^A'iY DORA ROGICK R9GICK COLLECTION 1^ 7G. - ^ - ) M^moires de TAcad^mie Royale des Sciences et des Lettres de Danemark, Gopenhagne, 7™« serie, Section de8 Sciences, t. X, a" 1. MARY DORA kUGICK STUDIES ON THE CYCLOSTOMATA OPERCULATA BY G. M. R. LEVINSEN WITH 7 PLATES AND 2 FIGURES IN THE TEXT D. Kgi.. Danske Vidensk. Selsk. Skrifter, 7. R.ekke, naturv. og Ihathematisk Afd. X. 1 '•<>»3J«(c>-«- ROGICK COLLECTION K0BENHAVN HOVEUKOMMISSION^R: ANDR. FRED. H0ST & S0N, KGL. HOF-BOGHANDEL BIANCO I.UNOS BOGTRYKKERI 1912 Prls: 4 Kr. 45 0re I Det Kgl. Danske Videnskabernes Selskabs Skrifter, 6*6 Rsekke. Naturyidenskabelig og mathematisk Afdeling. Rr. 0re I. med 42 Tavler, 1880—85 29. 50. 1. I'rjii, K. UiidersBgelser over Lysets Brydning i Dampe og tllsvarende Vaedsker. 1880 • 66. i. Boas, J. E. V. SUidier over Decapodeines Slaeglskabsforhold. Med 7 Tavler. Resume eii fraiifais. 1880 8. 50. 3. Steenstrup, Jap, Sepiadarium og Idiosepius, to nye Sliegter af .Sepieines Faiiiilie. Med Bemairkiiiiiger om to beslaegtede Former Sepioloidea D'Orb. og Spirula Lmk. Med 1 Tavle. Resume en frajieais. 1881 1. 35 4. Coldliig, A. Nogle Undersegelser over Slormen over Nord- og Mellem-Europa af 12'« — Hi^e Novb. 1872 og over deii derved fremkaldte Vandflod i 08tersoen. Med 23 Planer og Kort. Resume en franfais. I88I 10. • 5. Uoas, J. E. V. Om en fossil Zebra-Form fra Brasiliens Campos. Med et Tillacg om lo Arter af .Sliegten Hippldion. Med 2 Tavler. 1881 2. • 6. Steeii, A. Integration af en linear DitVerentialligning af anden Orden. 1882 50. 7. Krabbe, U. Nye Ridrag til Kundskab om Fuglenes Raendelorme. Med 2 Tavler. 1882 1. 35. 8. liaiinover, A. Den nienneskelige Hjerneskals Rygning ved Anencephalia og Misdannelsens Forhold til HJerneskallens Primordialbrusk. Med 2 Tavler. Extrait el explication des planches en franfais. 1882 I. 60. 9. Den menneskelige Hjerneskals Rygning ved Cyclopia og Misdannelsens Forhold til Hjerneskallens Primordialbrusk. Med 3 Tavler. Extrait et explic. des planches en franfais. 1884 4. 35. 10. Den menneskelige Hjerneskals Rygning ved Synolia og Misdannelsens I'orhold til Hjerneskallens Pri- mordialbrusk. Med 1 Tavle. Extrait et explic. des planches en franfais. 1884 1. 30. 11. Lehinanii, A. Forsog paa en Forklaring af Synsvinklens Indllydelse paa Upfattelsen af Lys og Farve ved direkte Syn. Med 1 Tavle. Resume en franfais 1885 1. 85. H, med 20 Tavler, 1881—86 20. ■• t. Wanning, Eu^. Familien Podoslemaceae. 1^'° Afhandling. Med 6 Tavler. Resume et explic. des planches ' en franfais. ISSI 3. 15. 2. Lorenz, L. Om Melallernes Ledningsevne for Varme og Elektricltet. 1881 1. 30. 3. Warming, Eug. Familien Podostemaceae. 2''«° Afhandling. Med 9 Tavler. Resume et explic. des planches en franfais. 1882 5. 30. 4. Chrlstensen, Odiu. Ridrag til Kundskab om Manganets liter. 1883 I. 10. 5. Lorenz, L. Farvespredningens Theori. 1883 60. 6. Gram, J. P. Undersegelser ang. Msengden af Primtal under en given Grsnse. Resume en franfais. 1884 4. • 7. Loreui, L. Restemmelse af Kviksalvsejiers elektriske Ledningsmodstande i absolut eleklromagnetisk Maal. 1885 80. 8. Traustedt, M. P. A. Spolia Atlantica. Ridrag til Kundskab om Salperne. Med 2 Tavler. Explic. des planches en franfais. 1885 3. » 9. Bohr, Cbr. Om lltens Afvlgelser fra den Royle-Mariotteske Lov ved lave Tryk. Med 1 Tavle. 1885 ... 1. » 10. Undersegelser over den af Rlodfarveslolfet optagne lUniajngde udferte ved Hja;lp af et nyt AbsorpUo- meter. Med 2 Tavler. 1886 1. 70. 11. Thicif, T.N. Om Deflnitionerne fer Tallet, Talarterne og de tallignende Restemmelser. 1888 2. » III, med 6 Tavler, 1885—86 16. • 1. Zeuthen, 11. C. Keglesnitslseren i Oldllden. 1885 10. • 2. Levlnseii, G. 111. R. Spolia Atlantica. Om nogle pelagiske Annulata. Med 1 Tavle. 1885 1. 10. 3. Hung, G. Selvregistrerende meteorologiske Instrumenter Med 1 Tavle. 1885 1. 10. 4. IHelnert, Fr. De eucephale Myggelarver. Med 4 dobb. Tavler. Resume el explic. des planches en francais. 1886 6. 75. IV, med 25 Tavler. 1886—88 21. 50. 1. Boas, J. E. T. Spolia Atlantica. Ridrag til Pteropodernes Morfologi og Systematik samt til Kundskaben om deres geografiske Udbredelse. Med 8 Tavler. Resume en franfais. 1886 10. 50. 2. Lehmann, A. Om Anvendelsen af MIddclgradationernes Melode paa Lyssansen. Med I Tavle. 1886. ... 1. 50. 3. llannoTer, A. Primordialbrusken og dens Forbening i Truncus og Extremiteter hos Mennesket for Fed- selen. Extrait en franfais. 1887 1. 60. 4. Lutken, Chr. Tillseg til ^Ridrag til Kundskab om Arterne af SIxgten Cyamus Latr. eller Hvallusene-. Med 1 Tavle. Resume en franfais. 1887 60. 5. Fortsatte Ridrag til Kundskab om de arkliske Dybhavs-Tudsefiske, siErligt SliBgteu Hiinantolophus. Med 1 Tavle. Resume en franfais. 1887 ■ 75. 6.. Kritiske Studier over nogle Tandhvaler af Slaegterne Tursiops, Orca og Lafjenorliijnchus. Med 2 Tavler. Resume en francais 1887 4 75. 7. Roefofd, E. Studier i Plalosoforbindelser. 1888 1. 30. 8. Warming, Eug. Familien Podostemaceae. 3<"^ Afhandling. Med 12 Tavler. Resume et explic. des planches en franfais. 1888 6. 45. V, med 11 Tavler og 1 Kort. 1889—91 15. 50. i. Lutken, Chr. Spolia Atlantica. Ridrag til Kundskab om de Ire pelagiske Tandhval-JJaegter Steno, Del- pliinna og Truddplnnnti. Med 1 Tavle og 1 Kort. Resume en franfais. 1889 2. 75. 2. Talenllner, D. De endelige Transformations-Cruppers Theori. liesunie en franfais. 1889 5. 50. 3. Hansen, U.J. Cirolanidx ct familiie nonnullx propinqua: Musci llauniensis. Et Ridrag til Kundskaben om nogle I'amilier af isopode Krebsdyr. Med 10 Kobbertavler. Resume en franfais. 1890 9 SO. 4. Lereui, L. Analytlske Undersegelser over PrinUalma;ngderne. 1891 • 75.- (Forl83ttes paa Omslagets S. 3.) MARY DORA ROGICK STUDIES ON THE CYCLOSTOMATA OPERCULATA BY G. M. R. LEVINSEN WITH 7 PLATES AND 2 FIGURES IN THE TEXT D. Kgi,. Danskk Vidensk. Selsk. Skhii- teh, 7. R^kke, naturv. or, mathematisk Afd. X. 1 •^X-SJUcS-. K0BENHAVN HOVEDKOMMISSION.EK: ANDR, FRED, H0ST&S0N, KGL. HOF-BOGHANDEL BIANCO LUNOS BOGTRYKKERI 1912 PKEFACE. Ma,„ ly years ago I undertook a study of the Danish cretaceous Brijozoa the results of which have not yet been pubiislied, and among the species studied by me were also a small number belonging to the family Melicerititidae (Eleidae d'Orb.) to the members of which d'Orbigny ascribes a calcareous operculum the presence of which, however, has been denied by all the later authors who interprete the supposed operculum as a closure-plate of the same nature as that which has been found both in the Cijclostomata and the Cheilostoinata. I came however to the result that d'Orbigny was right in his interpretation of the named structure, and I have published some remarks on this subject in my preliminary communication sStudies on Bryozoa« '). As later I wished to make a more comprehensive study of this interesting group I extended my investigations over a number of foreign species, and in this etTort I have been supported by several colleagues abroad. In the first instance I owe a debt of gratitude to Dr. F. Canu the author of so many valuable works on fossil Bryozoa, who has not only sent to me a large number of French species defined by him, but also helped me to acquire materials containing cretaceous bryozoa from a number of French localities. A similar material from a few other French localities has been sent me by Mr. G. Dollfus, and to Mr. M. FiLiozAT who has made a special study of the cretaceous Bryozoa from Vendome I owe the possession of a number of species from that locality. For the gift or loan of specimens I am also indebted to Mr. A. W. Waters, Dr. E. Pergens, Prof. Dr. G. Steinmann, Bonn and Prof. Dr. H. Wegner, Miinster, and lastly I have been able to acquire a collection of cretaceous Bryozoa from the Chatham chalk, by the aid of Mr. W. Gamble. To all these gentlemen I offer my sincere thanks. Zoological Museum of Copenhagen. /"* A/T R T August 16, 1912. ^- "^- ^- •L'EVINSEN. 16. Historical Introduction. GoLDFuss ') was the first author, who described a species belonging to this di- vision namely Ceriopora gracilis while the first genus Meliceritites has been instituted by RoEMER ^) for the three species Mel. gracilis (Goldf), Mel. Roemeri (Hag) and Mel. porosa Roemer. The name Meliceritites is derived from Melicerita, a genus founded by Milne Edwards'') for a fossil species of the present genus Celliilaria (Cellaria), Cel. Charlesworthii, and the main character upon which the genus was founded is the arrangement of the zooecia in continuous transverse series, a character which at the present state of systematic knowledge cannot be regarded as sufficient for the institution of a genus, and therefore the name Melicerita must be regarded only as a synonym to Cellularia. Roemer's genus is defined in the following manner: »Runde Slamme deren sechsseitige Zellen mit einer Ecke und nicht wie bei alien bisher beschriebenen Arten, mit einer Seite nach oben gerichtet sind und die gemein- schaftliche Scheidewand zweier Zellen der hoheren Reihe tragend«. After this de- finition he adds the following apparently contradictory information: »Gleiche Zellenbildung findet sich auch bei Eschara und hat Milne Edwards hier darauf die Gattung Melicerita gegrundet«. As stated above however it is not the form of the zooecial areas but the arrangement of them in transverse series which has induced Milne Edwards to found a new genus for this species. It is easy to understand that Roemer has derived the name of his new genus from Melicerita as the three species described by him agree with Mel. Charlesworthii in the possession of rhom- bic-hexagonal concave zooecial areas, surrounded by distinct ridges and, besides, arranged in transverse series, and as he has not seen the long tubular pari of the zooecia hidden within the colony his only motive to divide Meliceritites from Meli- cerita seems to be the different form of the colony, which in the latter is a two- layered plate while in the former it is ramose with rounded branches. V. Hagenow ') is the first author who refers these forms to a special division which he named Salpingina and characterized in the following manner: »Ange- wachsene verastelte Polypenstocke, mit langen Rohrenzellen, welche sich kurz vor der Miindung trompetenartig, fast sackformig erweitern, und ausserlich mil einer kalkigen oder hornagtigen Membran verschlossen sind. In dieser Meinbran befindet sich die kleine, verschieden gestaltete Miindung, die warscheinlich bei Allen mit einer Klappe versehen war und die bei einigen Arten noch erhalten ist.< At another place he names this >Klappe« operculum: vEin Operculum ist bei Einigen, vielleicht bei Allen vorhanden«. To this division he refers the two genera Escharites and Inversaria the former of which also comprises Roemer's Meliceritites. Apart from the circumstance ») 1. p. 35, pi. X, figs 11 a— c. ■-') 6, p. 18, pi. V, fig. 13. "■) 2, p. 34.5, pi. 12, (ig. 19. *) 5. p. 55. that Hagenow's division only embraces freely growing species it is founded on the examination of a few badly conserved species of which only a single Eschariles gra- cilis with certainty can be referred to the Eleidce, and as species belonging to this division have never been found in the tertiary formation the following statement of the author distinctly shows that his Salpingina also embraces common cyclosto- matous species: Alle sind fossil und gehoren dem Jura, der Kreide und der Ter- tiarbildung an«. The presence of an operculum in the Salpingina should be a real difference from the Cgclostomaia, but there is no evidence that the author has seen a real operculum in any of the species referred by him to this division. As a species provided with such an operculum he names Escharites fFeliceaJ velata Hag., but this species belongs to quite another division, the Ceidae, which are not provided with an operculum, and what Hagenow has seen is only a closure-plate. d'Orbigny ') in 1852 founded a division to which he gives the name Centri- fagines operculines and which he defines as follows: » Cellule centrifuginee toujours pourvue d'un opercule. Colonic tres variable dans sa forme composee de cellules generalement pen saillantes, mais toujours pourvues d'un opercule: partie testacee ou calcaire, s'ouvrant comma une porta pour laisser sortir I'animal*. As to the operculum he later adds: >>D'ailleurs s'il pouvait encore rester quelques doutes a cet egard la presence de ces opercules encore restes en place dans beaucoup d'es- peces fossiles, vient entierement les lever et donner la preuve que cet opercule exi- stait«. To this division d'Orbigny refers two families, the Eleidae and the Myrio- zoumidae, the last of which contains the exstinct genus Foricnla and the recent genus Myriozoum. The last named genus, however, belongs to the Cheilostomata. The above quotation leaves no doubt that d'Orbigny has founded his division Eleidae on the presence of an operculum, and it is a curious fact that all the modern authors with the exception of the present though accepting this division, at the same time deny that its member possesses an operculum, explaining the calcareous plate which may be found closing the aperture of more or less zooecia as a closure-plate. Not a single author even mentions this statement of d'Orbigny. Hamm "') in 1881, founded a very artificial division, the S/!(///!fl/o/)or(;!« to which he besides two inoperculate cyclostomatous genera Stiginatopora Hamm {-Hanimia Grey) and Cyrtopora Hag. also referred the operculate genus Melicerilites. He characterizes this division as follows: > Die Zellen sind ahnlich wie die der Tubuliporiden beschaffen. Statt dass sie aber in der mitlleren Axe des Slammes entspringen legen sie sich rings um ein senkrecht stehendes, im Querschnitt rundliches Biindel von langen cylindrischen unter einander parallelen R6hren«. He divides the Stigmatoporina in two groups the second of which (Melicerititesjhe characterizes by the trumpet-shaped distal enlargement of the zo- oecia: ». . zweitens in solche, deren Zellen lang, anfangs sehr dunn sind und erst an der Mundung sich plotzlich trompetenforinig erwaitern«. As we have seen Hagenow has al- ») 7, p. 605. '-] 8, p. 45, ready used this character for his Salpingina, but Hamm has overlooked that the nar- row tubes of the axial bundle are only the proximal parts of the zooecia, and as we shall see later, this error is repeated tjy Mahsson. As to the operculum spoken of by d'Orbigny he expresses the following opinion'): »Dieser Deckel ist indesscn kein deni beweglichen Deckel der chiloslomen Bryozoen analoges Organ, sondern eine einfache Kalkwand, mittelst deren sich das Thier bei zunehmendem Alter oder uni sich gegen aussere Einlliisse zu schutzen, in die Zelle einkapselt; er ist also ein blosses biologisches Produkt. Dem entsprechend findel sich derselbe nicht nur bei den Opercules von d'Ohbigny, sondern ebenso bei den Clausidae D'Orbg., Diaslo- pora, Terebellaria, Osculipora, Heleropora etc. also bei Formen von sehr verschiede ner Natur. Zur Systematik darf er daher nicht verwendet werden«. Marsson"), in 1888, united the Ceidea and the Eleidea into a division Metopopo- rina, and the Eleidea he characterized as follows: >. . . . die Hauplzellen bei den cylindrischen Arten auf der Aussenseite eines centralen cylindrischen Biindels langer, Rohrenzellen entspringend, (ob audi bei den llachen, blattartigen Formen die Haupt- zellen aus einer Schichl von Nebenzellen entspringen, bedarf einer weiteren Unter- suchung), auf der Oberllache des Stocks dicht an einauder gedrjingt, mit melir oder weniger rhombisch-sechseckigen Stirnseiten ausmiindend, die im vorderen Theile eine dreiseitige Miindung tragen«. In the diagnose of Melicertites he mentions the trumpet-shaped expansion ,of the zooecia, and the manner in which he speaks about the operculum seems to show that he inlerpretcs it as a preliminary calcareous co- vering which later disappears: sMiindung .... anfangs durch eine Kalkmembran geschlossen und mehr hervortretend <. Pergens ') in 1890, separated the two groups, the Ceidae and the Eleidae, united by Marsson in his Metopoporina, and the latter of them, to which he gives the name Melicertitina he defines in the following manner: »Les zoecies se dilatent vers I'extremile en forme de trompelle; Torifice n'occupe qu'une partie du dianietre trans- versal des zoecies et est situe a leur partie distale. Les ovicelles soul situees a I'in- terieur des colonies, entre les zoecies, ou occupant leur place, et communiquant avec I'exterieur par une ouverture triangulaire a base proximale et a pointe plus ou moins allongee«. The slructures which Pergens as d'Orbigny and Marsson re- gards as ovicells are as Waters has pointed out very like the Cheilostomalous avi- cularia, and the real gonozoocia have not been seen by Pergens, who, besides, speaks about Ihe supposed ovicells as if there were two differenl kinds, some placed within the colony and others taking the place of zooecia. This however is not the case. Pergens as Hamm and Marsson means that the zooecia of Meliceritites take their origin from a central bundle of narrow tubes, and like these authors he also denies the presence of a real operculum*): »les Opercules renferment deux families: les MyriozoLimidd' et les Eleidce. Le tj'pe de la premiere est reellement pourvu d'un opercule, c'est done un Cheilostome; les secondes n'ont ])as d'opercule, mais souvent 8, p. 9. 2, 10, p. 45. ■') 9, pp. 325, 391, ') 9. P- 320, une membrane calcareuse transversale, caractere qu'elles partagent avec des Hetero- pora, des Entalophora et une quantite d'autres genres. — Waters') who was the first to point out the likeness between the »cellules accessoires« or »cellules ovariennes« of d'Orbigny and the Clieilostoniatous avicu- laria is inclined to think, that the Eleidae have been provided with a chilinous operculum. He says about that, referring to transverse sections o{ Meliceritites Roy ana: »In transverse sections (figs 5 and 11) a contraction formed by a curved plate is seen on each side just below the opening. Possibly an operculum has an attachment here, but of this I have not been able to satisfy myself«. He does not mention d'Orhigny's calcareous opercula, and, therefore, no doubt regards them in the same way as the other authors, namely as closure-plates. On the whole Waters seems inclined to think that the Eleida; are nearer related to the Clieilostomata than to the Cyclostomata. Gregory-) who regards the Eleidae only as a family of his suborder Cyclosto- mata lubulata defines this family as follows: »Cyclostomata tubulata in which the ') 12, p. 48, pi. VI. 2) Gregory (14, p. 285) wlien speaking about d'Orbigny's institution of the family £/cidachas quite misunderstood this author. He says as follows: »D'Orbigny founded this family in 1853 for a series of Bryozoa with anomalous characters, of which the most striking was the presence of a series of modified zoiecia that he described as ^cellules accessoires<. These accessory structures are of two types, large superficial marsupial chambers, and cells with triangular or elongated apertures and a platform iiarallel to the surface of the zoarium. The marsupial chambers were described by d'Orbigny as Dcellules ovariennes's and they are clearly gono- cysts or gontecia. The nature of the second set is more important. D'Orbigny recognized that they sometimes occur on the same specimen as »cellules ovariennes«, and therefore cannot be ovarian. He suggested that they may be male cells or sperm-cells. Nevertheless Pergens and Marsson subsequently described them as ovicells . Every reader nuist understand the above statement in that manner 1) that r/ORBioNv has founded the family Eleidae chiefly on account of its possessing modified zooecia (licellules accessoiresi), and that he has only found such modified zooecia in the Eleidae. Further 2) that d'Orbigny has pointed out the pre- sence of marsupial chambers (gonozoa?cia) in the Eleidae, and 3) that he has interpreted some of the modified zotecia found in the Eleidae as „cellules males". As to the first ])ointthe above quotation of d'Orbigny shows that he has founded tfie division Centrifitgines opcrciilincs not on account of its [)Ossessing modified zooecia, but because it possessed calcareous opercula. and for i)oth the families belonging to tliis division he only mentions in somewhat different terms that they sometimes possess cellules accessoires* which structures, besides, he has found not only in this division, but also in the Cheilostomata and the Cyclostomata. As to the two other points d'Orbigny has nowhere mentioned or pictured an Eleid gonozosecium, and what he says about , cellules m;tles" only concerns the Cheilostomata. The structure of the avicularia be- ing at that time unknown d'Orbigny is evidently most inclined to look upon the modified zo- oecia as gonozotecia (»cellules ovariennes«), but as in Eschara') they are found together with zooecia wearing ooecia he cannot escape regarding the latter as »cellules ovariennes<, a view held further by all his i)redccessors, and therefore he asks whether they may not liere be regarded as DCellules m;iles«. In the representation he gives of the linjozoaires centrifutjines, a division which besides some C