Echinoids from the Triassic (St. Cassian) of Italy, Their Lantern Supports, and a Revised Phylogeny of Triassic Echinoids Porter M. Kier SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY • NUMBER 56 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon pubiication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, givjng an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." 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Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. S. Dillon Ripley Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY • NUMBER 56 Echinoids from the Triassic (St. Cassian) of Italy, Their Lantern Supports, and a Revised Phylogeny of Triassic Echinoids Porter M. Kier SMITHSONIAN INSTITUTION PRESS City of Washington 1984 ABSTRACT Kier, Porter M. Echinoids from the Triassic (St. Cassian) of Italy, Their Lantern Supports, and a Revised Phylogeny of Triassic Echinoids. Smithsonian Contributions to Paleobiology, number 56, 41 pages, 4 figures, 14 plates, 1984.— Three new species of Triassic echinoids are described from the St. Cassian (Karnian) beds of Cortina d'Ampezzo, Italy: Levicidaris furlani, L. pfaifferi, and Zardinechinus giidinii. Hundreds of echinoid fragments from the same beds show that 16 species lack apophyses (interambulacral lantern supports) and 7 possess them. Previously, paleontologists assumed that most Triassic echinoids had apophyses. Their absence from so many species and the presence of slightly developed auricles (ambulacral lantern supports) suggest that two echinoid lineages crossed from the Paleozoic to the Triassic: one, possessing apophyses, is ancestral to all modern cidaroids; a second, lacking apophyses, gave rise to all noncidaroid echinoids. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The trilobite Phacops rana Green. Library of Congress Cataloging in Publication Data Kier, Porter M. Echinoids from the Triassic (St. Cassian) of Italy, their lantern supports, and a revised phylogeny of Triassic echinoids. (Smithsonian contributions to paleobiology ; no. 56) Bibliography: p. Supt. of Docs, no.: SI 1.30:56 I. Sea-urchins, Fossil. 2. Paleontology—Triassic. 3. Paleontology—Italy. I. Title. II. Se- ries. QE701.S56 [QE783.E2] no. 56 560s [563'.95] 83-600346 Contents Page Introduction 1 Acknowledgments 1 Localities 1 Lantern Supports 2 Discussion 3 Nature of Lantern Supports in St. Cassian Species 3 Species without Apophyses 3 Species with Apophyses 6 Nature of Some Ambulacra in St. Cassian Species 8 A Biserial Ambulacrum 8 Species Discussion 8 Mikrocidarispentagona (Miinster) 8 Megaporocidaris mariana Kier 9 Tiarechinus princeps Neumayr 10 Levicidaris zardinia Kier 10 Levicidaris furlani, new species 10 Levicidaris pfaifferi, new species 11 Zardinechinusgiulinii, new species II Zardiyiechinus lancedelli (Zardini) 12 Literature Cited 13 Plates 1-14 14 111 Echinoids from the Triassic (St. Cassian) of Italy, Their Lantern Supports, and a Revised Phylogeny of Triassic Echinoids Porter M. Kier Introduction The best preserved Triassic echinoids come from the St. Cassian beds (Karnian) near Cortina D'Ampezzo, Italy. A large collection of these echinoids was described recently (Kier, 1977), but additional material has been found that in- cludes 3 new species and also fragments that show the nature of lantern supports in many of the species. Previously, it was believed that most Triassic echinoids had lantern supports formed from interambulacral plates (apophyses); how- ever, the majority of St. Cassian echinoid speci- mens studied have no apophyses and either had no lantern supports or have supports formed from ambulacral plates (auricles). This new in- formation requires revision of our understanding of the phylogeny and the higher taxonomic cat- egories of Triassic echinoids. ACKNOWLEDGMENTS.—I thank Rinaldo Zar- dini of Cortina d'Ampezzo who permitted me to study all his St. Cassian fragments, including Porter M. Kier, Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. newly collected material. All the specimens fig- ured in this paper in the section on lantern supports were collected by him. Giovanni Furlan of Treviso, Italy, arranged the loan of a large number of St. Cassian echi- noids collected by himself, Fabrizio Bizzarini, Eugenio Pfaiffer of Venice, and Saverio Giulini of Milan. I thank them for having the patience and skill necessary to find these rare fossils and for their willingness to lend them to me for study. These specimens are described in the "Species Discussion" section, including the type specimens of the three new species. David Pawson and Gordon Hendler of the Smithsonian Institution critically read the man- uscript, and Andrew Smith of the British Mu- seum (Natural History) shared his cladistic ap- proach with me and showed me how to use it to better understand the phylogeny of Triassic echi- noids. Most of the photography was done by Arnold L. Powell and the artwork by Larry B. Isham. The manuscript was edited by Donald C. Fisher, Smithsonian Institution Press, and I thank him for his suggestions. LOCALITIES.—All the echinoids were collected SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY from the following localities in the Dolomites region of Italy. AIpe di Species: SW of base of Monte Specie, approximately 12 km NE of Cortina d'Ampezzo. Boa Staohn: 1 km NE of Alvera, approximately 2.5 km NE of Cortina d'Ampezzo. Campo: approximately 2.01 km S of Cortina d'Ampezzo. Cianzope: 2.5 km E of Lago Bai di Dones, approximately 5.5 km W of Cortina d'Ampezzo. Forcella Giau: 2.5 km NE of Selva di Cadore, approximately 9 km SW of Cortina d'Ampezzo. Milires: 0.5 km W of Campo di Sotto, a town 2.25 km S of Cortina d'Ampezzo. Misurina: 2 km E of base of Mt. Cristallino, 10.5 km NE of Cortina d'Ampezzo. Rumerlo: 0.25-0.5 km S of Rumerlo, town 2.5 km W of Cortina d'Ampezzo. Stolla: 2.5 km NW of Monte Specie. Tamarin: 300 m NE of Lake Tamarin, approximately 3 km NE of Cortina d'Ampezzo. St. Cassiano: Badia Valley, near Rio Stuores. The specimens are deposited in the Museo di Cortina d'Ampezzo, Cortina d'Ampezzo, Italy (MCA); the Museo di Storia Naturale, Venice, Italy (MV); the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM); and in the Pfaiffer (private) collection in Venice, Italy. Lantern Supports The lantern of an echinoid is attached by two sets of muscles to the test near the inside edge of the peristomial opening. One set of muscles, the retractors, serve to pull the lantern back and the second set, the protractors, extend the lantern. The retractors in all modern cidaroids are at- tached to projections, called apophyses, at the edge of the interambulacra. In all noncidaroid, regular echinoids no apophyses are present, and the retractors are attached to projections, called auricles, in the ambulacra. The protractors are attached always to the interambulacra. In the cidaroid, these muscles attach to the medial re- gion in the apophyses; in the noncidaroids they are attached to a slight ridge along the peristo- mial edge ofthe interambulacra. Lantern supports are absent in lower Paleozoic echinoids. They first appear in the Permian where apophyses (interambulacral supports) are present (Kier, 1974:56) in Miocidaris keyserlingi (Geinitz) and Miocidaris connorsi Kier. Apophyses are present in the Early Triassic Lenticidaris utah- ensis Kier. It has been assumed that the vast majority of Triassic echinoids were cidaroids hav- ing apophyses; however, study of these St. Cas- sian fragments shows that most ofthe species had no apophyses and that some of those lacking apophyses had slightly developed lantern sup- ports in their ambulacra (auricles). They include the following 16 species: Zardinechinus lancedelli (Zardini) Z. giulinii, new species Levicidaris zardinia Kier L. furlani, new species L. pfaifferi, new species Megaporocidaris mariana Kier Tiarechinus princeps Neumayr Triadocidaris venusta (Miinster) Paurocidaris rinbianchi (Zardini) "Miocidaris' adrianae Zardini Mikrocidaris pentagona (Munster) 5 unnamed species The auricles, when present, are commonly formed by thickening of the first three plates at the adoral edge of each ambulacrum (Figure 1; Plate 1: figures 1-3). Although the edges ofthe adjacent interambulacra are thickened, they are not part of the auricle, which lies on top of the interambulacra. In many of these species a small protuberance is present in the middle of the inside of the interambulacra. The protractor muscles were presumably attached here. In mod- ern echinoids the protractor muscles are always attached medially in each interambulacrum rel- ative to the retractor muscles. Apophyses are present in the following St. Cassian species: Polycidaris regularis (Munster) "Miocidaris" arnpezzana Zardini Leurocidaris monlanaro (Zardini) 4 unnamed species NUMBER 56 . ■ - ' ••.•■•.;•;>;'»••»'•'•'.'.■■.•-";i!';.i FIGURE 1.—Interior and adradial views of adoral margin of an ambulacrum, MCA 5689, showing the auricles, X 20. Campo. Collector: Zardini. DISCUSSION The sixteen species lacking apophyses cannot be considered to be cidaroids. It is unreasonable to suggest that cidaroids gained apophyses in the Permian and lost them in the Triassic and then later regained them. If they had, it would be expected that there would be intermediate forms—species with vestigial apophyses and small auricles—but none have been found. The late Triassic echinoids with small auricles have no traces of apophyses. It is more reasonable to assume that two echi- noid lineages (Figure 2) crossed from the Per- mian to the Triassic. One, the cidaroids, had apophyses and the other, an unnamed suprataxa, had no lantern supports. This latter lineage de- veloped auricles in the Late Triassic and is an- cestoral to all living, noncidaroid echinoids. Reg- ular echinoids lacking lantern supports became extinct at the end of the Triassic. Nature of Lantern Supports in St. Cassian Species SPECIES WITHOUT APOPHYSES Zardiyiechinus lancedelli (Zardini) PLATE 1: FIGURES 4, 5 The lantern supports in this species consist of auricles formed by thickening of the three adoral-most ambulacral plates in each column. The highest part of each auricle rises from the third plate, forming a tab to which, presumably, were attached retractor muscles of the lantern. Although the interambulacra thicken at the ad- radial margin, the retractors presumably were not attached to the interambulacrum. The am- bulacral auricles (as viewed from the interior) rest on top of the adradial margin of the inter- ambulacra. No protuberance is present at the adoral margin in the middle of each interambu- SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY RECENT Cidaridae Noncidaroid regulars irregulars L*^^ 7 species TRIASSIC MIDDLE 1 EARLY Lenticidaris utoitensis Kier 1 1 16 species j /auricles Miocidaris ireyserlingi (Geintz) / ii4. connorsi Kier / PERMIAN \ / FIGURE 2.—Phylogeny of the echinoids based on character of lantern supports. lacrum. More than one hundred interambulacra of this species were examined and none of them had this protuberance. (See "Species Discus- sion.") Zardinechinus giulinii, new species The interior at the edge of the peristome is visible in the holotype and one of the paratypes. No apophyses are present. The ambulacra are thickened at the adradial suture, forming an auricle similar to that in Zardinechinus lancedelli. (See "Species Discussion.") Levicidaris zardinia Kier PLATE 1: FIGURES 6, 7 The lantern supports in this species are indis- tinguishable from those in Z. lancedelli; they are clearly visible in USNM 370501. There are no apophyses. The ambulacra and interambulacra NUMBER 56 are thickened at the adradial margin. The three adoral-most plates in each ambulacral column are thickened, forming an auricle. The highest part ofthe auricle rises from the third plate. (See "Species Discussion.") sammlung fiir Palaontologie und Historische Geologic, Munich, West Germany. No apophyses are present. Although the ambulacra are thick- ened at the adradial suture, no auricles are pre- served. Levicidaris furlani, new species The lantern supports are similar to those in L. zardinia. No apophyses are present. Slight auri- cles are formed by thickening of the adoral-most plate in each ambulacral column. (See "Species Discussion.") Levicidaris pfaifferi, new species The lantern supports are similar to those in L. zardinia. There are no apophyses. Slight auricles are formed by the thickening of the adoral-most ambulacral plates. (See "Species Discussion.") Megaporocidaris mariana Kier Several fragments, MCA 3146-S-Z and 3147- S-Z (Kier, 1977, pi. 15: figs. 6, 8), show the interior adoral edge of the test. No apophyses are present. Although thickening occurs at the edge of the adoral-most ambulacral plates, it is not clear whether or not auricles are present. (See "Species Discussion.") Paurocidaris rinbianchi (Zardini) No apophyses are visible on the holotype, MCA I25-M-Z. Tiarechinus princeps Neumayr On specimen MCA I93-S-Z no apophyses are present. The plates at the peristomial edge of the ambulacra are thickened, forming slight auricles. (See "Species Discussion.") Triadocidaris venusta (Munster) The peristomial region is preserved on the holotype (AS VII 436) in the Bayerische Staats- Species I PLATE 2: FIGURES 1-6 MATERIAL.—MCA 5692, 5693, 5694. Although the species cannot be generally iden- tified, it is distinguished from the other St. Cas- sian species by its wide interambulacra with per- forate tubercles adorally. Its tubercles have par- apets which are not crenulate except for two crenulations on the adoral side of each tubercle. This species differs from species 2 by its larger size and much narrower medial tract. All 15 fragments show the inside of the adoral edge of interambulacra. No apophyses are pres- ent. The interambulacra are thickened at the adradial suture. A protuberance is present in the middle of the interior of the adoral edge on most of the specimens (Plate 2: figure 4). This protu- berance probably was the point of attachment of the protractor muscles. No specimens of this species have been found with ambulacra, so it is not known whether or not auricles were present. Species 2 PLATE 3: FIGURES 1-6; PLATE 4: FIGURES 1,2 Triadocidaris species B.—Kier, 1977:25, pi. 18: figs. 3-5. MATERIAL.—MCA 168, 192, 5696, 5698. Ninety-three fragments can be referred to this species, which is easily distinguished from the other St. Cassian species by its small, wide test, perforate tubercles, and wide medial tract. This species has no apophyses. The interam- bulacra are thickened at the adradial suture, and on most specimens a protuberance is present in the middle of the inside adoral edge of each interambulacrum. No specimens are available with ambulacra; therefore, it is not known whether this species had auricles. SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY FIGURE 3.—Species 3 showing sutures, MCA 5699, X 20. Campo. Collector: Zardini. Species 3 FIGURE 3; PLATE 4: FIGURES 3, 4 MATERIAL.—MCA 5699. This species is based on only one fragment including the adoral portion of a half interam- bulacrum and ambulacrum. It is easily distin- guished from all the other St. Cassian species by the many columns of tubercles in the interam- bulacra. At the adradial suture both the inter- ambulacrum and ambulacrum are thickened, forming a ridge that was probably an auricle. This is the only specimen I have seen on a Trias- sic echinoid in which the auricle is partially inter- ambulacral in origin. A large protuberance is present at the midline of the interambulacrum. "Miocidaris" adrianae Zardini PLATE 4: FIGURES 5, 6; PLATE 5: FIGURES 1-6 Miocidaris adrianae Zardini, 1973:9, pi. 6: fig. 25. Species 1.—Kier, 1977:26, pi. 18: figs. 6-8, pi. 19: figs. 1, 2. MATERIAL.—Holotype: MCA 121. Paratypes: MCA 5700, 5701, 5702. This species is based on three specimens be- sides Zardini's holotype. Although I cannot generically identify it, it is easily distinguished from the other St. Cassian species by its strongly crenulate, perforate tubercles and deep pits on the interior of the interambulacra behind the primary tubercles. Nothing is known of the am- bulacra. No apophyses are present. The inter- ambulacra are only very slightly thicker at the adradial margin. A slight protuberance occurs adorally at the midline (Plate 5: figure 2). Species 4 Species 2.—Kier 1977:26, pi. 19: fig. 3. This species is based on only one fragment (MCA 3I54-R-Z) of an interambulacrum and cannot be identified generically. It differs from "Miocidaris" adrianae in having lower plates and more strongly elevated bosses lacking basal ter- races. It lacks an apophysis. Species 5 PLATE 6: FIGURES 1, 2 Mesodiddewaf.—Kier, 1977, pi. 19: figs. 4, 5 [fragments]. MATERIAL.—MCA 5704. Three interambulacral fragments are known of this species, which is easily distinguished from the other St. Cassian species by its more numer- ous interambulacral tubercles. No apophyses are present. The adradial margins of the interam- bulacra are thickened, and a protuberance is well developed medially near the adoral edge. Mikrocidaris pentagona (Munster) No apophyses are visible on any of the speci- mens I have seen of this species. (See "Species Discussion.") SPECIES WITH APOPHYSES Polycidaris regularis (Munster) The holotype, AS VII 435, is the only known specimen of this species and is in the Bayerische Staatssammlung fiir Palaontologie und Histo- rische Geologic, Munich, West Germany. The NUMBER 56 apophyses are low and inclined (Kier, 1977:16). No auricles are present. Species 6 PLATE 6: FIGURES 3, 4 Mesodiadema?.—Kier, 1977:27, pi. 19: figs. 6-9 [fragment]. MATERIAL.—MCA 3156-S-Z, 5705. This species is easily distinguished by its small, eccentrically situated primary tubercles and very small and numerous secondary tubercles. Two specimens are known; both are only adoral por- tions of an interambulacrum. Specimen MCA 3156-S-Z was illustrated by Kier (1977, pi. 19: figs. 6, 7). Its apophysis is bilobed, very large, occupying most of the adoral border of the in- terambulacrum. The other specimen, MCA 5705 (Plate 6: figures 3, 4), has the apophysis only partially preserved. It is massive, bilobed, and is formed by the thickening of the first three plates in each column of the interambulacrum, with most of the apophysis formed by the second plate in each column. There is no thickening of the interambulacrum at the adradial margin. Species 7 PLATE 6: FIGURES 5, 6 Mesodiadema?.—Kier, 1977:27, pi. 20: figs. 1, 2 [fragment]. MATERIAL.—MCA 3I59-S-Z, 5706. This species is represented by two specimens. The species resembles species 6 but differs in having higher plates. In specimen MCA 3159-S- Z, illustrated in Kier (1977, pi. 20: figs. 1, 2), the apophysis is massive, occupying all the adoral border between the adradial margin. It is bilobed with a narrow indented section medially. The adradial margin thickens only very slightly ador- ally. Each apophysis is formed by the thickening of the first three interambulacral plates of each column with most of the apophysis formed from the second plate. The adradial margin thickens slightly adorally. "Miocidaris" ampezzana Zardini PLATE 7: FIGURES 1, 2 Miocidaris ampezzayia Zardini, 1973:9, pi. 6: fig. 23. MATERIAL.—Holotype: MCA 119. Paratype: MCA 120. This species cannot be generically identified. It resembles most specimens of species 5 in its small secondary tuberculation but differs in hav- ing far larger primary tubercles. Both the holo- type and figured paratype show part of the lan- tern supports. The apophyses are large, includ- ing the full width of the adoral edge of each interambulacrum. They are bilobed and formed from the first three plates of each interambula- crum and are mostly formed from the second plate in each column. The adradial margin is not thickened adorally. Species 8 PLATE 7: FIGURES 3, 4 MATERIAL.—MCA 5708. This species cannot be generically identified. Only one fragment is known. Large crenulate tubercles and sparse secondary tuberculation dis- tinguish this species from the other St. Cassian species. Its apophysis is bilobed and wide, extend- ing across the full width of the interambulacrum at the adoral edge. It is formed of the first two plates in each column. Most of the apophysis is formed from the first plate in one column and the second plate in the other column. Species 9 PLATE 7: FIGURES 5, 6 MATERIAL.—MCA 5709. Although this species cannot be generically identified, it is easily distinguished from the other St. Cassian species that have apophyses. Its large tubercles have but few crenulations on the adoral side of each tubercle. The apophysis is bilobed, formed by the first two or three interambulacral SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY plates in each column. On first impression the first plate in one of the columns appears to be the main constituent, but this plate is really the second in the column—the first plate being al- most completely resorbed. A thickened flange borders the upper edge of the apophysis and extends down to the adoral margin. A small protuberance occurs medially between the lobes of the apophyses. Leurocidaris monlanaro (Zardini) Two specimens, MCA 3I42-S-Z and USNM 370502, show the inside ofthe adoral edge of an interambulacrum. A pronounced ridge occurs along the adoral margin which may be an apo- physis. Nature of Some Ambulacra in St. Cassian Species There are many fragments of ambulacra but most cannot be identified. Some are composed entirely of primary plates (MCA 5710, Plate 7: figure 7) but many have their plates compounded adorally. On some a tubercle occurs on every other plate (MCA 5711, Plate 7: figure 8). On some a tubercle covers a pair of plates with a single plate, lacking a tubercle, separating the pair (MCA 5712, Plate 7: figure 9). All the ambulacra have their porepairs uniserially ar- ranged except for one specimen with its pores biserially arranged. A BISERIAL AMBULACRUM (FIGURE 4; PLATE 8: FIGURES 1, 2) Material.—MCA 5713. This is the only example of biserial pores I have seen in the Triassic. The plates strongly bevel under the adjacent interambulacra as typ- ical in many other St. Cassian species such as Zardinechinus. Plates are enfolded around each other. The plates bearing the outer porepairs (even numbered on Figure 4) appear to be demiplates (not touching the perradial or medial suture), but this separation occurs only on the exterior of the test. As can be seen on an edge view of the ambulacrum along the perradial su- ture, these plates reach the perradial suture be- low the surface of the test. The plates that bear the outer porepairs extend over the tops of the adjacent plates that bear the inner pores, sepa- rating these plates from the adradial sutures (ex- ternally). The perradial portions of these sepa- rated plates are more and more deeply covered by the adjacent plates adorally. This is seen in the edge view along the perradial suture where these "demiplates" (even numbered) become shorter. In the upper, or more adapical, part of the ambulacrum these plates reach the exterior of the test (as in plate number 2 on Figure 4), whereas adorally (as in plate number 16) the plates are deeply covered by the adjacent ambu- lacral plates, with a height less than one-half the thickness of the ambulacrum. Likewise, the plates that bear the inner pore- pairs appear to be occluded (touching the per- radial but not the adradial sutures). This sepa- ration occurs only on the exterior of the test— internally these plates reach the adradial suture. The plates that bear the inner porepairs (odd numbered on Figure 4) extend externally over the plates that bear the outer porepairs, isolating them externally from the perradial suture. Presumably this intricate interlocking of the ambulacral plates would strengthen the ambula- crum and serve to resist the strain imparted by the contraction of the tubefeet when attached to the substratum or a heavy object. Species Discussion Mikrocidaris pentagona (Munster) PLATE 8: FIGURES 3, 4 Cidaris peyitagona Munster in Wissmann and Munster, 1841:42, pi. 3: fig. 8.—d'Orbigny, 1849:206.—Giebel, 1852:316.—Desor, 1855:4.—Laube, 1865a:280, pi. 9: fig. 3; 1865b:325. NUMBER 56 A B FIGURE 4.—Biserial ambulacrum, MCA 5713: A, exterior view; B, interior view; X 12. Campo. Collector: Zardini. Microcidaris pentagona (Munster).—Doderlein, 1887:39.— Lambert and Thiery, 1910:140.—Mortensen, 1928:64, fig. 34.—Zardini, 1973:9, pi. 5: figs. 18, 19.—Kier, 1974:17, 71, 79; 1977:13, fig. 4, pi. 2: figs. 3-6, pi. 3: figs. 1, 2, pi. 4: figs. 1. 2. Cidaris subpentagona Braun in Wissman and Munster, 1841:42, pi. 3: fig. 9.—d'Orbigny, 1849:206.—Giebel, 1852:316.—Desor, 1855:4.—Laube, 1865a:282, pi. 9: fig. 9; I865b:325. [Not Zardini, 1973:13, pi. 5: figs. 16, 17.] Mikrocidaris subpentagona (Braun).—Doderlein, 1887:39.— Lambert and Thiery, 1910:140.—Kier, 1974:17. MATERIAL.—MV 10084. One well-preserved specimen of this species has a diameter of 4.6 mm, height of 2.7 mm, with the diameter of the apical system 1.0 mm and the peristome 2.8 mm. An ambulacrum has 23 plates and an interambulacrum 12. OCCURRENCE.—Milieres. Collector: Giulini. Megaporocidaris mariana Kier PLATE 9: FIGURES 1-5 Megaporocidaris mariana Kier, 1977:23, pi. 15: figs. 6-8, pi. 16. MATERIAL.—MV 10085. Five other specimens (in private collections, unnumbered) can be referred to this species. The dimensions (in mm) are given below. Ht./ No. of No. of Diam. of diam. interamb. amb. apical Diam. of Diam. Height (%) plates plates system peristome 6.0 3.9 65 13 34 - - 8.4 5.4 64 14 34 - - 7.4 - - - 34 - - 6.2 4.1 66 13 36 3.0 3.0 5.3 3.4 64 12 34 2.5 -10 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY OCCURRENCES.—Alpe di Species, Misurina, Rumerlo. Collectors: Giulini, Furlan, Bizzarini. Tiarechinus princeps Neumayr Tiarechinus princeps Neumayr, 1881:169, pi. 2, fig. 4a-c.— Loven, 1883:11, 64, pi. 13: figs. 150-162.—Mortensen, 1935:519, fig. 312a-e.—Kier, 1977:24, fig. 7, pi. 17: figs. 1-5. Two specimens of this species are in a private, unnumbered collection. The best preserved spec- imen has a diameter of 3.1 mm, a height of 1.9 mm, and its periproct has a diameter of 0.6 mm. The diameter ofthe peristome is 1.9 mm and 22 plates are in an ambulacrum, 4 in an interam- bulacrum. OCCURRENCE.—Tamarin. Collector: Giulini. Levicidaris zardinia Kier PLATE 1: FIGURES 6, 7 Leiiicidaris zardinia Kier, 1977:21, pi. 14: figs. 1-5, pi. 15: figs. 1-5. MATERIAL.—MCA 5691, USNM 370501. Twelve other specimens of this species are in a private, unnumbered collection. Eight of them are well enough preserved to show the dimen- sions (in mm) of the test. No. of No. of Diam. of amb. interamb. apical Diam. of Diam. Height plates plates system peristome 13 (est.) 7.3 60 - - - 14.2 8.6 74 11 7.0 - 11.5 6.9 62 11 5.5 - 8.3 4.7 56 11 - - 13 - 66 11 5.7 - 8.2 4.1 48 10 - - 11.4 5.6 62 11 - 5.4 14.5 8.0 - - - - OCCURRENCES.—Misurina, Alpe di Species. Collectors: Bizzarini, Furlan, Pfaiffer, Giulini. Levicidaris furlani, new species PLATE 10: FIGURES 1-6; PLATE 11: FIGURES 1-4 MATERIAL.—Four specimens can be referred to this species. The holotype is slightly distorted but the surface detail is well preserved. Holotype: MV 10086. Paratype: MCA 5715. SHAPE AND SIZE.—The diameter varies from 6.7 to 11.4 mm, with a height from 46 to 52 (mean 48) percent of the diameter. APICAL SYSTEM.—No plates are preserved. The diameter ofthe system varies from 41 to 45 percent of the diameter of the test (D). AMBULACRA.—The ambulacra are straight. The greatest width at the margin is 13-15 per- cent D. The plates are all primaries (Plate 10: figure 3), each having a secondary tubercle along the perradial suture. These tubercles form a double vertical column running medially down each ambulacrum. The pores are in well-devel- oped peripodia, with a high ridge running trans- versely adapical to each porepair. A ridge sepa- rates the pores of a pair. An ambulacrum in the smallest specimen, 6.7 mm in diameter, has 44 porepairs; the largest specimen, 11.4 mm in di- ameter, has 58 porepairs. The outer pore of each pair is situated adapically to its partner. INTERAMBULACRA.—Each interambulacrum is composed of 10 plates in the smallest specimen to 12 plates in the largest. All the tubercles are imperforate except for the most adapical plate of the left-hand column of each interambula- crum. This tubercle is perforate. Its mamelon is smaller than on the tubercle adoral to it. The bosses of the most adapical tubercles are the largest, decreasing in size adorally. On the largest specimen, the mamelon on the most adapical plate of the right-hand column of an interambu- lacrum rises 0.6 mm above the parapet. A ring of scrobicular tubercles encircles each tubercle. They are partially confluent on some of the transverse sutures. The most adapical plate in the right-hand column of each interambulacrum bears no tubercle. PERISTOME.—The peristome has a diameter equal to 44-48 percent the diameter of the test. Slight notches, perhaps gill slits, occur at the edge of the interambulacra. LANTERN SUPPORTS.—No apophyses are pres- ent. Slight auricles are formed by thickening of the adoral-most ambulacral plates. COMPARISON WITH OTHER SPECIES.—The NUMBER 56 II other two species of Levicidaris, including the type-species (L. zardinia), have all their tubercles imperforate, whereas the adapical most tubercle in the left-hand column of each interambulacrum in L. furlani is perforate. This difference broad- ens the definition of Levicidaris but in all other important characters this species is similar to Levicidaris. This species further differs from Levicidaris zardinia and L. pfaifferi in having the tubercle on the most adapical plate in the left-hand column of each interambulacrum smaller than the tuber- cle on the plates adoral to it. Furthermore, the scrobicules are less depressed along their outer rims in L. furlani. OCCURRENCES.—Stolla, Alpe di Species, Mis- urina. Collectors: Furlan, Giulini, Bizzarini. Levicidaris pfaifferi, new species PLATE 11: FIGURES 5-7; PLATE 12: FIGURE 1 MATERIAL.—The holotype is very well pre- served and undistorted. It lacks its apical system and lantern. Holotype: Pfaiffer collection (private) 164. SHAPE AND SIZE.—The diameter of the test (D) is ll.l mm and the height 7.6 mm or 76 percent D. APICAL SYSTEM.—No plates are preserved. The sytem has a diameter of 5.6 mm (est.) or 50 percent D. AMBULACRA.—The ambulacra are straight to slightly sinuate. The greatest width at the ambi- tus is 1.8 mm or 17 percent D. The plates are all primaries (Plate 11: figure 6), each having a secondary tubercle along the perradial suture. These tubercles form a double vertical column running medially down each ambulacra. The pores are situated in well-developed peripodia. Each ambulacrum has 64 porepairs. The peri- podia are well developed with a transverse ridge running across the top of each porepair and a high ridge separating pores of a pair. INTERAMBULACRA.—Each interambulacrum is composed of 10-11 plates. All plates bear a pri- mary tubercle except the first adapical plate of the left-hand column (as viewed from above). None of the tubercles are perforate or crenulate. The mamelons are undercut and largest on the more dorsal plates. Their height is 0.5 mm or 4.5 percent D, and with a horizontal diameter of 1.0 mm. The bosses are well developed. A ring of approximately 15 tubercles surrounds each boss. A few of the more dorsal tubercles are confluent, with no secondary tubercles separat- ing tubercles in the same row. PERISTOME.—The diameter of the opening is 6 mm. Notches, presumably gill slits (Plate 12: figure 1), are present on the edges of the inter- ambulacra. LANTERN SUPPORTS.—Apophyses are absent. The edge of the ambulacra are thickened. COMPARISON WITH OTHER SPECIES.—Levici- daris pfaifferi differs from Levicidaris zardinia, also from the St. Cassian beds, in having wider am- bulacra, 17 percent D as opposed to 12 to 14 in L. zardinia. The interambulacra are narrower in L. pfaifferi, with only the tubercles in the scrobi- cular rings separating adjacent tubercles, whereas in L. zardinia the tubercles are separated by several rows of secondary tubercles. OCCURRENCE.—Misurina. Collector: Pfaiffer. Zardinechinus giulinii^ new species PLATE 12: FIGURES 2-6; PLATE 13: FIGURES 1-4 MATERIAL.—The holotype is well preserved, with the test not distorted. The apical system and lantern are absent. Two paratypes are also well preserved. Holotype: MV 10088. Paratypes: MV 10089 and one unnumbered specimen in a private collection. SHAPE AND SIZE.—The test is small; the holo- type has a horizontal diameter of 8.4 mm, with a height of 4.5 mm or 53 percent D. The mar- ginal outline of the test is pentagonal with inter- ambulacral apices. The test is flattened dorsally with the upper plates at the margin of the apical system horizontal. APICAL SYSTEM.—All the plates of the apical system are absent. The system was large with a diameter of 45-47 percent D. The outline ofthe 12 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY system is pentagonal with the apices in the am- bulacra. AMBULACRA.—The ambulacra are straight and have their greatest width at the ambitus, where the width of an ambulacrum is 13-15 percent D. The plates are all primaries (Plate 12: figures 5, 6) with no change adorally. The pores are situated in well-developed peripodia with a high ridge running transversely adapical to each porepair. A ridge separates the pores of a pair. Each ambulacrum in the holotype has 42 pore- pairs. The outer pore of each pair is adapical to its partner. Each plate bears a secondary tubercle between the porepair and the medial perradial suture. These tubercles form a double row run- ning down the middle of each ambulacrum. INTERAMBULACRA.—Each interambulacrum in the holotype is composed of 11 plates, each bearing a large noncrenulate tubercle, except for the first plate at the margin of the apical system in some of the interambulacra. The tubercle is largest on the most dorsal plates where the ma- melon is very high, rising 0.6 mm above the parapet. The tubercles decrease in size adorally. The mamelons are all perforate except for those below the margin. In each interambulacral area, 4 or 5 of the tubercles are perforate; 5 or 6 are imperforate. The perforations on the dorsal tu- bercles are elongated vertically. No scrobicules or scrobicular ring of tubercles are present. The area surrounding each boss is covered with sec- ondary tubercles of equal size. The first plate in the right-hand column of each interambulacrum (as viewed from above) bears the first tubercle (Plate 12: figure 5). PERISTOME.—The peristome in the holotype has a diameter of 3.6 mm. Slight notches, per- haps gill slits, occur at the edge of the interam- bulacrum. LANTERN SUPPORTS.—No apophyses are pres- ent. A slight thickening of the plates occurs at the edge of the ambulacra. COMPARISON WITH OTHER SPECIES.—This species differs from Zardinechinus lancedelli (Zar- dini), also from the St. Cassian beds, in lacking a scrobicular ring of tubercles, and scrobicules, and in having elongated perforations in the ma- melons. The adoral tubercles are imperforate in this species, whereas all are perforate in Z. lan- cedelli. OCCURRENCE.—Misurina, Stolla, St. Cassiano. Collector: Giulini. Zardinechinus lancedelli (Zardini) PLATE 1: FIGURES 4, 5; PLATE 13: FIGURE 5; PLATE 14: FIGURES 1-3 Cidaris lancedelli Zardini, 1973:12, pi. 5: fig. 24a,b. Cidaris admeto.—Zardini, 1973:12, pi. 5: fig. 34a,b, [not Braun in Wissmann and Munster, 1841]. Cidaris cf. admeto.—Zardini, 1973:12, pi. 6: fig. 1 la,b [not Braun in Wissmann and Miinster, 1841]. Zardinechinus lancedelli {Zardini).—Kier, 1977:16, pis. 9, 10, pi. 11: figs. 1-8. MATERIAL.—MCA 5690, 5716. In addition, there are three almost complete specimens (private collections, unnumbered) and hundreds of fragments (MCA, unnumbered) that can be referred to this species. The dimensions (in mm) of the three almost complete specimens are given below. No. of No. of Diam. of amb. interamb. apical Diam. of Diam. Height plates plates system peristome 6.6 3.7 42 11-12 3.3 3.8 8.8 5.1 52 12 - - 7.2 4.4 48 12 3.2 3.9 OCCURRENCES.—Milieres, Staolin, Campo. Collectors: Zardini, Furlan, Giulini, and Pfaiffer. Literature Cited Desor, P.J.E. 1855-1858. Synopsis des Echinides fossiles. 490 pages, 44 plates. Paris: Chez Ch. Reinwald. Doderlein, L. 1887. Die Japanischen Seeigel. . ./. Theil. Familie Cidaridae und Saleniidae. 59 pages, 11 plates. Stuttgart: E. Schweizerbortische Verlagshandlung. d'Orbigny, M.A. 1849. Prodrome de paleontologie stratigraphique universelle des animaux Mollusques et Rayonnes. Volume 1, 394 pages. Paris. Giebel, C.G.A. 1852. DeutschlandsPetrefacten: Ein systematisches Verzeichn- iss alter in Deutschland und den angrenzenden Ldn- dern vorkomrnenden Petrefacten, nebst Angabe der Syn- onymen und Fundorte. 706 pages. Leipzig. Kier, P.M. 1974. Evolutionary Trends and Their Functional Signif- icance in the Post-Paleozoic Echinoids. Journal of Paleontology, 48(2: memoir 5): 1-96, figures 1-78, table 1, chart 1. 1977. Triassic Echinoids. Switlisonian Contributions to Pa- leobiology, 30: 88 pages, 16 figures, 21 plates. Lambert, J., and P. Thiery 1909-1925. Essai de nomenclature raisonnee des Echinides. 607 pages, 15 plates. Chaumont: Librairie L. Fer- riere. Laube, G.C. 1865a. Die Fauna der Schichten von St. Cassian. Denk- schriften der Kaiserlichen Akademie der Wissenschaf- ten, Mathematisch-Naturwissenschaftliche Classe (Vi- enna), 24:223-296, plates 1-10. 1865b. Die Fauna der Schichten von St. Cassian. Sitzungs- brichte der Mathematisch-Naturwissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschaften, 50(l):319-326. Loven, S. 1883. On Pourtalesia, a Genus of Echinoidea. Kongelige Svenska Vetenskaps-Akademiens Handlingar, new se- ries, 19(7): 1-95, 16 unnumbered figures, plates 1-21. Mortensen, T. 1928. A Monograph ofthe Echinoidea, I: Cidaroidea. 551 pages, 173 figures, 88 plates. Copenhagen: CA. Reitzel. 1935. A Monograph of the Echinoidea, II: Bothriocidaroida, Melonechinoida, Lepidocentroida and Stirodonta. 645 pages, 377 figures, 89 plates. Copenhagen: CA. Reitzel. Neumayr, M. 1881. Morphologische Studien uber fossile Echinoder- men. Sitzungsberichta der Mathematisch-Naturwissen- schaftlichen Classe der Kaiserlichen Akademie der Wis- senschaften, 84(1): 143-176, plates 1, 2. Wissmann, H.L., and G. Munster 1841. Beitrage zur Geognosie und Petrefacten-Kunde des Su- dostlichen Tirol's Vorzuglich der Schichten von St. Cas- sian. 125 pages, 16 plates. Bayreuth. Zardini, R. 1973. Fossili di Cortina: Atlante degli echinodermi cassiani (Trias Medio-superiore) della regione dolomitica attorno a Cortina d'Ampezzo. 29 pages, 21 unnumbered figures, 22 plates. Cortina d'Ampezzo, Italy: Foto Ghedina. 13 14 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 1 Unidentified species 1-3. Adoral portion of an ambulacrum showing the auricles produced by the first three plates of each column, MCA 5689, x 14. Campo. Collector: Zardini. Zardinechinus lancedelli (Zardini) 4, 5. Exterior, interior views of adoral portion of ambulacrum and interambulacrum of MCA 5690, X 16. Note absence of apophyses and presence of auricles. Campo. Collector: Zardini. Levicidaris zardinia Kier 6, 7. Exterior, interior views of interambulacrum showing lack of apophyses, MCA 5691, X 12. Misurina. Collector: Zardini. NUMBER 56 15 16 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 2 Species 1 1,2. Exterior, interior views of MCA 5692 showing absence of apophyses, X 9. Campo. Collector: Zardini. 3, 4. Exterior, interior views of MCA 5693 showing absence of apophyses but presence of protuberance, which was probably the site of insertion ofthe lantern protractor muscles, X 10. Campo. Collector: Zardini. 5, 6. Exterior, interior views of MCA 5694 showing absence of apophyses, X 8. Campo. Collector: Zardini. NUMBER 56 17 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 3 Species 2 1, 2. Exterior, interior views of MCA 192 showing absence of apophyses, X 14. Cason dei Caai. Collector: Zardini. 3,4. Exterior, interior views of MCA 5696 showing absence of apophyses, x 16. Campo. Collector: Zardini. 5,6. Exterior, interior views of MCA 168 showing absence of apophyses, X 12. Rumerlo. Collector: Zardini. NUMBER 56 19 20 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 4 Species 2 1,2. Exterior, interior views of MCA 5698 showing absence of apophyses and presence of a protuberance presumably for attachment of protractor muscles, X 12. Campo. Collector: Zardini. Species 3 3, 4. Exterior, interior views of MCA 5699 showing absence of apophyses and presence of a large protuberance. Figure 3 (text) shows the location ofthe plate sutures, X 12. Campo. Collector: Zardini. "Miocidaris" adrianae Zardini 5, 6. Exterior, interior views of MCA 5700 showing absence of apophyses, X 16. Campo. Collector: Zardini. NUMBER 56 21 .^.*% 22 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 5 "Miocidaris" adrianae Zardini I, 2. Exterior, interior views of MCA 5701 showing absence of apophyses and presence of large protuberance presumably for attachment of protractor muscles, X 9. Campo. Collector: Zardini. 3,4. Exterior, interior views of MCA 5702 showing absence of apophyses, X 14. Campo. Collector: Zardini. 5,6. Exterior, interior views of MCA 121 showing absence of apophyses and presence of depression on interior opposite the primary tubercle, X 10. Misurina. Collector: Zardini. NUMBER 56 23 rif.r.'JfX ■*^1,f.r.'. 24 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 6 Species 5 1, 2. Exterior, interior views of MCA 5704 showing absence of apophyses and presence of a protuberance, X 10. Campo. Collector: Zardini. Species 6 3, 4. Exterior, interior views of MCA 5705 showing large apophysis, X 9. Misurina. Collector: Zardini. Species 7 5, 6. Exterior, interior views of MCA 5706 showing large apophysis, X 9. Campo. Collector: Zardini. NUMBER 56 25 26 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 7 "Miocidaris" ampezzana Zardini 1, 2. Exterior, interior views of MCA 120 showing large apophyses, X 6. Campo. Collector: Zardini. Species 8 3, 4. Exterior, interior views of MCA 5708 showing large apophyses, X 12. Alpe di Species. Collector: Zardini. Species 9 5, 6. Exterior, interior views of MCA 5709 showing well-developed apophyses, X 8. Campo. Collector: Zardini. Ambulacra of unknown species 7. Ambulacra composed of primary plates, MCA 5710, X 12. Campo. Collector: Zardini. 8. Ambulacra with a tubercle on alternating plates, MCA 5711, X 16. Misurina. Collector: Zardini. 9. Ambulacra with tubercles covering two plates periodically separated by a plate lacking a tubercle, MCA 5712, x 16. Misurina. Collector: Zardini. 7. NUMBER 56 27 28 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 8 Species ? 1,2. Exterior and perradial views of a half ambulacrum MCA 5713, showing biserial arrangement of porepairs, X 24. A drawing showing the arrangement of the plates is on Figure 4 (text), Campo. Collector: Zardini. Mikrocidaris pentagona (Munster) 3, 4. Top, side views of MV 10084, X 18. Milieres. Collector: Giulini. NUMBER 56 29 30 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 9 Megaporocidaris mariana Kier 1, 2, 4, 5. Side (1,2), dorsal (4), and ventral (5) views of MV 10085, X 12. Alpe di Species. Collector: Giulini. 3. Ambulacrum of same specimen, X 36. NUMBER 56 31 32 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 10 Levicidaris furlani, new species 1-4. Dorsal, ventral, and side views ofthe holotype MV 10086, X 6. Stolla. Collector: Giulini. 5,6. Dorsal, ventral views of paratype MCA 5715, X 6. Misurina. Collector: Furlan. Side views of this specimen are on Plate 11: figures 1, 2. NUMBER 56 33 34 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 11 Levicidaris furlani, new species 1, 2. Side views of paratype MCA 5715, X 6. Misurina. Collector: Furlan. Dorsal and ventral views of this specimen are on Plate 10: figures 5, 6. 3, 4. Side and dorsal view of paratype MV 10087, X 10. Alpe di Species. Collector: Giulini. Levicidaris pfaifferi, new species 5-7. Side and dorsal views ofthe holotype (Pfaiffer collection 164), X 7. Misurina. Collector: Pfaiffer. A ventral view of this specimen in on Plate 12: figure 1. NUMBER 56 35 36 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 12 Levicidaris pfaifferi, new species 1. Ventral view of holotype (Pfaiffer collection 164), X 7. Misurina. Collector: Pfaiffer. Zardinechinus giulinii, new species 2-4. Side, dorsal views ofthe holotype MV 10088, X 8. Stolla. Collector: Giulini. 5, 6. Ambulacrum ofthe same specimen at midzone and adorally, X 16. NUMBER 56 37 38 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 13 Zardinechinus giulinii, new species 1-4. Dorsal, ventral and side views of paratype MV 10089, X 10. Misurina. Collector: Giulini Zardinechinus lancedelli (Zardini) 5. Side view of MCA 5716, X 18. Campo. Collector: Zardini. More photographs of this specimen are on Plate 14. NUMBER 56 39 40 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 14 Zardinechinus lancedelli (Zardini) 1-3. Side, dorsal, and ventral views of MCA 5716, X 18. Campo. Collector: Zardini. 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