare and parallelones are unique. The columnals of this species are lowand disk-shaped. A t the same time, the columnalswith parallel ridges on both articula are prevalent inBurdigalocrinus maximus. The columnals of this speciesare, as a rule, relatively high. These circumstancesallow us to suppose that the short stem of some discoidalcolumnals was peculiar to the species with wideflattened calyxes. But a relatively long stem, consistingof high columnals, was typical for species with ahigh conical calyx.The order of columnal combination in the stem isunknown. However, one can assume, after the analogyto Bourgueticrinida, that the columnals were joinedso that the fulcral ridge had an alternating « swinging» position. I doubt whether one can speak, in thepresent case, of « spiral turning » as supposed byA rendt (1974, p. 75). In the species Loriolicrinusasper, cruciate and parallel columnals are found inapproximately equal numbers. One can think thatthey were joined in a stem alternately.A powerful fulcral ridge risen above the articulumplane and deep ligament fossae on both sides of themtestify to the considerable flexibility of the stem articulationsin Thiolliericrinidae (Gislén 1924, p. 206). Itresults in flexibility, as a rule, without any markedincrease of the separate columnal height as in Bourgueticrinidawith an analogous articulum construction.The high stem flexibility with the relatively low(strong) columnals agrees with the fact that Thiolliericrinidaeinhabited very mobil waters on the coral reef.The external sculpture of columnals correspondsthat o f the centrodorsal, in most cases. The columnalsof Loriolicrinus asper are often knobby (pl. 1, fig. 5)and those of Umbocrinus umbonatus are coveredsometimes with small granules. At the same time,smooth columnals are often found, belonging evidentlyto the ornamented calyxes. In this case, thedistal or proximal position of the columnal in a stemis probably the matter. Unfortunately, the lack offinds of stem fragments (i.e. some articulated columnals)does not allow one to make the nature of a stemvariation clear.If the cirriless calyxes of Thiolliericrinidae are wellpreserved they have also well preserved stem facet.However, available specimens of Thiolliericrinus(with poverful cirri) have obviously abraded and destructedcentrodorsal basis (fig. 5 A, 6A,B) even whenthe preservation as a whole is good. This circumstancecan mean that the cirriless forms were attached to thestem (and, accordingly, to the substratum) duringtheir whole life. On the contrary, the individualshaving cirri were detached from their stem and led afree life though preserving a biletarel stem facet onthe centrodorsal (Kirk 1912, p. 75). The poor preservationof the centrodorsal basis in these forms isexplained, possibly, by their activity. The separationof the stem during the life of an animal is confirmedby the absence (among over 500 columnals) of a singlecolumnal belonging to Thiolliericrinus belbekensis, T.elongatus or T. torosus.A r m s .Thiolliericrinidae brachials are known till now onlyafter Rem e!s (1905, pi. 7, fig. 7,8). In our collectionthere are 580 brachials plates of several species of thefamily (see table 1).Only muscular articula are developed on all the brachialswithout exception, including the lower face ofaxillaries. This leads to the conclusion that there wereno immovable articulations in the arms of Thiolliericrinidae.The axillaries of the first, second and third orderscan be determined with certain among available axillarplates. Consequently, arm branching was not lessthan threefold.Not a plate has been found which should be placedbetween the primaxillary and the radial. So, one canconclude that Thiolliericrinidae have only one primibrachial(axillary) with muscular articulation directlyjoined to the radial.The primaxillaries (IAx - see figs. 2 and 15) are thelargest plates in the crown, being greater than theradiais. The form of promaxillaries is various in differentgenera. The calyxes with high free radial surfaceand with a small angle of the inclination of the radialfacets have high axillaries with vertical external

— 654 —SIS rllAxsurface (Loriolicrinus, Burdigalocrinus). The calyxeswith low radiais whose facets are steeply inclined outsidehave wide and flattened primaxillaries hung overthe centrodrosal and absolutely covering radial plates(Umbocrinus, Heberticrinus). Two upper primaxillaryfacets are of an identical size and inclined outsidenearly parallel to the facets of the subjacent radiais.The lateral faces of all available axillar plates are flattened.Consequently, thesé plates formed a closedring. The sculpture of the external surface of a primaxillarycorresponds to that of the calyx. The pinnularsockets are absent on primaxillaries.IIBr3l!Br2IIBMThe brachials of most Articulata crinoids bear onepinnular socket placed on the upper inner margin ofthese plates. The fulcral ridges are not in parallels onlower and upper brachial faces. But these ridges canhave only two positions swinging from one articulumto another. The available secondbrachials were dividedinto right and left, even and odd, accordingly(fig. 15). If one looks at the primaxillary from itsexternal surface, the left odd (first) secondbrachialwill be on its upper face to the left and right odd (first)secondbrachial to the right. The odd secondbrachialsare cuneate with a thin margin at the inner side of aradius and with a thick one at the outer side where thepinnular socket is placed. Their external surface isvery convex. The thick lateral side of the secondbrachialbears a wide flat face, i.e. these plates were closeto the corresponding brachials of the adjacent radius.The even brachials are flattened and somewhat higherat the inner margin (where the pinnular socket is placed).They have convex external surface and flat lateralsides joined to corresponding secondbrachials ofthe adjacent radii. The number of secondbrachials inthe arm branch was apparently odd and hardly morethan seven or nine (according to the relatively greatnumber of the available secondaxillaries). The externalsecondbrachial surface has the sculpture analogousto that of the calyx but feebly expressed.The secondaxillaries (IIAx - see figs. 2 and 15),according to their construction, are analogous to theFig. 15 — A scheme of the construction of Thiolliericrinidae brachials(view from above, x 2). IAx - ring of five primaxillaries,IIBrl - first secondbrachial, IIBr2 - secondsecondbrachial, IIBr3 - third secondbrachial, IIAx-secondaxillar, IIIBrl - first tertibrachial, IIIBr2 -secondtertibrachial, SI - left arm series, Sr - right arm series.The number of secondbrachials is shown conditionally.Reconstitution de l’organisation d’un tronc brachial de Thiolliericrinidae(vue en dessus, x 2). IAx - cercle des cinq primaxillaires,IIBrl - première secundibrachiale, IIBr2 - deuxième secundibrachiale,IIBr 3 - troisième secundibrachiale, IIAx - secundaxillaire, IIIBrl -première tertibrachiale, IIIBr2 - deuxième tertibrachiale, SI -sériegauche d’un bras, Sr - série droite d’un bras. Le nombre des secundibrachialesreste 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 removed),Heberticrinus (in the centre) and Burdigalocrinus (from the right ; traces of parasites settlings are shown on the centralradius).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 innerupper facet is considerably greater than the outer one.The construction of tertibrachials is like that of thesecondbrachials. Only the wedge-shaped form ofseparate plates is more pronounced. Also the odd brachialshave their external surface like an obtuse horizontalridge. One failed to determine the number ofbrachials in the third arm series. The tertiaxillaries aresymmetrical having flattened lateral faces as before.The axillaries of the second and third orders have nopinnules. The tertibrachials have smooth external surfacewithout any dependence on the calyx ornamentation.The crown of Thiolliericrinidae was uniform on thewhole, differing only in details in different speciesand even genera. It formed, in the closed position, acontinuous tube at least up to the level of the tertiaxillaries(fig. 16). By that, relatively very small pinnuleswere enclosed into the crown and not visible from theoutside. One can say that the crown skeleton was atest being made of closely adjusted and firmly unitedbrachial plates. The Thiolliericrinidae crown resembled,in this respect, the crown of Triassic Encrinusliliiformis L a m a r c k but was considerably greater.6 — ONTOGENETIC VARIATIONS OF THE CALYXESAccording to available materials, the calyx ontogenycould be reconstructed for three Thiolliericrinidaespecies : Loriolicrinus asper, Umbocrinus umbonatusand Heberticrinus heberti. One fact attracts ourattention : in all three forms, in spite of their belongingto different genera, there are almost the samecalyxes in young specimens, namely : high-conical orcylindrical with high free radial surface. However,subsequently the development of the calyxes proceededby different ways in each of the three species.The radiais grew quicker than the centrodorsal in L.asper (fig. 17A). Therefore the centrodorsal of thisspecies remained low and its upper surface is convex.The young forms of U. umbonatus (fig. 17B) havestrongly pronounced though low basais and fullydeveloped fossae under radiais. Then the centrodorsaland the radiais grew at an about equal speed. Thereforethe upper centrodorsal surface is flat. Simultaneouslywith the preserving of the basais on the externalcalyx surface, the relative height of the free radialsurface considerably decreased. The sutural pores didnot disappear but even increased in number and size,so that they changed into irregular openingsFig. 17 — Schematic sections of some Thiolliericrinidae calyxesshowing the stages of the ontogenetic development (R -radial, В - basal, Cd, centrodorsal) : A - Loriolicrinus,В - Umbocrinus, С - Heberticrinus.Sections schématiques des coupes dorsales de quelquesThiolliericrinidae, montrant les stages du développementontogénétique.cdc

— 657 —occupying nearly the whole centrodorsal surface inthe oldest specimens. The stem facet, being flat in theyoung specimens, turned into concave in the adultones.The centrodorsal grew quicker than the radiais inH. heberti (fig. 17C). Therefore, the upper centrodorsalsurface is very concave. The basais, being visible inthe young specimens, disappeared in the adult ones.The height of the radiais decreased. The diameter ofthe centrodorsal was roughly equal to that of theradial ring in the young specimens. However, the centrodorsalwas considerably wider than the radiais inthe adult specimens. The stem facet was almostalways flat.7 — ORIGIN OF THE THIOLLIERICRINIDAEMany thiolliericrinids have the quite developed cirriwith the mobil bifascial articulation of cirrals on theircentrodorsals. Only Comatulida and Pentacrinida *may have such cirrus construction among postpaleozoiccrinoids. So specialized cirral organs couldnot originated twice in the phylogeny of crinoids.Therefore, the origin of Thiolliericrinidae must beconsider in the close connection with the origin ofcomatulids and pentacrinids in one phylogeneticsequence with these groups.Three hypothesis are possible in the comprehensionof the development of three given groups, namely :(1) stalked pentacrinids, (2) stalked thiolliericrinids or(3) stemless comatulids are initial. The embryologicalinvestigations showed that young stages of comatulidswere stalked (Thomson 1865 ; A .H . Clark 1921). Thefine and peculiarly constructed perforation on thelower centrodorsal face of some (relatively primitive?) comatulids (A.M. Clark 1973, fig. 1—3) are alsothe illustration of the stalkness of comatulid ancestors.Available facts testify to the origin of Comatulidafrom stalked forms and to the unacceptability ofthird (see above) hypothesis respectively.Earliest pentacrinids known beginning with earlyTriassic (Holocrinidae) had a number of specific features.One should note the following primitive signs :the distal attachement disc (replaced later into unattachedbulge), the infrabasals which can be seen on thecalyx surface, the narrow spot of the crown on theradial = brachial suture. The newly-gained feature isthe only one : the nodals in proximal stem part withshort cirri (mostly not more than three in a whorl).The recent Proisocrinidae are the analogue for theseancient stages. Known finds of young pentacrinids(Carpentier 1884, pi. 30a, 35, 36) and encrinids (Hagdorn1978, Abb. 21 ; 1982, Abb. 18, 19) also showthat pentacrinids have been developed from attached* Pentacrinida T o r t o n e s e (1938, p. 171, 177, 212) = IsocrinidaSie v e r t s -D o r e c k , 1952.and cirriless forms with quite large infrabasals, andthat they had many features common with encrinids.Then, the development of pentacrinids followed apath of perfection of the cirral organs (with a mountingof the five-rayed nodal symmetry and an appearanceof the cryptosymplectial sutures : Hagdorn1983, p. 357 ff.), transm utation of the calyx fromdicyclic to cryptodicyclic and disappearance of themorphological isolation of the calyx from the arms.Cited data show that origin and development of thepentacrinids followed without any connection withthe considerably more late Thiolliericrinidae. Consequently,only one of above mentioned hypothesisremains acceptable : the pentacrinids are the ancestorof the cirriferous Articulata.By subsequent consideration of this hypothesis, twoversions are possible : (1) Pentacrinida —» Thiolliericrinidae—> Comatulida (De Loriol 1880, p. 11 ; Kirk1912, p. 75 ; Jaekel 1918, p. 71 and others) or (2) Pentacrinida—> Comatulida —> Thiolliericrinidae (Rasmussen1978b, p. 314 ; 1978c, p. 867, 879).One can make out two stages in the skeleton developmentof the recent comatulids. Deltoid plates,large basais, minute infrabasals and columnals arelaid at early cystidean stage. Absence of cirri and theconstruction of the calyx basis permit to see, in thisstage, an analogy with the initial forms for articulatecrinoids (possibly, with Encrinidae). Further, thenodal plates on which cirri can developed subsequentlyare laid under the calyx in pentacrinoid stage.We can see here the transition from Encrinidae toPentacrinida. The cirri develop on all the proximalnodals of pentacrinids. The cirri appear but only onone or two most proximal nodals of comatulids,however some discoidal cirriless columnals which arehomological to pentacrinid nodals are formed belowthem. Then, the detachment of the comatulid larvafrom the stem takes place. By that, the free animalhad not carried yet that indivisible plate which shouldbe called as a centrodorsal. This very important circumstanceshows that the comatulid centrodorsal wasformed by some proximal nodal plates (Thomson1865, p. 536).

— 658 —At the same time, all the Thiolliericrinidae withoutexception possessed of a centrodorsal. And what ismore, this plate assumes the relatively greater size inyoung specimens (fig. 8 E, 9 C). This indicates thatThiolliericrinidae originated from the forms having awell developed large centrodorsal, and Comatulida -from the forms with the centrodorsal consisting ofseveral plates.Paracomatula Helvetica (H e s s , 1951, fig. 1—7) hasthe centrodorsal consisting of several nodal columnalsand crown already typical for comatulids, and servesa good example for the forms transitional from Pentacrinidato Comatulida (see fig. 18). Carpenterocrinusmollis (C a r p e n t i e r , 1884, p. 338, pi. 33, fig. 7-10) can be an example of such transitional formsamong recent crinoids.Thus, the hypothesis on the origin of the Thiolliericrinidaefrom comatulids and that on the origin of thelatter, in its turn, from pentacrinids seems to be themost substantiated ones.The explanation of the origin of Thiolliericrinidaefrom the family Solanocrinitidae seems to be mostconvincing. Adutl individuals of this latter familyhave large centrodorsals where, under each radial,two vertical rows of the cirrus sockets are placed.Each row can contain up to five sockets and more.The basais are well visible on the calyx surface, theradiais have high external surface and the centrodorsalbasis is flattened and smooth. In the enumerationof these features one can see an evident coincidencewith the calyx construction of Solonaerium, the firstand most primitive genus of the family Thiolliericrinidae(fig. 18).8 — DEVELOPMENT OF THE THIOLLIERICRINIDAEDuring settling of the reefs, one of the groups of theorder Comatulida (possibly Solanocrinitidae) obtainedthere the most favourable conditions for its life(pure water, an abundance of the light, heat, oxygen,food). There was no necessity to change the place oflife, and therefore the moment of the larval stem tearingoff began to be delayed. The individuals of Solonaeriumand Thiolliericrinus were free when theybecame adult. After detaching from a stem theseforms lived freely using their powerful centrodorsalcirri to attaching to the bottom (see fig. 16). Therefore,it seems to me that the determination of Thiolliericrinusas « a permanent larval form » (Carpenter1881b, p. 377) is quite good.The sessile mode of life appeared so successful, atthe time, that it began to be the cause of subsequentmorphological adaptations of the skeleton. The cirribegan to be lost as useless now (Gislén 1924, p. 206 ;Rasmussen 1978a, p. 273), the stem became morepowerful and more flexible. Here we witness thechange from cirriferous forms to Burdigalocrinus andUmbocrinus, having rudimentary cirrus fossae on thecentrodorsal, and then to Conoideocrinus and Heberticrinuswith no traces of cirri. The basais and partlythe radiais were reducing (Bather 1900, p. 195 ; Kirk1912, p. 78). The crown became more compact andmore strongly constructed (see fig. 16). The animalswere attached during the whole life. By that, theattachment organs (the stem and the distal disc) weresimilar to the larval structures of the « true » comatulids.Just so « Thiolliericrinidae appear to be aproterogenetic evolution from the Solanocrinitacea »(Rasmussen 1978b, p. 314). Thiolliericrinidae are evidentlynot an unique example of the conservation oflarval features by adult animals adapted to the life inthe reef conditions.The morphological variations of Thiolliericrinidaecalyxes are presented on fig. 18, according to theirsupposed phylogenetic correlation.Solonaerium seems to be the initial form with highradiais, large basais and a regular disposition of thecirrus sockets. Subsequently the disposition of thesockets lost the regularity but the basais were still preserved(Thiolliericrinus). The radiais can be high orlow. Then the number of the cirrus sockets decreased,some sockets turned into structureless fossae. Laterthe sockets disappeared completely, but the fossaewhich had been scattered on the centrodorsal remained(Burdigalocrinus). The full disappearance of thecirri on the centrodorsal, with the preservation ofhigh radiais and large basais, is typical for Conoideocrinus.The centrodorsal fossae can remain on theradial-centrodorsal suture. The gradual disapearanceof these fossae can be observed simultaneously withthe atrophy of the basais and the preservation of thehigh radiais (Loriolicrinus). Otherways, fossae andbasais could remain (and even decrease in size) withthe considerable reduction of the radial ring (Umbocrinus).The final disappearance of the fossae and thebasais, and the decrease of the relative size of radiais,are marked in the genus Heberticrinus in which theprincipal morphological types can be determinedaccording to the form and size of the centrodorsal.

— 659 —JURASSIC L. CRETACEOUSFig. 18 — A scheme of the origin and the development of main morphological types of the Thiolliericrinidae calyxes (the dotted arrowssymbolize the continued independent development).Schéma de l’origine et du développement des types morphologiques principaux des coupes dorsales des Thiolliericrinidae.P - Pentacrinus, Pc - Paracomatula, St - Solanocrinites, S - Solonaerium, T - Thiolliericrinus, U - Umbocrinus, H - Heberticrinus,B - Burdigalocrinus, С - Conoideocrinus, L - Loriolicrinus.

— 660 —AcknowledgementsThis article was written in the Historical Geology Chair ofthe Leningrad Mining Institute. A part of the collections servingthe materials for this work was handed to the author byDr. T.N. Bogdanova and Dr. S.V. Lobachiova (All-UnionGeological Institution, Leningrad). The author consultedthem on the Lower Cretaceous stratigraphy of the Crimea,Dr. B.T. Yanin (Moscow University) on the stratigraphicalrange of late Jurassic and early Cretaceous rudistids. Thebrachiopods from some localities were determined by Dr.S.Y. Lobachiova. The information on the Czecho-SlovakThiolliericrinidae was received from Dr. J. Zitt (Mnichovice,Czechoslovakia). The manuscript was reviewed by Dr.A.S. Tarakanov (All-Union Geological Institution, Leningrad)and by Dr. S.N. Alekseev (All-Union GeologicalPetroleum Institution, Leningrad). The translation of thearticle into English was made by Mgr. Igor F. Naftul’eff andR.D. Kirpichnikov. The author thanks Prof. M. Roux forhis help in supervising the manuscript.REFERENCESARENDT Y.A. (1974) - Sea-lilies Cyrtocrinids. TrudyPaleontologicheskogo Instituta, Moscow, 144, 250 p. (inRussian).ARENDT Y.A. & YANIN B.T. (1964) - On Late Jurassicand Early Cretaceous crinoids of the Crimea. J. Paleontol.,Moscow, 3 : 140-142 (in Russian).BATHER F.A. (1900) - The Echinoderma. In : LankesterE.R., A Treatise on Zoology, pt. 3, London, 344 p.CARPENTER P.H. (1881a) - On the genus SolanocrinusGOLDFUSS, and its relations to recent Comatulae. J.Linn. Soc. London, Zool., 15 : 187-217.CARPENTER P.H. (1881b) - Interesting new Crinoids.Nature, London, 23 : p. 377.CARPENTER P.H. (1884) - Report on the Crinoidea collectedduring the Voyage of H.M.S. Challenger. StalkedCrinoids. Report on scientific results o f the Voyage o fH.M.S. Challenger, Zoology, London, 11, 442 p.CLARK A.H. (1913) - Post-Paleozoic Crinoidea. In : Textbooko f Paleontology edited by C.R. Eastman, adaptedfrom the german of K.A. Zittel, vol. 1, London - NewYork : 225-242.CLARK A.H. (1921) - A monograph of the existing crinoidsvol. 1. The Comatulids, pt. 2. Bull. U.S. Nat. Museum,Washington, 82, 795 p.CLARK A.M. (1973) - Fossil and recent Comatulid Crinoidswith coelomic extensions penetrating the centrodorsal.Bull. Brit. Mus. (Natur. Hist), Zool., London, 24, 9 :441-446.DACQUÉ E. (1933) - Wirbellose des Jura. In : Gtirich G.,Leitfossilien, Berlin, 7, 1.1., 272 p.ÉTALLON A. (1857) - Esquisse d’une description géologiduHaut-Jura, et en particulier des environs de St-Claude. Ann. Sci. phys. natur. agricult. industr., Lyon,(3), 1 : 247-354.ÉTALLON A. (1859) - Études paléontologiques sur le Haut-Jura. Rayonnés du Corallien. Mém. Soc. Emul. Doubs,Besançon, (3), 3 : 401-553.ÉTALLON A. (1862) - Études paléontologiques sur le Haut-Jura. Monographie du Corallien. Mém. Soc. Emul.Doubs, Besançon, (3), 6 : 53-260.GISLÉN T. (1924) - Echinoderm Studies. Zool. BidragUppsala, 9 : 330 p.GOLDFUSS A. (1826-1833) - Petrefacta Germaniae, Tl.lDüsseldorf, 252 p.HAGDORN H. (1978) - Muschel-Krinoiden Bioherme imOberen Muschelkalk (mo 1 Anis) von Crailsheim undSchwabisch Hall (Südwestdeutschland). N. Jb. Geol.Palaont. Abh., Stuttgart, 156, 1 : 31-86.HAGDORN H . (1982) - Chelocrinus schlotheimi (QUENS-TEDT) 1835 aus dem Oberen Muschelkalk (mo 1, Anisium)von Nordwestdeutschland. Verôff. Naturk. Mus.Bielefeld, 4 : 5-33.HAGDORN H . (1983) - Holocrinus doreckae n.sp. aus demOberen Muschelkalk und die Entwicklung von Soilbruchstellen im Stiel der Isocrinida. N. Jb. Geol.Palaont. Mh., Stuttgart, 6 : 345-368.HESS H. (1951) - Ein neuer Crinoide aus mittleren Doggerder Nordschweiz (Paracomatula Helvetica n.gen. n.sp.).Eel. geol. Helv., Basel, 43, 2 : 208-216.

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PLATE

PLATE 1Fig. 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 crêtes fulcrales croisées.16. CK-51-313 : articular face of a distal cruciate columnal (x 3.4). Facette articulaire d’un article distal à crêtes fuicraies croisées.Crimea, Belbek Valley : Berriasian.

Geobiosn° 20, fasc. 5Pl. 1V.G. Klikushin