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ADDITIONS TO THE LIBRARY
OF THE
Connecticut Academy of Arts and Sciences,
By GIrT AND EXCHANGE FROM JULY 1, 1895, To Oct; 1, 1899.
ALBANY.—New York State Library. Annual report. LXXVI-LXXX, 1893-97. 8°. Bulletin. Additions. No. II-LY, 1894-96. 89°. Legislation. No. VI-X, 1895-99. 89. Bibliographies. No. II-IV, 1897. 8°. New York State Museum of Natural History. Annual report. XLVII-XLIX, L. 1, 1894-96. 8°. Bulletin. No. 14-23, 1895-98. 89°. University of the State of New York. Extension bulletin. No. IX-X XVIII, 1895-98. §9°. American Association for the Advancement of Science. Proceedings. Meeting XLIV—XLVII, 1895-98. Salem, 1896-98. 8°. ANNAPOLIS.— United States Naval Institute.
Proceedings. Vol. XIX 1, XXI 2-4, XXII-XXIV, XXYV. 1, 2, 1893-99. 8°.
AustTiIn.— Texas Academy of Science. : Transactions. Vol. I. 5, 1897. 8°. BALTIMORE.—Johns Hopkins University. American chemical journal. Vol. XVII. 8-10, XVIII-XXI, XXII. 1-3, 1895-99. 8°. University circulars. No. 121-141, 1895-99. 4°. BuLvuE Hitu.—Veteorological Observatory. Bulletin. No. I, III, 1899. 49. Boston.—American Academy of Arts and Sciences. Proceedings. Vol. XXX-XXXIV, XXXV. 1-3, 1894-99. 8°. —— Massachusetts State Board of Agriculture.
The gipsy moth. A report on the work of destroying the insect in the Commonwealth of Massachusetts. By E. H. Forbush and C, H. Fer- nald., Boston, 1895. 8°.
Society of Natural History.
Memoirs. Vol. V. 1-5, 1895-99. 4°.
Proceedings. Vol. XXVI.4, XXVII, XXVIII, X XIX. 1-5, 1895-99. 8°.
Occasional papers. IV. Vol. I. 1, 2, 1893-94. 8°.
_ BurraLo.—Society of Natural Sciences. Bulletin. Vol. V.5, VI. 1, 1897-98. 8°.
CAMBRIDGE.— Harvard College.
Annual reports of the president and treasurer. 1894-5, 1895-6, 1896-7,
1897-8. 8°. 13 (4)
vi Additions to the Library.
CAMBRIDGE. —Astronomical Observatory of Harvard College. Annals, Vol. XXIII. 2, XXVIII. 1, XXX. 4, XXXI. 3, XXXIV-XXXVI, X XXVIII. 1, XXXIX.1, XL. 4, 5, XLI. 3-5, XLII. 1, 1894-99. 40. Annual report. L-LIIT, 1895-98. 8°. . —— Museum of Comparative Zoilogy at Harvard College. Memoirs. Vol. XIX-XXII, XXIII. 1, 1895-97. 4°. Bulletin. Vol. XXV. 9, XXVIII. 2-7, XXVIII-XXXI, XXXII, XXXITI, XXXYV. 1-2, 1894-99. 8°. Annual report. 1894-5, 1895-6, 1896-97, 1897-8 1898-9. 8°. —_—Fntoniological Club. Psyche. No, 231+248, 1895-96. 8°. Cuare, Hrw—Zvisha Mitchell Scientific Society. Journal, “Vol. XI, 2,\XII, XIII. 2, XIV. 2, XV. 2, 1894-98, 8°. Cur1caGco.—Academy of Sciences, Bulletin... Vol. I, I. 1, 2, 1883-95. 8°. Anntal report. XX XVIII-XL, 1895-97. 8°. Geographical and Natural History Survey. Bulletin. No. I, If, Il. 1, 1896-98, A naturalist in Mexico, being a visit to Cuba, Northern Yucatan, and Mexico, | By Frank Collins Baker. Chicago, 1895. 8°. — Field Columbian Museum. Publications. No. 2-39, 1894-99. 8°. —— University of Chicago. Journal of geology. Vol. IIL. 4-8, IV-VI, VII. 1-5, 1895-99, 89.
CINCINNATI.— Observatory. \ Publications. No. XIV, 1899. 8°. ~ Ss Historical sketch, 1843-1893. 8°. [ |
—— Society of Natural History.
Journal. Vol. XVIII, XIX, 1896-98. 8°. Ce.
CoLorabo SpriInGs.— Colorado College Scientifie Society. Colorado College studies. VI, 1896. 8°. re! | 0 Des Mornes.—Jowa Academy of Sciences. F Proceedings. Vol. II-VI, 1894-98. 8°. ¢r. | Towa Geological Survey. Annual report. IV, V, 1894-95. 8°. ‘GRANVILLE.—Denison University. * Bulletin of the scientific laboratories. Vol. IX, X, XI. 1-8, 1895-99. 8°. Journal of comparative neurology. Vol. V. 2-4, VI-VIII, IX. 1, 2, 1895-99. Sa. HArrispure.—Second Geological Survey of Pennsylvania. Summary finalreport. Vol. III. 1,2, and general index. 1895. 8°. Har trorp.—Connecticut Historical Society. Annual report, May 26, 1896. 8°. ; IrHaca.—Journal of physical chemistry. Vol. I, IL, III. 1-6, 1896-99. 8°. JEFFERSON City.—Missouri Geological Survey. Publications. Vol. IV-IX, 1894-96, 8°. LAWRENCE.— University of Kansas. Kansas University quarterly. Vol. IV-VII,. VIII, A. 1-8, 1895-99. 89°. University Geological Survey of Kansas. Vol. I-IV, 1896-98. 8°. Mineral resources of Kansas. 1897, 8°. University of Kansas Experiment Station. Annual report. I-V, 1891-95. 8°, Kansas Board of Irrigation, Survey and Experiment. Report. 1895-96. 8°. Common injurious insects of Kansas. By Vernon L. Kellogg. 1892. 8°. Quarter-centennial history of the University of Kansas, 1866-1891. Tope-- ka, 1891, 8°,
ee Ce eee
y
Additions to the Library.
Mavison.— Washburn Observatory. Publications. Vol. IX, X. 1, 1893-98. 8°. Wisconsin Academy of Sciences, Arts and Letters. fransactions. Vol. X, XII. 1, 1891-98. 8°. MANCHESTER, N. H.—Historic Association. Collections, Vol. I. 1, 1896. 8°. MERIDEN.— Scientific Association. Transactions. Vol. VII, VIII, 1895-98. 8°. MINNEAPOLIS. —Minnesota Geological and Natural History Survey. Annual report. XVI-XXIIIT, 1887-94. 8°. Bulletin. No. I, V—VIII, IX. 3, X, 1889-94. 8°. Geology of Minnesota. Final report. Vol. III. 1, 1885-92. 40, Mr, Hamittron.—Lick Observatory. Contributions. No. IV, V, 1895. 8°. New Yorxk.—Academy of Sciences. Annals. Vol. VIII. 6-12, IX, X, XI, 1895-98. $e. Transactions. Vol. X, XIV-XVI, 1594-97. 8°. Memoirs. Vol. I, 1, 1895. 4°. - — American Geographical Society.
Bulletin. Vol. X XVII. 2-4, XXVIDIJ-XXX, XXXI. 1-3, 1895-99.
American Museum of Natural History. Bulletin. Vol. VII-X, XI. 1, 1895-98. 8°. Annual report. 1894-98. 89, Memoirs. Vol. I. 2, 3, II. 1-3, 1895-99. 4°. Linnaean Society. Abstract of proceedings. 1894-5, 1895-6, 1896-7, 1897-8. 8°, Scientific Alliance. Annual directory. VII, VIII, 1897-98. 8°. Torrey Botanical Club.
Bulletin. Vol. XXII. 6-12, XXIJI-XXV, XXVI. 1, 1895-99. 8°.
OBERLIN.— Oberlin College. Laboratory bulletin. No. 3-5, 9, 1885-98. Wilson Ornithological Chapter of the Agassiz Association. Wilson bulletin. No. 8-12, 19-27, 1896-99. 8°. PasaDENA.—Academy of Sciences. Publications. Vol. I. 1-3, 1897-98. 8°. PHILADELPHIA.—Academy of Natural Sciences. Journal. Vol. X. 3, 4, XI. 1, 2, 1894-99. 40°. ——American Entomological Society.
Transactions. Vol. XXII. 2-4, XXIJI-XXV, XXVI. 1, 1895-99.
Franklin Institute. Journal. Vol: CXL-CXLVII, CXLVIII. 1-3, 1895-99. 8°. University of Pennsylvania.
8°,
8°.
Vil
Contributions from the zodlogical laboratory. The embryology of Cre-
pidula, by #. G. Conklin. Boston, 1897. 8°. PirrsBurGH.— Carnegie Museum. Publications. No. V. 1899. 8°. PORTLAND, Mu.—Society of Natural History. Proceedings. Vol. II. 3, 4, 1895-97. 8°. PovuGuKEEPsi£.— Vassar Brothers Institute. Transactions. Vol. VII, i894-96. 8°.
e RocHEstTER.— Academy of Science.
Proceedings. Vol. If. 4, III. 1, 1895-96. 8°. Rock Istanp.—Augustana Library. Publications. No.1, 1898. 8°.
viii Additions to the Library.
Sr. Lovurs.—Academy of Science. Transactions. Vol. VII. 4-20, VIII, IX. 1-4, 1895-99. 8°. —— Missouri Botanical Garden. Annual report. VIII-X, 1896-99. 8°. SaLtem.—Zssex Institute. Bulletin. Vol. XXVI. 7-12, XXVII-XXX, 1894-98. 8°. Annual report. 1898-99. 8°. San Francisco.— California Academy of Sciences. Memoirs. Vol. II, 5, 1896. 8°. Occasional papers. V, VI, 1897-99. 8°, Proceedings. Ser. II. Vol. V, VI. Ser. III. Math.-phys., Vol. I. 1-4; Geology, I. 1-6; Botany, I. 1-9; Zodlogy, I. 1-12. 1895-99. 8°. Zoe: a biological journal. Vol. I, If, 1890-92. 8°. California State Mining Bureau. Report. XIII, 1895-96. 8°. Bulletin. No. VII-XII, 1895-96. 8°. Technical Society of the Pacific Coast. Transactions and proceedings. 1895-96. 89. TopeKka.—Kansas Academy of Science. Transactions. Vol. XIV-XVI, 1898-98. 8°. Tufts College. Studies. IV-V, 1895-98. 89°. Urpana.—Zlllinois State Laboratory of Natural History. Bulletin, Vol. LV. 2-5, V. 1-6, 1895-98. 8°. Biennial report. 1897-98. 8°. W ASHINGTON.— Biological Society. Proceedings. Vol. X. pp. 1-83, 109-174, XI, XII. pp. 1-170, 183-190, XIIT. pp. 1-59, 1896-99. 8°. : “ United States Department of Agriculture. Division of entomology. Bulletin. N.S. No. 1, 3-19, 1896-99. Insect life. Vol. VII. 5, and general index (vol. I-VII), 1895-97. 8°. Report of the chief of the weather bureau. 1895-96. 8°. —— United States Geological Survey. Annual report. XIV-XVIII, XIX. 1, 4, 6, 1892-98. 8°. Bulletin. No. 112-156, 1893-98. 8°, Geological atlas of the United States. Fol. 1-48, 1894-98. we Monographs. Vol. XXITI-XXXJ, XXXV, 1894-98. 4°, United States National Museum. Annual report. 1894-96. S$°. Bulletin. No. XXXII, XXXIV, XLVII. 1-3, X LIX, 1887-96. 8°. Proceedings. Vol. XVIII-XX. 1895-98. S°.
Oceanic .ichthyology. By George Brown Goode and Tarleton H. Bean.
1895. 4°. —— United States Naval Observatory. Astronomical and meteorological observations. 1890. 4°. Astronomical papers prepared for the use of the American Ephemeris and Nautical Almanac. Vol. VIII. 2, 1898. 4°. Report of the superintendent. 1898. 8°. Smithsonian Institution, Bureau of Ethnology. Annual report. XIII-XVI, 1891-95. 8°. WILKES-BARRE.— Wyoming Historical and Geological Society. Proceedings and collections. Vol. IV. 1,1398. 8°. WorcESTER.—American Antiquarian Society. Proceedings. New series. Vol. X-XII, 1895-99. 89,
Additions to the Library. ix
Amiens.—Société Linnéenne du Nord dela France. Bulletin. No. 259-292, 1894-96. 8°. Mémoires. Tome VIII, IX, 1889-98. 8°. AMSTERDAM.—Kon. Akademie van Wetenschappen. Jaarboek. 1894-97. 8°. Verhandelingen. Afdeel. Natuurkunde. Sectie I. Deel V-VI. 1-5. Sectie II. Deel IV, V, VI. 1-2. 1894-98. 6°. Verslagen van de zittingen (gewone vergaderingen) van de wis- en natuur- kundige afdeeling. Deel IV—VI, 1894-98. 89°, ‘Kon. Zodlogisch Genootschap ‘* Natura Artis Magistra.” [Feestschrift] 1838-1888. 4°. AvuGSBURG.—Naturhistorischer Verein fiir Schwaben und Neuburg. Bericht. XXXII, XXXIII, 1896-98. 8°. Australasian Association for the Advancement of Science. Report. 6th meeting, Brisbane, 1895; 7th meeting, Sydney, 1898. 8°. AUXERRE.—Société des Sciences Historiques et Naturelles del Yonne. Bulletin. Tome XLVIII. 2, XLIX-LI, LIT. 1, 1894-98. 8°. BaseEL.—WNaturforschende Gesellschaft. Verhandlungen. Theil XI, XII. 1, 1895-98. 89°. Batavia.—Kon. Natuurkundige Vereeniging in Nederlandsch-Indié. Natuurkundige tijdschrift. Deel LIV-LVIII, 1895-98. s°. Supplement catalogus (1883-1895) der bibliotheek. 1895. 8°. Boekwerken ter tafel gebracht, 1896, 1897. 8°. Magnetical and Meteorological Observatory. Observations. Vol. XVII-XX, 1894-97. 4°. Wind and weather currents, tides and tidal streams in the East Indian Archipelago. By J. P. Vander Stok. Batavia, 1897. f°. BERGEN.— Wuseum. Aarbog. 1893-98. 8°. Publication. V, 1894. 4°. Account of the Crustacea of Norway. By G.O.Sars. Vol. II. Isopoda. 1896-99. 89. - BeRLIN.—Konigliche Sternwarte. Berliner astronomisches Jahrbuch. 1895-99. 89. Veroffentlichungen des kon. astronom. Rechen-Instituts. No. 4-10,
1897-99. 8°. Naturae novitates. Jahrg. XVII. 6-24, XVIII-XX, XXI. 1-14, 1895-99. So.
Botoena.—R. Accademia delle Scienze dell’ Istituto di Bologna. Rendiconto. Anno 1894-95, 1895-96. N.S. Vol. 1, 1896-97. 8°. BomBay.— Bombay Branch of the Royal Asiatic Society. Journal. No. LI-LIYV, 1895-98. 8°. —— Government Observatory. Magnetical and meteorological observations. 1894, 1896. 4°, Bonn.—WNaturhistorischer Verein der preussischen Rheinlande, Westfalens wid des Reg.—Bezirks Osnabriick. Verhandlungen. Jahrg. LI. 2, LIII-LV, 1894-98. 8°. Sitzungsberichte der niederrheinischen Gesellschaft fiir Natur- und Heil- kunde. 1895-98. 8°. BorpEaux.—Académie Nationale des Sciences, Belles-Lettres et Arts. Actes. Année LV-LVII, 1893-95. 8°. Cartulaire de l’église collégiale Saint-Seurin de Bordeaux, publié avec une introduction et des tables par Jean-Auguste Brutails. 1897, 8°. Société Linnéenne. Actes. Tome XLVII-LII, 1894-97. 8°.
x Additions to the Library.
BorDEAUX.—Société des Sciences Physiques et Ni uturelles.
Mémoires. 4¢sér. Tome V. 5¢sér. Tome I-IV. 1895-98. 8°. Proeés-verbaux. Année 1894-5, 1895-6, 1896-7, 1897-8. 8°.
BrEeMEN.—WNaturwissenschaftlicher Verein.
Abhandlungen. Bd. XIV, XV. 2, XVI. 1, 1895-98. 8°.
Meteorologische Station.
Ergebnisse der meteorologischen Beobachtungen. Jahrg. VI-IX, 1895- 98. 4°. BRESLAU.-—Sehlesische Gesellschaft fiir vaterldndische Cultur. Jahres-Bericht. LXXII-LXXYV, 1894-97. 8°. BrisBaNne.— Queensland Branch of the Royal Geographical Society of Australia. Proceedings and transactions. Vol. X-XIII, 1894-98. 8°. Queensland Musewn. Annals. No. IV, 1897. 8°. Briinn.—Naturforscher Verein. Verhandlungen, Bd. XXXITI-XXXY, 1894-97. 8°. Bericht der meteorologischen Commission. XIII-XYV, 1893-95. $89. BRUXELLES.—Académie Roy yale des Sciences, des Lettres et des Beawx-Arts de Pe Mémoires. Tome L. 2, LI- LIII, 1893-98. 4°. Mémoires couronnés et mémoires des savants étrangers. Tome LII- LVI, 1894-98. +0°. Mémoires couronnés et autres mémoires. Tome XLVII-LY, LVII, 1892- OStaoe. Bulletins. 3° sér. Tome XXIV-XXXVI, 1893-98. Tables générales, tome J-XXX._ 8°. Annuaire. Année LX-LXV, 1894-99. 8°. Notices biographiques et bibliographiques. 4° éd. 1896. 8°. Réglements et documents concernant les trois classes. 1896. 8°. Sociéte Belge de Géologie, de Puléontologie et a Hydrologie. Bulletin. Année VII-IX, X. 1, XI. 1, 1893-97. 5°. Société Hntomologique de Belgique. Annales. Tome XX XIX-XLIT, 1895-98. 8°. Mémoires. III-VI, 1895-97. 8°.
Société Royale Belge de Géographie.
Bulletin. Année XIX-XXII, XXIII. 2, 1895-99. 8°. :
Socicté Royale de Botanique.
Bulletin. Tome XXXII-XXXYI, 1893-98. 8°.
Société Royale Malacologique de Belgique.
Annales. Tome XXVII-XXX, XXXI. 1, 1892-95. 8°.
Bucuirest.—ZJnstitut Wétéorologique de Roumanie.
Annales. Tome IX-XIII, 1898-97. 40°.
Bupapast.—Kon. wag. Central-Anstalt fiir Meteorologie wud Erdmagnetismus. Jahrbiicher, Jahre. XXII, XXIV-XXVII, XXVIII. 2, 1892-98. 4°. Publicationen, Bd. I, 1898. 4°.
BuENos ArrEs.—Sociedad Cientifica Argentina,
Aniles. Tomo XXXIX=XLI, XLII 1, 2,5, 6, XLIIL, XLIV, XV. 1-4 XLVI, XLVII, XLYVIII. 1, 2, 1895-99. 89. ' —— Museo Nacional. Anales.” Tomo IV-VI, 1895-99. 8°. Comunicaciones. Tomo I. 1-3, 1898-99. 8°. Memoria. 1894-1896. 8°.
CABEN.—Société Linnéenne de Normandie.
Bulletin. 4° sér, Vol. IX, X. 5e sér, Vol. I. 1895-97. 8°.
CALCUTTA.—Asiaiie Society of Bengal.
Journal. Vol. LXIV. pt. ii, no. 2-4, LXV-LXVII, 1895-98. Bee Proceedings. 1895, no. 4— 10, 1896-98, 1899, no. 1-3, 8°, Annual address, 5 Feb. 1896. 8°,
Additions to the Library. xi
CaLcutrTa.— Geological Survey of India, 2 Paleontologia Indica. Ser. XIII, vol. Il, pt. 1; XV. vol. I, pt. 3,4; vol. pid, 2) 43) OV. vole pts leyaen 890-7. 0. Memoirs. Vol. XXV-XXVII, XXVIII. 1, 1895-98. 8°. i Records. Vol. XXVIII. 3, 4, XXIX, XXX, 1895-97. 8°. ——WMeteorological Department of the Government of India. Indian meteorological memoirs. Vol. V. 7-9, VI. 2,4, VII. 3, 7, VIII. 1, 2, TX. 1-9, X. 1, 2, 1894-99. f°. Monthly weather review. 1895-98, 1899 Jan.—Apr. f°. Rainfall of India. 1894-97. f°. ~CAMBRIDGE.— Philosophical Society. = Transactions. Vol. XVI, XVII. 1-3, 1896-99. 4°. Proceedings. Vol. IX, X. 1, 2, 1895-965. 8°. CaTANIA.—Accademia Gioenia di Scienze Naturali. Att. Ser, TV. Vol, VII-XI, 1894-98. 4°. Bullettino mensile. Nuovaserie. Fasc. 36-56, 59, 1894-98. 89. a CHEMNITZ. —Natur EEE aie Gesellschaft. ag Bericht. XIII, 1882- ier, e CHERBOURG. — Société aeaier" a Sciences Naturelles. 7 Mémoires. Tome XXIX, XXX, 1892-97. 6°. ; 4 CHRISTIANIA.—Kong. Norske Chinen sitet. Norrénaskaller. Crania antiqua in parte orientali Norvegiz meridionalis inventa. Af Justus Barth. 1896. 8°. Fauna Norvegie. Bd.I. Phyllocarida og Phyllopoda. Ved G. O. Sars. 1896. 4°. : Norwegische Commission der Europdischen Gradmessung. Astronomische Beobachtungen und Vergleichung der astronomischen und geoditischen Resultate. 1895, 4°. Resultate der im Sommer 1894 in dem siidlichsten Theile Norwegens ausgefiihrten Pendelbeobachtungen, 1895. 8°. — ——WNorwegisches meteorologisches Institut. eke Jahrbuch. 1893-97. 4°. —— Norwegian North-Atlantic Expedition, 1876-78, ay Publication XXIII, XXIV, 1896-97. 40°.
Any
_ Forhandlinger. 1895-98. 8°. ~ Oversigt. 1894-1898. 8°. Skrifter. 1894, i, ii. 8°. un.—Naturforschende Gesellschaft Grawbiindens. _ Jahresbericht. Neue Folge. Jahre. XXXVIII-XL, 1894-97. 8°. 90BA.— Academia Nacional de Ciencias. - Boletin. Tomo XIV. 2-4, XV. 1-3, XVI. 1, 1894-99. 8°. G.—WNaturforschende Gesellschaft. Schriften. Neue Folge. Bd. IX, Hes 98... 8°.
aan sér. Lome ravi 1905- 98, 8°.
Pe etciochericlite. 1895. 80, Verhandlungen. Bd. XVI. 4, XVII, XVIII, 1896-98. 8°. Naturforscher- Gesellschaft bei der Universitat Dorpat. _ Archiv fiir die Naturkunde Liy-, Ehst- und Kurlands. Ser. II. Bd. XI. 1, 2, 1895-97. S°. Sitzungsberichte. Bd. X. 3, XI, XII. 1, 1894-99. 8°. ‘Schriften. VIII, 1X, 1895-96. 8°. Archiologische Karte yon Liy-, Est- und Kurland, entworfen von J. Sitzka. 1896. 8°.
xii Additions to the Library.
Drespen.—Naturwissenschaftliche Gesellschaft Isis. Sitzungsberichte und Abhandlungen. 1895-i898. 8°. Verein fiir Erdkunde. Jahresbericht. XXIV-XXVI, 1894-98. 8°. Wissenschaftliche Veroffentlichungen. Bd. III. 2, 1897. 8°. Literatur der Landes- und Volkskunde des Kénigreichs Sachsen, hrsg. von Paul Emil Richter. Nachtrag II. 1894, 8°. Dusiin.—Royal Lrish Academy. Transactions. Vol. XXX. 15-20, XX XI. 1-7, 1892-99. . 4°. Proceedings. Ser. III. Vol, II. 4-5, IV, V. 1-2, 1895-97. 89°. Todd lecture series. Vol. VI, 1896. 8°. List of members. 1895, 1896, 1898. EpinpurGH.—Botanical Society. Transactions and proceedings. Vol. XX. 2, 3, XXI. 1-3, 1895-99. 8°. Geological Society. Transactions. Vol. VII. 2-4, 1895-99. 8°. Royal Physical Society. Proceedings. Vol. XIII, XIV. 1, 1894-98. 8°. Royal Society. Proceedings. Vol. XX, X XI, 1895-97. 8°. Emprn.—Naturforschende Gesellschaft. Jahresbericht. LXXIX-LXXXII, 1893-97. 8°. Klein Schriften. XIX, 1899. 8°. ErFurt.—Kon. Akademie gemeinniitziger Wissenschaften. Jahrbiicher. Neue Folge. Heft XXII-X XIV, 1896-98. 8°. FIRENZE.— Biblioteca Nazionale Centrale. Bollettino delle pubblicazioni Italiane ricevute per diritto di stampa. No. 227-828, 1895-99. 8°. —— R. Istituto di Studi Superiori Pratici e di Perfezionamento, Pubblicazioni. Sezione di filosofia e filologia. Le opere latine di Giordano Bruno esposte e confrontate con le italiane. Da Felice Tocco. 1889. 8°. La filosofia dell’ inconsciente metafisica e morale. Per Adolfo Faggi. 1890. 89°. Notizie storico-biografiche intorno al Conte Baldassare Castiglione. Studio del Dott. Camillo Martinati. 1890. 8°. — Sezione di scienze fisiche e naturali. Fisiologia del digiuno, Studisull’ uomo per Luigi Lanciani. 1889, 8°. Le pieghe delle Alpi Apuane. Contribuzione agli studi sull’ origine delle montagne per Carlo de Stefani. 1889. 8°. Sopra i resti di un coccodrillo scoperto nelle lignite mioeceniche de Montebamboli. Nota paleontologica del Dott. Giuseppe Ristori. 18905 18°: ; Sull’ origine e decorso dei peduncoli cerebellari. Pel Dott. Vittorio Marchi. 1891. 8°. — Sezione di medicina e chirurgia. Archivio della scuola d@’anatomia normale e patologica. Vol. V, 1, %, - 1889-90. 8°. Il triennio 1883-85 nella clinica ostetrica e ginecologica di Firenze. Parte I. 1888. 8°: L’acido carbonico dell’ aria e del suolo di Firenze. Indagine siste- matiche eseguite nel 1886 del Dott. Giorgio Roster, 1889. . 8°. Sul lichen rosso. Studio del Dott. Alfonso Minuti. 1891. 8°. FRANKFURT A. M.—Deutsche malakozoologische Gesellschaft. Nachrichtsblatt., Jahre. XXVII. 7-12, XXVIII. 1, 3-12, XXIX. 3-12, XXX. 1-4, 7, 8, 11, 12, XXXI. 1-8, 1895-99. 8°.
Additions to the Library. xiii
FRANKFURT A. M.—<Senckenbergische naturforschende Gesellschaft.
Abhandlungen. Bd. XIX, XX. 1, XXII-XXIV, 1895-98. 4°.
Bericht. 1895-95. 8°.
Katalog der Reptilien-Sammlung. Theil II. 1898. 8°.
FRANKFURT A. O.—WNaturwissenschaftlicher Verein des Regierungsbezirks Frankfurt. Helios. Abhandlungen und monatliche Mittheilungen. Jahrg. XIU-XVI, 1895-99. 8°. Societatum litterae. Jahrg. IX. 4-12, X-XII, 1895-95. 8°. FREIBURG IN B.—WNaturforschende Gesellschaft. Berichte. Bd. 1X, X, XI. 1, 1894-59. 8°. GENEVE.—Jnstitut National Genevois. Bulletin. Tome XXXIIIJ, XXXIV, 1895-97. 8°. Société de Physique et d’ Histoire Naturelle. Mémoires. Tome XXXII, X XXIII. 1, 1894-98. 4°. GENoVA.— Museo Civico di Storia Naturale. Annali. Vol. XXXIV-XXXVIII, 1895-98. 8°. GiEssEN.— Oberhessische Gesellschaft fiir Natur- und Heilkunde. Bericht. XXX, XXXI, 1895-96. 89°. Guascow.—WNatural History Society. , Proceedings and transactions. N.S. Vol. IV. 2,3, V.1, 2. 1894-98. 8°. Philosophical Society. Proceedings. Vol. XXVI-XXIX, 1894-98. 8°. GorxLITz.— Naturforschende Gesellschaft. ; Abhandlungen. Bd. XXI, XXII, 1895-98. 8°. GOTEBORG.—Kon. Vetenskaps och Vitterhets Samhdille.
Handlingar. Ny tids. Haft. XXX-XXXII. 4defolj. Haft. I. 1895-
Sisk ties GOTTINGEN.—Konigl. Gesellschaft der Wissenschaften.
Nachrichten. Philologisch-histor. Klasse, 1894, ivy, 1895-98, 1899, i; Mathe- matisch-physikal. Klasse, 1895, ii-iv, 1896-98, 1899, i; Geschaftliche Mittheilungen, 1895, ii, 1SY6-98, 1899, i. 8°.
Gutstrow.— Verein der Freunde dev Nuturgeschichte in Mecklenburg. Arehiv. Jahrg. XLIX-LII, LIII. 1, 1895-99. $9. HapBana.—Academia de Ciencias Médicas, Fisicas y Naturales. Anales. No. 418, 1899. 8°. Real Colegio de Belen. Obseryaciones magneticas y meteorologicas. 1891-97. 4°. Investigaciones relativas a la circulacion y traslacion ciclonica en los huracanes de las Antillas. Por el P. Benito Vines. 1895. 5°. Hawiax.—Nova Scotian Institute of Natural Science. Proceedings and transactions. Vol. VIII. 4, IX, 1893-98. 89°, —— Department of Mines, Nova Scotia. Report. 1896, 1897, 1898. 89. Ores of Nova Scotia. Gold, lead and copper. By E. Galpin. 1898. &°. Hatie.—Kais. Leopoldinisch-Carolinische deutsche Akademie der Naturforscher. Leopoldina. Heft XXX-XXXIV, 1894-98. 4°. Naturforschende Gesellschaft. Abhandlungen. Bd. XXI, 1898-99. 5°. Naturwissenschaftlicher Verein fiir Sachsen und Thiiringen. Zeitschrift fiir Naturwissenschaften. Bd. LXVIII. 1,2,5,6, LXIX, LXX, LXXI. 1-5, 1895-99. 8°. HampBure.—Deutsche Scewarte.
Aus dem Archiv. Jahrg. XVIII-XXI, 1895-98, 4°.
Deutsches meteorologisches Jahrbuch, 1894-97. 4°.
Ergebnisse der meteorol. Beobachtungen, 1891-95, 1886-95. 4°.
Xiv Additions to the Library.
Hampure.—Naturwissenschaftlicher Verein. Abhandlungen. Bd. XIV, XV, 1896-97. 8°. Verhandlungen. III. Folge. I-VI, 1893-98. 8°. Hannover.—Naturhistorische Gesellschaft. Festschrift zur Feier des 100jahrigen Bestehens der Naturhist. Gesell- schaft. Geschichte und 44.-47. Jahresbericht. 1897. 89. Katalog der systematischen Vogelsammlung des Provincial-Museums in Hannover. 1897. 8°. ; Katalog der Vogelsammlung aus der Provinz Hannover. 1897. 8°. Verzeichniss der im Provincial-Museum vorhandenen Saugethiere. 1897. 8°. Verzeichniss der in der Provinz Hannover vorkommenden Gefiasspflanzen nebst Angabe ihrer Standorte. Zusammengestellt von W. Brandes. 1897. 8°: Hartem.—Musée Teyler. Archives. Série II. Vol. IV. 4, V, VI, 1895-99. 8°. Société -Hollandaise des Sciences. Archives néerlandaises des sciences exactes et naturelles. Tome XXIX, 2-5, XXX. 2% sér, Tome I, If, 1895-99) “89: Le Havre.—Sociélé Géologique de Normandie. Bulletin. Tome XVI, XVII, 1892-95. 89. HELSINGFORS.—Societas Scientiarum Fennica. Acta. Tom. XX-X XIII, 1895-97. 4°. Ofversigt af forhandlingar. XXXVI-XXXIX, 1893-97. 8°. Bidrag till kinnedom af finlands natur och folk. Haft. LIV-LVI, 1894-5. 8°. Institut Météorologique Central, ; Observations météorologiques (stations finlandaises). 1889-90; tome sup- plémentaire, 1881-90; résuiné, 1891-90. HERMANNSTADT.— Siebenbiirgischer Verein fiir Naturwissenschaften. Verhandlungen und Mittheilungen. Jahre. XLIV-XLVI, 1894-96. 8°. Der siebenbiirg. Verein nach seiner Entstehung, seiner Entwicklung und seinem Bestande. 1896. 8°. JENA.— Medicinisch-naturwissenschaftliche Gesellschaft. Jenaische Zeitschrift fiir Naturwissenschaft, Bd. XXIX. 3.4, XXX- XXXII, XXXIIL. 1, 1895-99. 8°, Kasan.—Sociélé Physico-Mathématique de V Université Impériale. Bulletin. 2¢sér. Tome IV. 3, 4, V-VII, VIII. 1-3, 1895-98. 8°. KuARKOW.—Société des Sciences Physico-Chimiques. Travaux. Tome I, II, 1894-95 8°. Société de. Médecine Scientifique et @ Hygiene. Travaux. 1896, 1897. 89°. Vingt-cinquiéme anniversaire. : 1898. 8°. Kiew.—Konigl. Christian Albrechts- Universitdt. Schriften aus dem Jahre 1894-95, 1895-96, 1896-97, 1897-98. 8°. —— Naturwissenschaftlicher Verein fiir Schleswig-Holstein. Schriften. Bd. X. 2, XI. i, 1895-97. 8°. : Kirv.—Kievskie Obshchestuo Testestvoispytatelei. Zapiski. Tom. XIII-XV, 1894-98. 8°. KINGSTON, — Ji stitute of Jamaica. Annual report for year ended 31 March, 1896. f°. KJOBENHAVN.—Kon. Danske Videnskabernes Selskab. Oversigt over forhandlinger, 1894. iii, 1895-98, 1899. i-iii. S°. —— _ Naturhistorisk Forening.
Videnskabelige meddelser. Aaret 1895-98, 8°.
Additions to the Library. XV
K6ONIGSBERG.—Konigl, physikalisch-dkonomische Gesellschaft. Schriften. Jahrg. XXXV-XXXIX, 1894-98. 4°. Krakow.—XK. k. Sternwarte. Materyaly do klimatografii Galicyi. Rok 1894-97. 8°. Stan wody narzekach Galicyjskich. Rok 1893-94. $°. La Puata.—Direccion General de Estadistica de la Provincia de Buenos Aires. Anuario estadistico. Ano 1896. 4°. Memoria demografica, Ano 1895, 4°. L’agriculture, l’élevage, Vindustrie et le commerce en 1895. 4°. Museo. Revista. Tomo Y-VIII, 1894-98. 8°. Anales. Seecion de antropologica. I, II, 1896-97. f°. Seccion zoologica. II, II, 1895-99. f°. Paleontologia Argentina. III, IV, 1895-96. f°. LAUSANNE.—Société Vaudoise des Sciences Naturelles. Bulletin. 3¢sér. No. 116-131, 1894-99. 8°. LrEEDs.— Yorkshire Geological and Polytechnic Society. * Proceedings. Newseries. Vol. XII. 5, XIII. 2, 1894-97. 89, LewwEn.—WNederlandsche Dierkundige Vereeniging. Tijdschrift. Ser. II. Deel V, VI. 1, 1896-98. 8°. Catalogus der bibliotheek. 4deuitg. 1897. 8°. Compte-rendu des séances du 8¢ congrés international de zoologie, Leyde, 16-21 Sept., 1895. Leyde, 1895. 8°. Sternwarte. Annalen. Bd. VII, 1897. 4°. Verslag. 1894-96. 5°. Lerezie.— Konigl. stichsische Gesellschaft der Wissenschaften. Berichte. Math.-physische Classe. Bd. XLVII. 2-6, XLVIII, XLIX, L, math. 3-5, naturwiss., LI, math. 1-4, 1895-99. 8°. Namen-und Sachregister der Abhandlungen und Berichte der math.- phys. Classe, 1846-1895. 4°. Zur fiinfzigjahrigen Jubelfeier der kon. sachs. Gesellschaft am 1 Juli, | 1896. 8°. 4 Naturforschende Gesellschaft. . Sitzungsberichte. Jahrg. XIX-XXI, 1892-94. 8°. Verein fiir Erdkunde. Mittheilungen. 1895-98. 8°. Wissenschaftliche VerOffentlichungen. Bd. III, 1896-99. 8°. Zoologischer Anzeiger. No. 478-596, 1895-99. 8°. LisBoa.— Sociedade de Geographia. Boletim Serie XIII. 1, 2, 10, 11, XIV, XV, XVI. 1-11, 1894-97. 8°. Os descobrimentos Portuguezes e os de Colombo. Por Manuel Pinheiro Chagas. 1892. 8°. _ Lonpon.—Geological Society. , Quarterly journal. Vol. L13, 4, LIL. 1-3, LID, LIV. 1,4, LV. 1-3. Gen- . eral index, vol. I-L. 1895-99. 8°. Linnean Society. Journal. Zoology. No. 158-175, 1894-99. 89. Journal. Botany. No. 209-238, 1894-99. 8°. Proceedings. Noy. 1893-June 1897. 8°. List. 1894-95—1898-99. 8°. Mathematical Society. Proceedings. No. 509-672, 1895-99. 8°. Royal Historical Society. Transactions. Newseries. Vol. [IX-XI, 1885-97. 8°. The domesday of inclosures, 1517-1518. London, 1897. 2y. 8°.
;
xvi Additions to the Library.
Lonpon.—Royal Microscopical Society. Journal. 1895, iv-vi, 1896-98, 1899, i-iv. 8°. Royal Society. ; Philosophical transactions. Vol. CLXXXV, A, B; CLXXXVI, A, B; CLXXXVII, A, B; CLXXXVIII, A, B; CLXXXTX, A. 1894-97740. Proceedings. No. 346-396, 398-418, 1894-1899. 8°. List of council and fellows. 1894-97. 4°. Year-book. I, II, 1897-98. 8°. Record. No.1, 1897. 8°. LovuvaIn.—La Cellule: Tome XI-XV, XVI. 1, 1895-99. 89. LunpD.— Universitet. Acta. Tom, XXXI-XXXIV, 1895-98. 4°. Festskrift med anledning af hans Majestat Konung Oscar II* regerings jubileum, 1872-97. Afdel. III. 1897. 4°. LuxemBourG.—Institut Royal, Grand-Ducal. Publications. Section des sciences naturelles et mathématiques. Tome MXIV, 1896. (8°. Lyon.—Académie des Sciences, Belles-Lettres et Arts. Mémoires. Sciences et lettres. 3° sér. Tome III-V, 1895-98. $°. Mapras.— Government Observatory. ‘ Daily meteorological means. 1896. 4°, Report for 1897-98. 8°. i Maprip.—Comision del Mapa Geologico de Espana. Boletin. Tomo XX-XXIV, 1893-97. Indice Tom. I-XX. 8°. Memorias. Descripcion fisica y geologica de la provaniia de Logrono. Por D. Rafael Sanchez Lozano. 1894. 8°. Explicacion del mapa geologico de Espana. Por L. Mallada. Tomo I- III, 1895-98. &°. —— Observatorio. Observaciones meteorologicas. 1894-95, 1896-97, 8°. Resumen de las observaciones meteorologicas efectuadas en la peninsula. 1891-92, 1893-94, 1895-96. 8°. Real Academia de Ciencias Bractas Pisicas y Naturales. Anuario. 1896-98. 16°. Memorias. Tomo XVI, 1895. 8°. . MAncCHESTER.—Literary and Philosophical Society. Memoirs and proceedings. Series IV. Vol. VIII. 4, IX, X, 1894-96. Vol. XL-XLII, XLIII. 1-3. 1897-99. §9. Marsure.— Gesellschaft zur Befirderung der gesammten Natuwrwissenschafien. Sitzungsberichte. Jahre. 1894-97. 8°. MARSEILLE.—Faculté des Sciences. Annales. Tome VY, VI. 1-3, VII, 1896. 4°. Metz.—Académie. : Mémoires. 8° sér. Année XXII-XXV, 1892-96. 8°. Mex1co.—Asociacion de Ingenieros y Arquitectos. Anales. Tomo IV. 4-13, V-VII, 1895-98. 8°. Instituto Geolégico de México. Boletin. No. I-XI, 1895-98. 4°. Expedicion cientifica al Popocatepetl. 1895. 8°, — Instituto Médico Nacional. Anales. Tomo I, 8, II. IIT. 1-11, 14-22, IV. 1, 1896-99. 4°. ——- Observatorio Me teoroldgune: Magen Central.
Boletin mensuel. 1895, no. 4-12; 1896, no.1-3, 6-12; 1897; 1898: 1899, no. 1-4, 40°, — Secretario de Fomento.
Biblioteca Botanico-Mexicana. Por Dr. Nicolas Leén, 1895. 8°.
oe cay cee
Additions to the Library. XVli
Mexico.—Sociedad Cientifica “‘Antonio Alzate.” Memorias y revista. Tomo VIUII-XI, XII. 1-8, 7-10, 1894-99. 8°. Sociedad de Geographia y Estadistica. Boletin. EpocalV¥. Tomo III. 3-9, 1894-95. 8°. Sociedad Mexicana de Historia Natural. La naturaleza. Ser. II. Tomo II. 8-11, 1894-96. 4°. MIDDELBURG.—Zeeuwsch Genootschap der Wetenschappen. Geschiedskundige beschrijving van Tholen en omstreken. Door A, Hollestette. 1897. 8°. Zelandia illustrata. 2* vervolg. 1897. 8°. Mizano.—Real Istituto Lombardo di Scienze e Lettere. Rendiconto. Serie II. Vol. XX VII-XXXI, 1894-98. 8°. Reale Osservatorio di Brera. Riassunto delle osservazione meteorologiche. 1895-98, 4°. Societa Italiana di Scienze Naturali. Atti. Vol. Il, 1V—XII, XX. 1,2, XXI. 1, 2, XXIII. 1, 2, XXXV, XXXVI, XXXVII. 1, 2, 1881-98. . 8°. Memorie. Tomo VI. 1, 1897. 4°. Mopena.—Regia Accademia delle Scienze, Lettere ed Arti. Memorie. Serie II. Tomo X, XI, XII. 1, 1894-96. 40°. Societa dei Naturalist. Memorie. Serie III. Vol. XII. 3, XIII-XV, XVI. 1, 2, 1894-98. 89°. Mont Buianc.— Observatoire Meétéorologique. Annales. Tome II, III, 1896-98. 8°. MOontTEVIDEO.— Museo Nacional. Anales. Fasc. VII-XI, 1896-99. 4°. MONTPELLIER.—Académie des Sciences et Lettres. Mémoires. Section des lettres. Sér. IJ. Tome I. 5-7, II. 1, 1895-97. 8°. Section dessciences. Sér. II. Tome II. 2-4, 1895-96. 8°. Moscovu.—Société Impériale des Naturalistes. Bulletin. Année 1894. ivy, 1895-97, 1898, i. 8°. MUtncHen.— Kon. bayerische Akademie der Wissenschaften. Sitzuugsberichte. Philosph.-philolog. und histor. Classe. 1894. ii-iii, 1895-97, 1898, Bd. I. 89. Mathemat.-physikal. Classe. 1894. iv, 1895-97, 1898. i-iii. 8°. Ueber die Bedeutung wissenschaftlicher Ballonfahrten. Festrede von L. Sohnke. 1894, 4°. Ranke und Sybel in ihrem Verhialtniss zu Konig Max. Festrede von Alfred Dove. 1895. 4°. Der churbayerische Kanzler Alois Freiherr yon Kreittmayr. Festrede von August von Bechmann. 1896. 4°. Ueber die wechselseitigen Beziehungen zwischen der reinen und der angewandten Mathematik. Festrede von Walther Dyck. 1897. 4°. Die Bedeutung der deutschen Philologie fiir das Leben der Gegenwart. Festrede von Hermann Paul. 1897. 4°. Der bayerische Geschichtsschreiber Karl Meichelbeck, 1669-1734. Fest- rede von Franz Ludwig Baumann. i897. 4°. Gedachtnissrede auf Ludwig von Seidel, von Ferdinand Lindermann. 1898. 4°. Ueber Studium und Auffassung der Anpassungserscheinungen. Festrede von Karl Goebel. 1898, 4°. —Konigliche Sternwarte im Bogenhausen. Neue Annalen. Bd. III, 1898. 4°. Minster.— Westfilischer Provincial- Verein fiir Wissenschaft und Kunst. Jahresbericht. XXXIII-XXXYI1, 1894-98. 58°.
”
Xvill Additions to the Library.
Nanoy.—Académie de Stanislas. Mémoires. 5¢sér. Tome XII-XV, 1894-97. 8°. Napour.—R. Accademia delle Scienze Fisiche e Matematiche. Atti. Ser. II.’ Vol. VII, VIII, 1895-96. 4°. Rendiconto. Ser. III. Vol. I-IV, V. 1-7, 1895-99. 4°. —— Real Istituto @ Incoraggiamento alle Scienze Naturali, ete. Atti. Ser. IV. Vol. VII, VII, X, XI, 1894-98. 4°. NEvucHATEL.—Société des Sciences Naturelles. Bulletin. Tome XXI-XXV, 1893-97. 8°. NewcAstLE-UPON-LTYNE.—JNorth of “England Institute of Mining and Mechanical Engineers. Transactions. Vol. XLIV. 4,5, XLV-XLVII, XLVIII. 1-4, 1895-99. 8°. Annual report of the council. 1896-97, 1897-98, 1898-99. 8°. Account of the strata of Northumberland and Durham as proyed by bor- ings and sinkings. U-Z. 1897. 8°. Ntrnperc.—WNaturhistorische Gesellschaft, Jahresbericht nebst Abhandlungen. Bd. X. 3-5, XI, 1894-97. 89°. Ovessa.—Société des Naturalistes de la Nouvelle Russie. Zapiski. Tom. XIX-XXI. 1, XXII. i, 1894-98. 40. Matematicheskoe otdielente. Tom. XVII, 1895. 8°. Université Impériale. Annales de l’obsery. magnétique et météorologique. Année LV, 1897. 4°. OsnaBRUCK.—M ee wissenschaftlicher Verein. Bericht. X, XIII, 1893-98. 8°. Snare ie Coney and Natural History Survey of Canada. Annual report. New series. Vol. VI-IX, 1892-96. 8°. Maps: Nova Scotia, 15 sheets; British Columbia, 9 sheets; Quebec, 1 - sheet; Ontario, 1 sheet. Royal Society of Canada. Proceedings and transactions. Vol. XII, 1894. 4°. OxrorD.—Radcliffe Library. Catalogue of books added 1895-98. 8°. ——-adcliffe Observatory. Results of astronomical and meteorological observations. Vol. XLVI, XLVII, 1888-91. 8°. - PALERMO.—&. Accademia di Scienze, Lettere e Belle Arti. Atti. Ser. III. Vol. II-IV, 1893-96. 4°. Pel terzo centénario della morte di Torquato’Tasso. Adunanza del 19 Maggio, 1895. 4°. Paris.—Evole Normale Supérieure. Annales scientifiques. 3° sér. Tome XII. 7-12, XIII-XV, XVI. 1-8, 1895-99, 4° Ecole Polytechnique. Journal, 2° sér, Cahier I-1V, 1895-98. 4°. cana Guimet. Annales. Tome XXYVI. 2, 3, XXVII-XXIX, 1895-97, 4°. Bibliotheque des études. Tome III, V-VII, 1895-98. 8°. Revue de Vhistoire des religions, Tome XXIX. 3, XXX-XXXVII, 1895- 98. 89. Muséum @ Tistoire Naturelle. Bulletin. Année 1895, no. 4-8, 1896-98, 1899, no. 1-5. 8°. Observatoire National. Rapport annuel, 1895-98. 4°, Société Mathématique de France. Bulletin, Tome XXIII. 4-10, XXIV-XXVI, XXVIL. 1, 2, 1895-99. 8°. « Ocuyres mathématiques d’Eyvariste Galois. Paris, 1897. 8°.
e
Additions to the Library. xix
PaRis.—Société Nationale d@ Acclimatation.
Revue des sciences naturelles appliquées. Année XLII. 12-17, XLII. 1
3-12, XLIV, 1895-97. 8°. Société Zoologique de France. Bulletin. Tome XX=X XIII, 1895-98. 8°. Mémoires. Tome VIII-XI, 1895-98. 89°. PENZANCE.—Royal Geological Society of Cornwall, Transactions. Vol. XII. I-4, 1896-99. 8°. Pisa.—Societd Toscana di Scienze Naturali. Memorie. Vol. XIV—-XVI, 1895-98. 8°. Processi verbali. Vol. IX. pp. 243-310, X, XI. pp. 1-157, 1895-99. Se, PorspamM.—Astrophysikalisches Observatorium. Publicationen. Bd. XI, XIII, 1895-99. 4°. Photographische Himmelskarte. Bd. I, 1899. +°. Prac.—Kon. bihmische Gesellschaft der Wissenschaften. Sitzungsberichte der math.-naturwiss. Classe, 1894-98. 8°. Jahresbericht, 1894-98. 8°. — _ K. k. Sternwarte.
b
Magnetische und meteorologische Beobachtungen. Jahrg. LVI-LIX,
‘ 1895-98. 4°. REGENSBURG.— Naturwissenschafilicher Verein. Berichte. Heft V, VI, 1894-98. 8°. Historischer Verein von Oberpfalz und Regensburg. Verhandlungen. Bd. XLVII-L, 1895-98. 8°. Riga.—Naturforscher Verein. Correspondenzblatt. Jahrg. XXXVIII-XLI, 1895-98. 8°.
Festschrift in Anlass seines 50jahrigen Bestehens, am 27 Miirz (8 April),
1895. 8°. Die Bodentemperaturen bei Riga. Bearbeitet von G. Schweder. Riga, 1899. 4°. Rio DE JANEIRO.—Jnstituto Historico Geographico Brazileiro. Revista trimensal. Tomo LVI. 2, LVII, LVIII, 1894-96. 8°. Commission centrale de bibliographie Brésilienne. Année I, 1894. 8°. La ROCHELLE.—Société des Sciences Naturelles de la Charente-Inférieure. Annales. 1894-98. 8°. Roma.—Accademia Pontifica de’ Nuovi Lincei. Atti. Anno LXVII. 4-6, XLVIII-LI, 1894-98. 4°. Reale Accademia dei Lincei.
Atti. Serie V. Rendiconti. Classe di scienze fisiche, matematiche e
naturali. Vol. IV. 8, 10-12, V-VII, VIII. i, ii. 1-3, 1895-99. 4°.
Rendiconto dell’ adunanze solenne. 1895-99. 4°. Reale Comitato Geologico @ Italia. ;
Bollettino. Vol. XXV-XXIX, 1894-98. 8°. Societa degli Spettroscopisti Italiani.
Memorie. Vol. XXIV. 7-12, XXV-XXVII, XXVIII. 1-6, 1895-99. 4°. Societd Italiana delle Scienze.
Memorie. Ser. III. Tome VIII-XI, 1892-98. 4°. RorrerDamM.—Bataafsch Genootschap der Proefondervindelijke Wijsbegeerte.
Nieuwe verhandelingen. Reeks II. Deel IV. 2, 1897; buitengewone
aflevering, 1895. 4°. St. GALLEN.—WNaturwissenschaftliche Gesellschaft.
* Bericht. Jahrg. 1893-96. 8°.
St. Jonn.—WNew Brunswick Natural History Society. Bulletin. No. XIV-XVII, 1886-99. 8°.
ne 2 Additions to the Library.
S. PaoLto.—Commissao Geographica e Geologica de S. Paolo. Boletim. No. 10-14, 1895-97. 8°. Seecao metvorologica. Dados climatologicos. 1893-97. 8°. Museu Paulista. Revista. ‘Vol. J, III, 1895-98. Sr. Pererspure.—Acad. Impériale des Sciences. Bulletin. 5esér. Tome II. 3-5, III-VIII, [X. 1, 1895-98. 4°. Mémoires. 7° sér. Tome XLII, 7-14, 1894-99, 40°. , 8° sér. Classe phys.-math. Tome I-IV. 1, 2, 4-13, VI. 1-10, 1894-97. .
4°,
Versuch eines Worterbuches der Tiirk-Dialecte. Von W. Radloff. Lief. VII-X, 1895-98. 8°.,
Die alttiirkischen Inschriften der Mongolei. Von W. Radloff. Lief. III. u. Neue Folge, 1895-97. 4°.
Atlas der Alterthiimer der Mongolei. Von W, Radloff. Lief. IIT, 1896. ile.
Proben der Volkslitteratur der nordlichen turkischen Stamme, gesam- melt und iibersetzt von W. Radloff Theil VII, 1896. 8°.
Reisen und Forschungen im Amur-Lande, 1854-56, hrsg. von L. von Sehrenck. Bd. III. 38,1875. 4°. :
Syrisch-nestorianische Grabinschriften aus Semirjetschie, Neue Folge, hrsg. von D. Chwolson. 1897.
*Abdulqidiri Bagdadensis Lexicon Sahnamianum. Ed. ©. Saleman. Tom. I. 1, 1895.
Bibliotheca Friedlandiana. Catalogus librorum impressorum hebraeo- rum in Museo Asiatico Acad. Sci. Petropol. asservatorum. Opera et studio Samuelis Wiener. Fase. II, III, 1895-97. 40°. ;
Bibliotheca Buddhica. Cikshamuccaya, a compendium of Buddhistic teaching compiled by Cantideva, Ed. by C. Bendall.- I. 1897. 8°.
Das Manava-Grhya-Sitra, nebst Commentar, brse. von Friedrich Knauer- 1897 (See.
Arkheologicheski dnevnik poiezdki v Sredniuiu Mongoliu vy 1891 godu. D. A. Klements. 1895. 89.
Geografia Tibeta. V. Vasiliev. 1895. 8°.
Snoshenia Petra Velikago s armianskim narodom. G. A. Ezoy. 1898. 8°.
Sbornik trudoy orkhonskof ekspeditsii. III. Kitaiskia nadpisi na orkh- onskikh namiatnikakh. V. P. Vasiliev. IV. Drevne-tiurskie namiat- niki. V. V. Radltov iP. M. Melioranski. 1897. 2yv. 8°.
Istoricheski obzor arkheologicheskikh izsledovani i otkryti na taman- skom poluostrove s kontsa XVIII stoletia do 1859g. K. K. Hertsa. Izd. 2-e. 898s" 8°:
Comité Géologique.
Mémoires.. Vol. IX. 4, X. 3, 4, XIV, XV. 2, XVI. 1, 1895-98. 4°.
Bulletins. Vol. XIII. 4-9, XIV-XVI, XVII. 1-5, 1894-98. $9.
Bibliotheque géologique de la Russie. 1893-96. 8°.
Hortus Petropolitanus. Acta. Tom. XIV, XV. 1, 1895-96. 8°. ——Imp. Russ. Geograf. Obshtchestuo.
Izviestiya. Tom. XXXI-XXXIV, 1895-98. 8°,
Otchet. God 1894-97. 8°,
Beobachtungen der russischen Polarstation an der Lenamiindung. Theil I. Astron. und magnet. Beobachtungen, 1882-84. 4°.
Physikalisches Centralobservatorium,
Annalen. Jahrg. 1894-96. 4°,
Russisch—Kaiserliche Mineralogische Gesellschaft.
Verhandlungen, Ser, Il. Bd, XXXI-XXXV, 1894-98. 8°.
Additions to the Library. xxi
Sr. PeETERSBURG.—Russisch-Kaiserliche Mineralogische Gesellschaft.
Materialien zur Geologie Russlands. Bd. XVII, XVIII, 1895-97. 8°. Systemetisches Sach-und Namenregister zu der zweiten Serie der Ver- handlungen und den Materialien zur Geologie Russlands, 1885-1895, 8°.
Université Impériale, Observatoire Astronomique.
Mesures micromeétiques d’étoiles doubles faites a St. Pétersbourg et a
Domkino. Sér. I, III, 1895-97. 8°. San SALvApDOR.—Observatorio Astronomico y Meteorologico. Anales. 1895. 4°. Observaciones meteorologicas. Enero-Marzo, 1897. SantTiago.—Z/nstituto de Hijiene. Revista Chilena de hijiene. Tomo I-IV, 1894-99. 89. Boletin de hijiene é demografia. AnoI, 1898. 8°. Consejo superior de hijiene. Sesiones 1896, 1897, Société Scientifique du Chili. Actes. Tome IV. 5, V—VII, VIII. 1-4, 1895-98. 89. Congress cientifico jeneral Chileno de 1894. 8°. — Universidad di Chile. Anales. 1898, Nov., Diciem.; 1899, Enero-Abril. 8°. Schweizerische noturforschende Gesellschaft. Verhandlungen. Jahresversammlung LXXVIII, 1895. 8°. STAVANGER.— Museum. Aarsberetning. 1893. 8°, SrockHotm.—Hntomologisk Forening. Entomologisk tidskrift. Arg. XVI-XIX, 1895-98, 8°. —-Kongl. Bibliotek.
Sveriges offentliga bibliotek Stockholm, Upsala, Lund, Gdéteborg.
Accessions-katalog, X-XII, 1894-97. Register, 1886-95. 8°. Kongl. Svenska Vetenskaps-Akademie. Handlingar. Ny foljd. Bd. XXVI-XXX, 1894-98. 4°. Bihang till handlingar. Bd, XX—XXIIT, 1895-98. 8°. Ofversigt af férhandlingar. Bd. L-LIV, 1894-97. 8°. Meteorologiska jakttagelser. Bd. XXXIJJ-XXXIV, 1891-92. 4°.
Carl yon Linné’s brefyexling. Férteckning uppriittad af Ewald Ahriing. g S$ upy
1885. 8°.
C. W. Scheele. Efterlemnade bref och anteckningar utgifna af A. E.
Nordenskidld. 1892. 8°.
Om sveriges zoologiska hafsstation Kristineberg. Af H. Theel. 1895, 8°.
STRASSBURG.—Laiserliche Universtits-Sternwarte. Annalen. Bad. I, 1896. 4°. STUTTGART.— Verein fiir vaterlindische Naturkunde in Wiirttemberg. Jahreshefte. Jahrg. LI-LV, 1895-99. 8°. SyDNEY.— Australian Museum. Records. Vol. III. 1-5, 1897-99. 8°. Report. 1898. f°. Government Observatory.
Results of rain, river and evaporation observations during 1894-95, 1897.
Ser
Nine papers (extracts) by H. C. Russell, 1894-98. Linnean Society of New South Wales.
Proceedings. Series II. Vol. XVI-XXIII, XXIV. 1, 1891-99. 8°. Royal Society of New South Wales.
Journal and proceedings. Vol. XXVIII-XXXI, 1894-97. 8°. TACUBAYA.— Observatorio Astronomico Nacional.
Anuario. Ano XV—XVIII, 1896-98. 8°.
XXii Additions to the Library.
TacuBaya.— Observatorio Astronomico Nacional. Boletin. Tomo I. 22-25, Il. 1-5, 1895-09. 4°. Observaciones meteorologicas. 1895. 4°. THRONDHJEM.—Aon. Norske Videnskabers Selskab. 4 Skrifter. 1893-97. 8°. TirLis.—Physikalisches Observatorium. Beobachtungen. 1893-96. 4°. Beobachtungen der Temperatur des Erdbodens, 1890. 8°. Toxyo.—Jmperial University of Japan. Journal of the college of science. Vol. VIII. 2, IX, X, XI. 1-3, XII. 1-3 1895-97. 4°. Calendar. 1894-5—1897-8. 8°. Torino.—Musei di Zoologia ed Anatomia Comparata. Bollettino. No. 193-353, 1895-99. 8°. Toronto.—Canadian Institute. Transactions. Vol. IV. 2, V, 1895-98. 8°. Proceedings. Newser. Vol.I, If. 1, 1897-99. 8°. TouLousE.-—Académie des Sciences, Inscriptions et Belles-Lettres. Mémoires. 9¢ sér. Tome VII-IX, 1895-97. 8°, Bulletin. Tome I. 1-38, 1897-98. 8°. Faculté des Sciences. Annales. Tome I-XI, XII. 2-4. 2°sér. Tome I. 1887-99. 4°. TRIESTE.— Osservatorio Astronomico-Meteorologico. Rapporto annuale. Vol. X-XII, 1893-95, 4°. Troms0.— Museum. Aarsberetning. 1893-96. 8°. Aarshefter. XVII-XIX, 1895-96, 8°, Upsata.—Kongl. Universitet. Arsskrift. 1894-96. 8°. Bulletin of the geological institution. Vol. II, III. 1, IV. 1, 1894-98. 8°. Observatoire météorologique, Bulletin mensuel. Vol. XXX, 1895. 4°. Zoologiska studier. Festskrift Wilhelm Lilljeborg tillegrad pa attonde fodelsedag af svenska zoologer. 1894. 4°. , Regia Societas Scientiarum. Nova acta. Ser. III. Vol. XV. 2, XVII, XVIII. 1, 1895-99. 4°. ’ Urrecut.—Kon. Nederlandsch Meteorologisch Instituut. Nederlandsch meteorologisch jaarboek. Jahrg. XLV—XLVIII, 1893-6, 4°. Provinciaal Utrechtsch Genootschap van Kunsten en Wetenschappen. Verslag van het verhandelde in de algemeene vergadering. 1895-98. 8°. Aanteekeningen van het verhandelde in de sectie-vergaderingen. 1895-98. 8°. ; : ; VENEZiA.—Istituto Veneto di Scienze, Lettere ed Arti. Atti. Ser. VII. Tomo YV. 4-10, VI-VIII, IX. 1-7, 1894-98. 8°. WELLINGTON.—New Zealand Institute. Transactions and proceedings. Vol. XXVII-XXX, 1893-95. 8°. The students’ flora of New Zealand and the outlying islands. By Thomas Kirk. Wellington, 1899. 4°. WieEn.--Kais. Akademie der Wissenschaften. Sitzungsberichte. Mathemat.-naturwiss. Classe. Abth. I. Bd. CIII. 4 10, CIV-CVI, CVII. 1-5, 1894-98, 8°. ~—— K. k. Central-Anstalt fiir Meteorologie und Erdmagnetismus. Jahrbiicher. Neue Folge. XXX-XXXIII, XXXIV. 1, 1898-97. 4°, K. k. geologische Reichsanstalt. Abhandlungen. Bd. XVII. 4, XVIII. 1, 1895-97. 4°. Jahrbuch. Bd. XLIV. 3, 4, XLV-XLVII, XLVIIL. 1, 2, 1894-98. 8°,
ee ee ee :
——
onsen.
a oe
Additions to the Library. XXIll
Wien.—AXK. k. geologische Reichsanstalt. Verhandlungen. Jahrg. 1895, no. 6-18; 1896, no. 1-12, 16-18; 1897; 1898. \ 8°. _ +— K. k. naturhistorisches Hofmuseum. 3 Annalen. Bd. X-XII, 1895-97. 89°. ——-K. k. Universitdts-Sternwarte. Annalen. Bd. X—-XIII, 1898. 8°, = ——K. k. zoologisch-botanische Gesellschaft. Verhandlungen. Bd. XLY. 5-10, XLVI- —XLVIII, XLIX. 1-7, 1895-99. 8¢ WIESBADEN.—Wassauischer Verein fiir Naturkunde. Jahrbucher. Jahrg. XLVIII-LI, 1895-98. $09. Wiirzeure.—Physikalisch-medicinische Gesellschaft. | }: Sitzungsberichte. Jahrg. 1895-98. 8°, Lis
trice. —Naturforschende Gesellschaft. Vierteljahrschrift. Jahrg. XL. 24, XLI-XLIII, XLIV. 1, 2, 1895-99. 8°.
_. Bessey, Charles E. The phylogeny and taxonomy of angiosperms. 1897. 8°. a From the Botanical Society of America. Chamberlin, T. C. A group of hypotheses bearing on climatic changes. Chicago, 1897 18°. From the Author. Cobb, N. A. Agricultural experiment work. Sydney, 1897. 8°. From the Author. Coulter, John M. The origin of gymnosperms and the seed habit. 1898. 8°. From the Botanical Society of America. Darapsky, Luis. Las aguas minerales de Chile. Valparaiso, 1897. 8°. Las zeolitas de la coleccion mineralojica del Muséo Nacional. Santiago, 1888. 8°. La lengua Araucana. Santiago, 1888. 8°. Prom the Author. Fitzpatrick, T. J. Ferns of Iowa and their allies of Iowa. Lamoni, 1896. 8°. ; From the Author. Freire, Domingos. Mémoire sur la bactcériologie, pathogénie, traitement et pro- phylaxie de la fiévre jaune. Rio de Janeiro, 1898. 8°. From the Author. _ Hayden, Everett. Clock-rates and barometric pressure as illustrated by the mean- time clock and three chronometers at the Mare Island observatory. San Francisco, 1899. &°. From the Author. Honoré, Charles. Loi de rayonnement thermique solaire, ses principales consé- | quences et tables du soleil. Montevideo, 1896. 8°. From the Author. i Janet, Charles. Etudes sur les fourmis, les guépes et les abeilles. Note 14, 15, : 16. Limoges, ete., 1897. 8°. | Notice sur les trayaux scientifiques présentés par M. Charles Janet a l’Acad- émie des Sciences au concours de 1896 pour le prix Thore. Lille. 8°. From the Author. Klossoysky, A. Vie physique de notre planéte devant les Jumicres de la science : contemporaine. Odessa, 1899, 8°. Kuntze, Otto. Geognostische Beitrige. Leipzig, 1895. 8°. From the Author. Lamprecht, Guido. Wetterperioden. Bautzen. 1897, 8°. From the Author. Le Jolis, Auguste. Remarques sur la nomenclature bryologique. Cherbourg, 1895. 8°. From the Author. Lemoine, E. Mélanges sur la géométrie du triangle. Paris, 1895. 8°. Questions relatives a la géométrie du triangle. Paris, 1896. 8°. Note sur une construction approchée de déyelopement de la circonférence et remarques diverses. Paris, 1896. 8°. From the Author. ewis, Margaret. Studies on the central and peripheral nervous systems of two polychaete annelids. Boston, 1898. 8°.
XXIV Additions to the Library.
Lewis, Margaret. Clymene producta sp. noy. Boston, 1897. 8°. From the Author. Mueller, F. von. Index perfectus ad Caroli Linnzi species plantarum nempe earum primam editionem (Anno 1753), Melbourne, 1880, 8°. Wattle bark. Report of the board of inquiry. Melbourne, i692. 8°. From the Author. Nelson, E. W. Description of three new squirrels from South America. New
York, 1899: ‘8°: From the Author. Palmer, A. deF. On an apparatus for measuring very high pressures. New Haven, 1898. 8°. From the Author.
Rajna, Michele. Sull’ escursione diurna deJla declinazione magnetica a Milano in relazione col periodo delle macchie solari. Milan, 1895. 8°. From the Author, Riem, J. Ueber cine friihere Erscheinung des Kometen 1881 II Tebbut. Im Anschluss an die chinesische Annalen dargestellt von Dr. Joh. Riem. Gottingen, 1896. 8°. From the Author. Schiapareli, G. V. Osservazioni astronomiche e fisiche sull asse di rotazione e sulla topografia del pianeta Marte. Memoria TV, V. Roma, 1896-97. 4°. tubra canicula. Considerazioni sulla mutazione di colore che si dice ayve- nuta in Sirio. Rovereto, 1896. 8°. Origine de] sistema planetario eliocentrico pressi i Greci. Milano, 1895. 4°. From the Author. See, F. J. J. Researches on the evolution of the solar systems. Vol. I. On the universality of the law of gravitation and on the orbits and general characteristics of the binary stars. Lynn, 1896. 4°. Hrom the Author. Socolow, Serge. Nouvelles recherches astronomiques. Moscow, 1896. 8°. Des planétes se trouvant vraisemblemant au dela de Mercure et de Neptune. Moscow, 1897. 8°. Corrélations réguliéres du systéme planétaire avee Vindication des orbites des planétes inconnus jusqu’ici. Moscow, 1899. 4°. (4 copies.) From the Author. Stearns, Frederick. Hyalodendron navalium. A new genus and species of em-
plectellid sponge. Philad., 1898. 8°. From the Author. Tannert, A. C. Der Sonnenstoff als Zukunftslicht-und Kraftquelle. Eine physi- kalische Entdeckung. Neisse, 1896. 8°. Prom the Author.
Trelease, William. Botanical opportunity. 1896. 8°. From the Botanical Society of America. Verbeek, R. D. M. and Fennema, R. Description géologique de Java et Madoura. Amst., 1896. 2 vols. 8°, and atlas. f°. From the Minister of Colonies, Kingdom of the Netherlands. 4 Wadsworth, M. E. Eighteen papers, extracts from periodicals. vy. y. From the Author. Die Litteratur des Jahres 1892 iiber Morphologie, Systematik und Verbreitung der Phanerogamen, nebst Register. Berlin, 1895. 8°. Prom Die Verlagsbuchhandlung Gebriider Borntraeger, Berlin.
A REVISION OF THE NortH AMERICAN SPECIES OF FRULLANIA, 5 A Genus oF Hepatics. By AtexanperR W. Evans. Wir the single exception of Jungermannia itself, as defined by most recent writers, the genus Fiw/lania is the richest in species of all our hepatic genera, and the plants belonging to it are so distinct in their appearance and in their mode of life that the genus is one of the earliest which students of the liverworts learn to recognize. All ‘of our species attain their best development in rather exposed locali- ties, some of them on the trunks and branches of trees and bushes, others on rocks ; and, unless we see them soon after a shower or on a moist, cool day, they appear quite shriveled up and lifeless. At such times the plants are dark red or brownish-green in color, most of them adhere closely to bark or rock, and their stems, toward their extremities, look like fine, radiating, branched lines with round- ish irregularities produced by the leaves; in Frullania squarrosa the dry leaves are appressed to the stems and give them a some- what worm-like appearance. As soon as the plants absorb water, they become strikingly different ; their stems and leaves are no longer shrunken and brittle, but are turgid and flexible, and their colors are more lively and distinct. Several of our species are not absolutely restricted to exposed situations but are able to exist in more sheltered places; we find them, for example, on damp, shaded rocks, on rotten logs, or creeping over or through tufts of mosses. ‘Such plants are rarely satisfactory for study, their leaves are more ‘scattered than is normal, they reproduce almost entirely by vegeta- tive means, and they often fail to develop the water-sacs which are so characteristic of our genus. | As in nearly all large and natural genera, the species of Frudlania are difficult to define. Many of them are widely distributed and extremely variable, and the confusion to which these conditions “naturally give rise has been increased by the tendency among older writers of magnifying slight or temporary differences between plants into specific characters and, at the same time, of disregarding more f important points of distinction. In the Synopsis Hepaticarum of Fottsche, Lindenberg and Nees von Esenbeck (published from 1844 to 1847), twelve species are accredited to us; four of these are . synonyms of the common F. Eboracensis, leaving us, therefore, only “eight good species. During the forty years following the publication ‘Trans. Conn. Acap., VOL. X. May, 1897.
of this work, new North American species were described from time to time, mainly by Austin, until, in 1884, Professor Underwood * was able to ascribe to our region twenty species of the genus. Of these twenty species, however, three are synonyms and two of the others, FH Pennsylvanica and FE. Hutchinsie, var. (in reality the same plant’), have been transferred to the closely allied genus Jubula. This leaves sixteen species known at that time, including F. inflata, which is omitted in Professor Underwood’s paper. The few new species which have been added since 1884 and the few described in the present paper, increase the number to twenty-two, of which several are still known to us only from scanty or incomplete
2 A, W. Evans—North American Species of Frullania. |
j i j 4 4 material. |
The generic characters of /ullania are so well stated in accessible — literature, particularly in the writings of Spruce*® and of Schiffner’*, that it would be superfluous to detail them here. The remarks which follow are simply to call attention to certain interesting peculiarities in leaf and perianth and to make clearer the specific descriptions given later on.
The leaves of Frullania are unequally complicate-bilobed, and the antical or “ dorsal” lobe, which is called simply the “lobe,” is larger . than the other, spreads obliquely from the stem and is more or less orbicular in shape. In most cases, one side of this lobe arches over the stem and is often produced at the base into a cordate or auriculate expansion; the other side passes by a short and abrupt fold (except | in F/. arietina) into the postical or ‘ ventral” lobe of the leaf. This lobe, in turn, is deeply divided, usually to the very base, into two — unequal segments. The outer segment or “lobule” is the so-called “auricle” of older writers ; it is an extremely variable organ, but, in all of our species, is normally hooded over and inflated, sometimes throughout its whole extent, into a galeate or clavate structure, which serves as a sac or reservoir for the temporary retention of water.° The inner segment or “stylus” is usually much smaller than the lobule and is reduced in some cases to a minute, subulate process consisting of only three or four cells; in #4. Asagrayana, however, and in a few species allied to it, the stylus is larger and forms a disc-like cellular plate of considerable size. In & Caroli-
-
1 Bull. Illinois State Lab. Nat. Hist., ii: 61-68. 1884.
* Cf, Underwood, Bull. Torrey Bot. Club, xix: 301. 1892.
’ Hep. Amaz, et And., 3. 1884,
* Engler and Prantl, Die natiirlichen Pflanzenfamilien, Lief. 112: 132. 1896. * Goebel, Ann. du Jard, Bot. de Buitenzorg, vii: 21. 1888.
A. W. Evans—North American Species of Frullania. 3
niana@ and in one or two others, the first leaf of a branch sometimes develops both lobe and lobule into water-sacs; such a leaf is always quite covered over by other leaves.
It has already been noted that in sheltered places some species of Frullania may fail to develop water-sacs; the lobule under these circumstances is explanate and appears as a small, lanceolate, plane or slightly concave process; transitional forms may frequently be found between these explanate lobules and the typical inflated ones. : In every species, however, there are three leaf-modifications where explanate lobules normally occur. These are (1) the leaves from whose axils branches spring, (2) the perichetial bracts, and (3) the perigonial bracts. In the first of these, the stylus is about as large _as the lobule and the whole postical lobe is very like an underleaf ; occasionally the modified stylus bears a small tooth or secondary _ stylus on its inner edge. Even in this situation, although the stylus always retains its modified form, the lobule is sometimes inflated as in ordinary leaves. In the perichetial bracts, the lobule, which is _ often nearly as long as the lobe, is attached to it by a broad fold, and the stylus, which can usually be distinguished even in toothed bracts, appears on the inner edge of the lobule at some little distance from the base; occasionally the stylus is a segment of considerable _ size. In the perigonial bracts the lobe and the lobule are subequal and are connected by a broad fold to above the middle; the bracts are inflated so as to form hollow pouches for the antheridia, and the stylus, which is carried up on the inner edge of the lobule, usually remains minute.
The perianth of Frullania belongs to the hypogonianthous type as described by Spruce.’ In this type of perianth there are three keels, two lateral and one postical. The significance of this becomes evident if we consider that a perianth is normally formed by the coalescence of three floral leaves or “anthophylls,”* including two side-leaves and one underleaf. In case a species has flat leaves and bracts, the anthophylls are flat, their united edges give rise to the keels, and the perianth which results is triangular in section with an -antical keel and a flat, postical face corresponding with the flat _underleaf. This is the “epigonianthous” type and is well repre- sented by Lophocolea, In case a species has complicate leaves, the anthophylls are folded, the folds and not the united edges give rise
1 On Cephalozia, 5. 1882. The structure of the perianth is fully discussed in this paper, but the points brought forward are so important that it has seemed advisable _ to call attention to them again. 2 Spruce, l. ¢., 3.
4 A. W. Evans—North American Species of Frullania.
to the keels, and the perianth which results is triangular in section with a postical keel corresponding with the folded underleaf. This is the hypogonianthous type. In case no underleaf takes part in the formation of the perianth, the other two anthophylls unite postically and give rise to a perianth which is flattened, either laterally as in Plagiochila or antico-postically as in Radula. The typical charac- ter of the perianth of Hrudlania is often obscured by the interposition of supplementary keels or ridges.
A much less extensive coalescence is sometimes to be found in the involucre, where a bracteole may be connate on one or on both sides with the corresponding bracts. In several of our species the degree of such coalescence is by no means constant and it is only occasion- ally to be relied on as a specific character. In # Bolanderi, for example, the bracteole may be connate on both sides or on only one side; in J. Kunzet, it may be connate on one side or entirely free; while, in &? Virginica, all three conditions may be found.
The inflorescence of Frudlania is a character of great importance and should be determined wherever possible. The sporophyte, on the contrary, which is usually difficult to obtain and very uniform in structure, is of little value in distinguishing species and is not made use of in the following descriptions.
Frullania attains its greatest development in the tropics, where it is represented by numerous species in both hemispheres. In 1884, Spruce’ divided it into six subgenera, several of which are typically tropical. Five of these subgenera are represented in our flora, one being confined to Florida. Some of Spruce’s subgenera are connected by intermediate species and it is impossiblé to draw rigid lines of distinction between them ; still, they are for the most part natural assemblages of forms and are very convenient and useful. In Europe, eight species of Frullania have been recorded, three of them belonging to the subgenus Zrachycolea and the others to Thiop-— siella. Only one of these species, # Zumarisci, has been certainly found in North America, although two others have been accredited tous. <A striking peculiarity of our rullanie is the large number
of monoicous species among them ; no fewer than nine exhibit this character, while the eight European species are all dioicous.
Key to our Subgenera. Lobule inflated in the upper part only, connected with the lobe by a long fold subparallel with the stem; inflorescence paroicous ; perianth trapezoidal in section. SubgenusI. CHonanTHEtia.
' Hep. Amaz. et And., 7.
A, W. Evans—North American Species of Frullania. 5
_Lobule inflated throughout its whole extent or nearly so, connected with the lobe by a short fold approximately at right angles to the stem.
Lobule about as broad as long.
Underleaves not cordate at base; perianth typically triangular or trapezoidal in section, usually with tubercles or supplementary ridges or both,
Subgenus II. Tracuycorea.
Underleaves cordate at base ; perianth simply trigonous and smooth. Subgenus III. Homorropanrna.
Lobule decidedly longer than broad; perianth simply tri- gonous and smooth. Inflorescence dioicous ; perianth terminal on a sim- ple lateral branch. Subgenus IV. Turopsrewa.
Inflorescence autoicous ; perianth terminal on the stem or a main branch. Subgenus V. Drasronosa.
Supcenus I.—CHONANTHELIA Spruce. Represented by the single species :—
1. Frullania arietina Tayl. in G. L. et N. Syn. Hep., 413. 1845. PLATE I. figs. 1-6,
Paroicous: plants closely appressed to matrix, green, often tinged with yellow or brown: stems irregularly branched : leaves imbri- cated, the lobe orbicular-ovate, arching over the stem but scarcely cordate at base, plane or slightly decurved at the rounded apex, con- nected with the lobule by a long fold, subparallel to the stem ; lobule tubulose-inflated in the upper half only, the lower forming a plane, irregular or subrhomboidal, subentire expansion ; stylus minute, sub- ulate: underleaves contiguous or subimbricated, plane, orbicular, scarcely or not at all auriculate at base, shortly bifid at apex (4-4 with subobtuse lobes and sinus, entire or slightly crenulate on the margins; leaf-cells in middle of lobe rather thin-walled but with conspicuous trigones; toward the base the trigones are still more con- Spicuous and intermediate thickenings become abundant: @ inflores- cence terminal on a short lateral branch; bracts in one to three pairs, highly connate with the corresponding bracteoles, unequally bifid, the lobe ovate, acute, sparingly toothed, lobule ovate, acute, narrower than the lobe but similarly toothed and bearing a dis- tinct tooth or stylus at or below the middle of its inner edge ; brac-
6 A, W. Evans—North American Species of Frullania.
teoles bifid with narrow, acute or acuminate lobes, obtuse sinus and entire margins ; perianth immersed to or above the middle, com- pressed, oblong, abruptly narrowed into a short, broad beak, with two deep postical, and two less pronounced antical keels: @ bracts
in two or three pairs below the involucre, corresponding bracteoles
connate on one side.
Stems 0°18™" in diameter; lobes of leaves 1:20™™ long, 1:15™™ wide, lobules 0°82™™ long, 0°37"™ wide, inflated part 0:45™™ long; under- leaves 0°65"™ long and wide; leaf-cells from edge of lobe 0:017™™, from middle 0:029™", and from base 0:038™™ in diameter ; bract I,
lobe 1°65"™ long, 0°80"™ wide, lobule 0:80™™ long, 0°45™™ wide (to
point of coalescence with lobe and bracteole) ; bracteole I, 0°60™™ Jong, 0'40™™ wide (to point of coalescence); bract II, lobe 1-20™™ long, 0°70" wide, lobule 0°50™" long, 0°27"" wide, bracteole II, 0°37™™ long, 0°25"" wide; perianth 1°80™™ long, 0:90" wide.”
On trees ; Caloosa River, Florida (J. Donnell Smith).
Chonanthelia is better represented in the American tropics than in any other part of the world, and the range of our only known species extends as far south as Chili. Other members of the sub- genus should be looked for in southern Florida, Taylor’s original J arietina was apparently a composite species, and I have followed Spruce in restricting the name to plants with paroicous inflorescence.* This very unusual character and the peculiar lobules will serve at once to distinguish the plant from all our other Wrwllanie.
Suscenus Il.—TRACHYCOLEA Spruce. Key to the Species. Autoicous. Lobule more than half the size of the lobe. 2. F. Oakesiana.
Lobule less than half the size of the lobe, often explanate. Perianth truncate and abruptly narrowed into a short beak ; bracteoles free from the bracts. 4. FE! inflata.
Perianth not truncate, gradually narrowed into a short beak ; bracteoles connate on one side with bracts. 5. J. Cataline.
' The Roman numerals refer to the position of bract or bracteole: thus, I signifies the bract or bracteole next to the perianth; II, the bract or bracteole of the next outer row; and so on.
* The species of Frullania are of course not constant in size, and the measurements, which are taken from average-sized plants, are merely of comparative value.
* The specimens of /. arietina in the Taylor herbarium, all of which came from Demerara, the second of the localities mentioned in the Synopsis, are paroicous and agree with those described and distributed by Spruce,
A, W. Kvans—North American Species of Frullania, 7
Dioicous.
Lobes not cordate at base ; leaf-cells in middle of lobe with incon- spicnous trigones and no intermediate thickenings; branches often terminating in upright, leafless flagella with squarrose underleaves. 3. EH Bolanderi.
Lobes cordate or auriculate at base; leaf-cells in middle of lobe with conspicuous trigones and occasional or frequent interme- diate thickenings ; branches not terminating in leafless flagella.
Leaves strongly squarrose when moist. 7. E squarrosa.
Leaves scarcely or not at all squarrose when moist. Lobule usually explanate. 6. Ef. riparia.
Lobule usually inflated.
Lobule inflated in upper and outer parts, compressed below ; underleaves dentate or crenate on the sides above the middle ; perianth strongly tuberculate with more or less distinct supplementary ridges.
8. Lf. Britionie.
Lobule inflated throughout, underleaves entire or uniden- tate on the sides. Perianth with one or more distinct roughened supple- mentary ridges both antically and postically. 9. FF Virginica.
Perianth smooth and without distinct supplementary ridges. 10. & EHboracensis.
2. Frullania Oakesiana Aust., Proc. Acad. Phila. for 1869: 226. Puate I. figs. 7-15.
Autoicous: plants closely appressed to matrix, reddish-brown, vary- ing to greenish: stems irregularly pinnate; leaves imbricated, the lobe orbicular to ovate, slightly squarrose, arching over the stem but not cordate at base, slightly decurved at the rounded apex, entire or nearly so; lobule large, galeate, truncate at base, close to the stem; stylus minute: underleaves distant or subimbricated, plane, obovate or rhombic, bifid about one-third with acute lobes and narrow sinus, margin entire or unidentate on one or both sides: leaf-cells in middle of lobe with rather thick walls and inconspicuous trigones but with- out intermediate thickenings: @ inflorescence terminal on the stem or a main branch ; bracts in about two pairs deeply and unequally bifid, the lobe ovate, rounded at the apex, entire, lobule about as long as the lobe but narrower, ovate, obtuse or acute, entire but bearing a
8 A, W. Evans—North American Species of Frullania,
minute tooth or stylus at about the middle of its inner edge; brac- teole connate on one side, ovate, bifid with acute lobes and sinus, otherwise entire; perianth long-exserted, inflated or slightly com- pressed at sides, obovate, narrowed into a short, broad beak, with a broad usually two-angled postical keel and often with additional antical and postical ridges interposed: ¢ bracts in about two pairs occupying a short lateral branch near the involucre.
Stems 0°10™" in diameter; lobes of leaves 0°45™™ long, 0-35mm wide, lobules 0.23™™ long and: wide ; underleaves 0°18™™ long, 0°14™™ wide; leaf-cells from edge of lobe 0°013™", from middle 0:019"™ in diameter, and from base 0:025™™" long, 0°015™™ wide ; bract I, lobe 0°55™™ long, 0°30™™ wide, lobule 0°55™™ long, 6°23"™" wide ; bracteole — I, 0°45™™ long, 0:018™™ wide; bract II, lobe 0°45™™ long, 0°23™™" wide; bracteole II, 0°30"™ long, 0°12™" wide; perianth 1-00" long, 0°65™™ wide.
On trees, mostly at high altitudes; White Mountains, New Hamp- shire (Oakes, Austin, etc.): Mount Mansfield, Vermont (Farlow). Distributed in Hep. Bor.-Amer. n. 105c, and in Hep. Amer. n. 49.
This delicate little species seems to be quite local ; it has been col- lected perhaps a half dozen times in the White Mountain region, where it was discovered many years ago by Oakes, but I have seen specimens from only one other locality. Except for its small size, it would not be difficult of detection ; its reddish color usually serves to distinguish it from /? /’boracensis, a very common mountain species at lower altitudes, and there is little danger of its being confused with #. Asagrayana, the only other species found in the White Mountains. Aside from its color, #2 Oakesiana differs from F. Eboracensis in its autoicous inflorescence, in the areolation of its leaves, in its large lobule, and in the additional ridges on its perianth.
3. Frullania Bolanderi Aust., Proc. Acad. Phila. for 1869: 226. PLATE II.
Frullania Petalumensis Gottsche in Bolander, Catalogue of the Backs growing in the vicinity of San Francisco, 1870.
Frullania Hallii Aust., Bull. Torrey Bot. Club, vi: 20. 1875.
Dioicous: plants closely appressed to matrix, dark green, varying to reddish : stems irregularly pinnate, the branches often prolonged at right angles to the matrix as flagella without leaves, except a few toward the extremity, and with squarrose underleaves : leaves distant or subimbricated, the lobe ovate, somewhat squarrose when moist, arching over the stem but not cordate at base, rounded at the apex ; lobule large, galeate, truncate at base, close to the stem; stylus
A, W. Evans—North American Species of Frullania. 9
minute: underleaves distant, rhombic-ovate, bifid about one-third with subacute lobes and sinus, margin entire or bearing one or two teeth on the sides : leaf-cells of lobe with rather thick walls, trigones more conspicuous near the margin, intermediate thickening not de- veloped: @ inflorescence terminal on the stem or a main branch ; bracts in two or three pairs, unequally bifid, the lobe ovate, rounded at the apex, entire or nearly so, lobule narrower than the lobe, ovate, rounded or obtuse, entire but bearing a minute tooth or stylus near the middle of its inner edge ; bracteole connate on one or both sides, very variable, ovate, normally bifid about one-third with acute lobes and sinus, but sometimes rounded or merely emarginate at apex, sometimes with three or four more or less distinct teeth; perianth about half-emersed, obovate, narrowed into a short, broad beak, with a distinct, usually two-angled, postical keel and one or more antical and postical supplementary ridges: ¢ bracts in six to ten pairs, occu- pying a short lateral branch and forming an oblong spike.
Stems 0°10™ in diameter; lobes of leaves 0°35"™ long, 0°30™™ wide, lobules 0:25"™ long and wide; underleaves 0°18™™ long, 0°15™™ wide; leaf-cells from edge of lobe 0:016™", from middle 0:027, and from base 0:035"™ in diameter ; bract I, lobe 0°80™™ long, 0°50™™ wide, lobule 0°50™" long (to point of coalescence), 0°25™" wide ; bracteole I, 0°65™™ long, 0°35"" wide; bract II, lobe 0°65™™ long, 0°40™" wide, lobule 0°35™™ long, 0°15"™ wide; bracteole IJ, 0°60™™ long, 0°20™™ wide; perianth 1:25™™ long, 0°80™™ wide.
On trees ; west of the Rocky Mountains, from California to British Columbia. Distributed in Hep. Bor-Amer. 7. 1056, and in Hep, Amer. n. 28.
Frullania Bolanderi is the most widely distributed Zrachycolea of the Pacific Coast region and is common in many places. ‘There is little danger of confounding it with any other western species, and the remarkable, upright, leafless flagella, which are usually produced in greater or less profusion, are a ready means of distinguishing it from the eastern species which it most closely resembles. Its nearest ally is perhaps F: Oakesiana, which, aside from the absence of flagella, has a narrower perianth than / Bolanderi and an autoicous inflorescence. 2 Eboracensis occasionally produces flagella-like branches, but they are always leafy and are a rather unusual feature of the plant; its smaller lobule, different areolation, and smooth trigonous perianth will also serve to distinguish it. Mrullania Hallit is said by its author to be monoicous; the specimens of Hall and of Macoun, however, which I have been able to examine are all dioicous
10 A, W. Evans—North American Species of Frullania.
so that the monoicous character is at least very exceptional. Another point of distinction which Austin gives between his two species is in the character of the innermost bracteole—in / H/allii this is said to be entire or slightly emarginate at the apex, while in / Bolanderi it is said to be “acutely 2 (-4) toothed.” All of these conditions are sometimes found together, and the other less important differences given are no more constant.
4. Frullania inflata Gottsche in G. L. et N. Syn. Hep., 424. 1845. Puate III.
Autoicous: plants closely appressed to matrix, brownish-green varying to reddish: stems irregularly pinnate: leaves imbricated, the lobe orbicular, arching over the stem but not cordate at base, decurved at the rounded apex, entire ; lobule galeate, truncate at base, inflated especially in the upper and outer parts, separated from the stem by about one-fourth its width; stylus minute, subulate: under- leaves distant, orbicular or obovate, bifid about one-third with acute — or obtuse lobes and sinus, entire or nearly so: leaf-cells of lobe rather thick-walled with inconspicuous trigones and no intermediate thickenings : 9 inflorescence terminal on the stem or a main branch ; bracts in two or three pairs unequally bitid, the lobe ovate to obovate, rounded at apex, entire, lobule shorter and narrower than the lobe, ovate, rounded to subacute at the apex, bearing a small tooth or stylus below the middle of the inner edge, otherwise entire; bracteole free from bracts, ovate, deeply bifid with acute or obtuse lobes and sinus, entire or bearing one or two minute teeth toward the base; perianth exserted, more or less compressed when young, inflated when old, obovate, abruptly narrowed into a short, broad beak, with a distinct, angled, postical keel and usually with one or more supple- mentary antical and postical ridges: ¢ bracts in about two pairs, occupying a short lateral branch near the involucre and forming a short ovoid spike.
Stems 0°12™" in diameter; lobes of leaves 0°55™™ long, 0°60™™ wide, lobules 0:25™" long and wide; underleaves 0:30™™ long and wide; leaf-cells from edge of lobe 0°017™™, from middle 0:024"™™ and from base 0:°032™" in diameter; bract I, lobe 0°85™™ long, 050mm wide, lobule 0°65™" long, 0°40™™ wide; bracteole I, 0°65™™ long, 0:40" wide; bract II, lobe 0°65™™ long, 0°50™™" wide, lobule 0°60™™ long, 0°30"™" wide ; bracteole II, 0°65™™ long, 0:40™™ wide ; perianth 0°90™™" long, 0°65™" wide.
On trees; “ Whastite Red River (Beyrich)”: Baton Rouge, Louisi- ana (Joor): Point a la Hache, Louisiana (Langlois): Georgetown,
A, W. Evans—North American Species of Frullania. 11
D. C. (Coville): Austin, Texas (Underwood). On cypress pickets ; St. Martinsville, Louisiana (Langlois). Distributed as Prullunia Virginica in Hep. Amer. x. 68.
The determination of the plants which I have called Frullania inflata is based on a small scrap so named from Austin’s herbarium. This material was collected in Mississippi by E. Hall, and no nearer indication of its station is given. The species is apparently not rare in the western Gulf States and it is probably commoner elsewhere than collections would seem to indicate. At first sight the perianth of &. inflata resembles that of #. Virginica, especially when young, in having distinct supplementary ridges, but these ridges are never tuberculate as in that species and are usually quite smooth. It also differs from /. Virginica in its autoicous inflorescence, in its leaf- lobes, which are scarcely if at all cordate at the base, and in its areolation, the cells of the lobes having more uniformly thickened walls. There is little danger of confusing it with any other southern Trachycolea. Our New England F Oakesiana is a much smaller plant than / inflata and its large lobule and different perianth will readily serve to distinguish it.
5. Frullania Catalinee n. sp. PLATE IV.
Autoicous: plants growing in depressed tufts, reddish-brown, sometimes tinged with greenish: stems irregularly pinnate: leaves imbricated, the lobe ovate, squarrose when moist, arching over the stem but not cordate at base, slightly decurved at the rounded or obtuse apex, entire ; lobule broadly galeate, inflated, often imper- fectly developed as a water-sac or wholly explanate, separated from the stem by about one-fourth its width; stylus subulate, usually minute: underleaves distant, broadly rhombic, bifid to about the middle with obtuse or subacute lobes and sinus, entire or more commonly unidentate on the sides: leaf-cells with slightly thickened walls, inconspicuous trigones and no intermediate thickenings : 2 in- florescence terminal on the stem or a main branch; bracts in two or three pairs, unequally bifid, the lobe ovate, rounded or obtuse at the apex, entire or vaguely crenulate at base; lobule shorter and narrower, ovate, rounded to subacute at the apex, bearing a distinct tooth or segment (stylus) at or above the middle of the inner edge, otherwise entire ; bracteole connate on one side with bract, narrowly ovate, bifid one-third or more with lanceolate lobes and narrow sinus, otherwise entire ; perianth about half-exserted, somewhat compressed at least in the upper part, fusiform to pyriform, gradually narrowed
12 A, W. Hvans—North American Species of Frullania.
into a short, broad beak, with a distinct angled postical keel and one or more less pronounced antical and postical ridges: 2 bracts in one or two pairs, occupying a short branch near the involucre and form- ing a short, oval spike.
Stems 0°10™" in diameter; lobes of leaves 0°60"™ long, 0°55™™ wide, lobules 0:16"™ long, 0°23"™ wide (when explanate, 0°30™™ long,
0°23™™ wide); underleaves 0°30™™" long, 0°27"™ wide ; leaf-cells from:
edge of lobe 0:016™", from middle 0:020™™ in diameter and from base 0:030™" long, 0:020™" wide ; bract I, lobe 1:20™" long, 0°70™™ wide, lobule 0:80"™" long, 0°45™™ wide ; bracteole I, 0°75™™ long, 0°30™™ wide, bract I, lobe 0:90" long, 0°60"™ wide, lobule 0°60™™ long, 0°30"™" wide; bracteole II, 0°65"™ long, 0:20" wide; perianth 1:50™™ long, 0°75"™™ wide.
On rocks in a cafion; Catalina Island, California (McClatchie).
It will be seen from the foregoing description that F. Cataline is closely related to J” inflata. Its leaves, however, are much more
squarrose than in that species, its bracteoles are connate on one side, —
and its perianth, antheridial spike, and underleaves are different in shape. From & Bolanderi it differs most strikingly in its larger size, autoicous inflorescence and absence of flagella.
The first four species which I have placed in Zrachycolea form a rather distinct group by themselves and have the following characters in common :—(1) the leaf-lobes are scarcely or not at all cordate at base ; (2) the cells of the lobes are pretty uniformly thickened, hay- ing neither conspicuous trigones nor intermediate thickenings; (3) the postical keel of the perianth is more or less two-angled, so that the perianth is typically trapezoidal in section, although this condi- tion is usually obscured by the interposition of supplementary ridges ; (4) the keels and ridges of the perianth are not tuberculate, although they are sometimes slightly roughened or sinuous on the edges. In all of these points they differ from such typical Zrachy- coleue as F dilatata, F. Virginica and F. squarrosa, and seem to find their nearest allies in the last three South American Ayudlanic® which Spruce includes under Chonanthelia. Spruce’ suggests, how- ever, that these three species might better be placed in Zrachycolea, with which they certainly seem to have more in common.
' Hep. Amaz, et And., 29. 1884. we SRP i!)
a Sot de See f Oe Le’ Heh ee
SE ———— ee ee ee
~~
—— =
— a Se
ae
A. W. Evans—North American Species of Frullania., 13
6. Frullania riparia Hampe in Lehmann: Pugillus, vii, 14, 1838.
Frullania wolotis Mont. et Nees in Nees: Europ. Leberm., iii: 210. 1838 (nomen nudum).
Frullania eolotis Nees in G. L. et N. Syn. Wep., 417. 1845.
PLATE V.
Dioicous: plants growing in depressed tufts, green, sometimes tinged with brownish : stems loosely and irregularly pinnate : leaves distant to somewhat imbricated, the lobe ovate, slightly squarrose when moist, arching over the stem and cordate at base, plane or slightly decurved at rounded apex, entire or vaguely sinuate ; lobule when inflated a galeate sac truncate at base, when explanate (the usual condition) a small lanceolate lamina ; stylus minute: underleaves distant, rhombic to orbicular, bifid one-third or more with subacute lobes and sinus, entire or subdentate on the edges: leaf-cells in mid- dle of lobe with slightly thickened walls, distinct trigones and occa- sional intermediate thickenings, the latter disappearing and the tri- gones becoming more pronounced toward the base : @ inflorescence terminal on the stem or a main branch ; bracts in two or three pairs, deeply and unequally bifid, the lobe ovate, obtuse, entire ; lobule shorter and narrower, lanceolate, acute, bearing a small tooth or stylus on the inner edge near the base, otherwise entire ; bracteole free from bracts, narrowly ovate, bifid about one third with subacute lobes and narrow sinus, irregularly dentate or subentire on margin : perianth and ¢ spike not seen.
Stems 0°12™™ in diameter; lobes of leaves 0°60™™ long, 0°48™™ wide, lobules (when explanate) 0°30™" long, 0°12™" wide; under- leaves 0°30" long, 0°30™" wide ; leaf-cells from edge of lobe 0:015™", from middle 0:018™™ and from base 0:030™™ in diameter.
On trees and rocks, mostly in shaded places ; from New England westward to Minnesota and southward to the Gulf of Mexico. Dis- tributed in Muse. Alleg. n. 268 (as & dilatata, var. 2), in Hep. Bor.- Amer. n. 101 (as F- eolotis), and in Hep. Amer. . 140 (as &: wolotis).
Frullania riparia was first described from sterile material ; and, although a description of the involucre is added in the Synopsis Hepaticarum, the perianth and antheridial plant are apparently still unknown. In the absence of these data it is not possible to point out definitely the relationships of the species within the genus, but the general characters of leaves and underleaves, the dioicous in- florescence and the position of the female flowers show with little doubt that it isa true Zrachycolea. Its nearest relative seems to be the Italian #. Cesatiana De Not.,’ which is likewise incompletely
1 Mem. Accad, delle Sci. di Torino, II, xxii: 383, pl. 5. 1865.
14 A, W. Evans—North American Species of Frullania.
known, and Professor Massalongo' suggests that the two species may be identical. The Italian plant shows the same general appearance as ours, the same characters in lobes, underleaves and areolation, and the same great variability in the lobules, but it shows also slight differences in bracts and bracteoles. It seems safest, therefore, to keep the plants apart until both are better known, JF. riparia is most readily distinguished from its American allies by its lobules, which are rarely inflated but usually wholly or partially explanate. Of course a character of this sort is not very satisfactory, as sev- eral other species may show a similar variability in the shape of the lobule if growing in sheltered places. &: riparia also differs from F. squarrosa in its looser habit, less squarrose and narrower lobes and in its free bracteoles; from J! Virginica and F? Eboracensis, in its larger size, ovate lobes, and broader underleaves.
7. Frullania squarrosa (Bl. R. et Nees) Dumort., Recueil d’ Obs. sur les Jung., 13> 1835. ;
Jungermannia squarrosa Bl. R. et Nees, Nova Acta Acad. Caes. Leop., xii: 219. 1824.
PuaTeE VI.
Dioicous : plants closely appressed to matrix or more commonly growing in loose, wide mats, green, varying to reddish-brown: stems irregularly pinnate: leaves densely imbricated, the lobe rolled about the stem when dry, strongly squarrose when moist, very fragile in texture, broadly ovate, arching over the stem and cordate or auricu- late at base, rounded at the apex, entire; lobule galeate, inflated, especially in upper and outer parts, compressed at base, separated from the stem by about one-fourth its width ; stylus minute: under- leaves subimbricated, orbicular, plane or nearly so, entire or spar- ingly repand-dentate : leaf-cells from middle of lobe rather thick- walled with distinct trigones and intermediate thickenings: @ in- florescence terminal on a short lateral branch ; bracts in about three pairs, unequally bifid, the lobe ovate to orbicular ovate, rounded at the apex, entire; lobule ovate or broadly lanceolate, acute, bearing one or more small subulate teeth near the base on the inner edge, otherwise entire ; bracteole connate on one or both sides with bracts, approximately orbicular, deeply bifid to the middle or beyond with acute lobes and sinus, entire or slightly toothed or lobed toward the base, often revolute on the borders: perianth oblong, compressed, narrowed into a short broad beak, strongly unicarinate postically
* Atti del Congr, Nazionale di Bot. Crittog. in Parma, 10 (sep.) 1887.
A. W. Evans—North American Species of Frullania. 15
and bearing numerous scattered tubercles or scales, especially on the keels: ¢ plant not seen,
_ Stem 0°15™" in diameter; lobes of leaves 0:80™™ long and wide, lobule 0°23™" long, 0:20™™ wide; underleaves 0:45™™ long, 0°35™™ wide; leaf-cells from edge of lobe 0:016™" in diameter, from middle 0-027" long, 0°019"™ wide, and from base 0°030™™ long, 0:023™™ ' wide; bract I, lobe 1:00" long, 0-80" wide, lobule 0°75™™ long, 0-40™™ wide ; bracteole I, 0-75™™ long, 0:60" wide; bract II, lobe 0:90™™ long, 0°75™™ wide, lobule 0:65™" long, 0:30™ wide; bracteole II, 0°55"™ long, 0°30" wide; perianth 1:50™™" long, 0:90" wide.
On trees and rocks, from Connecticut to Ohio and southward ; common in the Southern States. Distributed in Hep. Bor.-Amer. nm. 100 and in Hep. Amer, 7. 94.
Frullania squarrosa is the most cosmopolitan of all our species, occurring almost everywhere in the warmer parts of the earth. The species 18 commonly sterile and plants with perianths are extremely ‘rare, although female plants without perianths are not unusual, Even in a sterile state there is no difficulty in distinguishing the plant, because the densely imbricated leaves, closely appressed to the stem when dry and strongly squarrose when moist, are unlike any- thing found in our other species. In the Southern States a form with the lobules pretty uniformly explanate sometimes occurs: this is apparently Frullania ericoides Nees, but there seems to be no good reason for keeping it distinct from /! squarrosa even as a variety.
8. Frullania Brittoniz n. sp. Frullania dilatata Underw. in Gray: Manual of Botany, sixth edition, 706. 1890
(not (L.) Dum.). Puate VII, figs. 1-12.
Dioicous : plants growing in wide depressed tufts, reddish-brown varying to greenish: stems irregularly pinnate: leaves imbricated, the lobe reniform-orbicular arching over the stem and strongly cordate or auriculate at base, plane or decurved at the rounded apex, entire ; lobule galeate, close to the stem, truncate and compressed at base, inflated in upper and outer parts ; stylus subulate, three to five cells wide at base: underleaves distant, broadly orbicular or elliptical, bifid about one third with obtuse, acute or apiculate lobes and acute sinus, irregularly dentate or crenulate on the sides above the middle: leaf-cells at margin of lobe with rather thin walls, distinct trigones and occasional intermediate thickenings, the last becoming fewer, the walls thicker and the trigones more conspicuous as we pass inward: @ inflorescence terminal on the stem or a principal branch ; bracts in two or three pairs, unequally bifid, the lobe ovate, rounded
16 A. W. Evans—North American Species of Frullania.
or obtuse (sometimes apiculate), entire or slightly crenulate, lobule
shorter and narrower, ovate or lanceolate, subacute or apiculate, —
bearing a small tooth or stylus at or below the middle of the inner edge, otherwise entire ; bracteole free or slightly connate on one side with bract, ovate, bifid one fourth or more with acute lobes and sinus, entire or unidentate on one or both sides; perianth emersed, obovate, truncate above and abruptly narrowed into a long, slender beak, compressed at the sides and with abroad postical keel and one or more short, supplementary antical and postical ridges, the whole surface being provided with scattered tubercles especially numerous on keels and ridges: ¢ bracts in many pairs, occupying a short lateral branch and forming an oblong spike.
Stems 0°18"" in diameter ; lobes of leaves 0°60™™ long, 0°75™™ wide, lobules 0:25"™ long and wide; underleaves 0°30™™ long, 0°37™™ wide; leaf-cells from edge of lobe 0:014™™", from middle 0:022™™ in diameter and from base 0:032™" long, 0°025™" wide ; bract I, lobe 1°35™™" long, 0°75™™ wide, lobule 1:00™™ long, 0°50™™ wide ; bracteole I, 0°85™™ long, 0°50™™ wide ; bract IT, lobe 1:05"™ long, 0°65™™ wide, lobule 0°75™" long, 0°65™™ wide; bracteole II, 0°70™™ long, 0°35™™ wide ; perianth 1°90™™ long, 1°20™™ wide.
On rocks and trees; Central New York (Underwood) : Holston River and Slemp’s Creek, Virginia (Mrs. Britton and Miss Vail) : Meriden, Connecticut (Evans): Canton, Illinois (Wolf): Easton, Pennsylvania (James). Distributed in Hep. Amer. x. 48 (as F. dilatata); in some sets there is admixture with #. Hboracensis.
In a sterile condition the present plant strikingly resembles the European /. dilatata, and it is little wonder that they have been considered the same. The involucre and perianth, however, afford safe points of distinction : in / dilatata the lobes of the bracts are broader than in our plant, the innermost bracteole is bifid with its
.
lobes deeply cut into two or three segments, and the perianth is sim-~
ply trigonous and narrowed into a short, broad beak. The long, slender beak of the perianth is indeed a most peculiar feature of F, Brittonice and serves, together with the numerous tubercles, to dis- tinguish the species from all other North American Prullanie. But, even in the absence of inflorescence, there is little danger of mistak- ing the present species, for the points which ally it with / dilatata separate it from other Zrachecolec, viz., the larger size of the plant, the curious inflation of the lobule and the broad underleaves with their peculiar dentation. I take pleasure in naming this distinct and beautiful species in honor of Mrs. Elizabeth G. Britton, whose care- ful work on American mosses is so highly appreciated by bryologists.
A, W. Evans— North American Species of Frullania. 17
9. Frullania Virginica Gottsche in Lehmann, Pugillus, viii: 19. 1844. Frullania saxicola Aust., Proc. Acad. Phila. for 1869: 225. Frullania Sullivantit Aust., Proc. Acad. Phila. for 1869: 226. PLATE VIII. Dioicous: plants closely appressed to matrix, green, varying to _ brownish : stems irregularly pinnate: leaves imbricated, the lobe _ suborbicular, arching over the stem and cordate at base, decurved at the rounded apex, entire; lobule galeate, truncate at base, somewhat inflated throughout, separated from the stem by about one-sixth its width ; stylus minute, two or three cells wide at base: underleaves distant, rhombic-ovate, bifid about one-third with subacute lobes and acute sinus, entire or rarely unidentate on the sides: leaf-cells of lobe rather thick-walled with conspicuous trigones and intermediate thickenings, especially toward the base: 2 inflorescence terminal on the stem or a main branch; bracts in two or three pairs, unequally bifid, the lobe squarrose, ovate to orbicular, rounded at the apex, entire, lobule ovate to lanceolate, acute or apiculate, bearing a small tooth-like segment or stylus at or above the middle of the inner edge, otherwise entire ; bracteole free or connate on one or both sides, ovate, bifid one-fourth to one-third with acute lobes and sinus, mar- gins entire, crenulate or slightly dentate; perianth hali exserted, somewhat compressed on the sides, obovate, abruptly narrowed into a short, broad beak, with a distinct angled postical keel and usually with two or more supplementary antical and postical ridges, more or less tuberculate, particularly on keels and ridges: ¢ bracts in many pairs, occupying the end or middle part of a short lateral branch and forming an oblong spike. Stems 0°10™" in diameter ; lobes of leaves 0°55™™ long, 0°45"™ wide, lobules 0°28™" long, 0°18™" wide; underleaves 0°22™" long, 0°15™™ wide; leaf-cells from edge of lobe 0:014"™, from middle 0°018™, and from base 0°025™" in diameter; bract I, lobe 0°80" long, 0°65™™ wide, lobule 0:75™™ long, 0°35"" wide ; bracteole T.0:302™stong, 0°55™"= wide; bract II, lobe 0°65™™ long, 0°55™™ wide, lobule 0°65"™" long, 0:30™" wide; bracteole II, 0-60"™ long, 0°25™™ wide ; perianth 1.85™™ long, 1°00™™ wide. On trees or, more rarely, on rocks; from Canada to the Gulf of Mexico: rare in the north but becoming abundant southward. Dis- tributed in Muse. Alleg. x. 267 (as F. dilatata, var. 1) and in Hep. Bor.-Amer. n. 103 and n. 104 (as &. saxicola). It will be seen that there have been included under this very varia- ble species two forms which were considered distinct by Austin. May, 1897.
TRANS. Conn. ACAD., VOL. X. 2
18 A, W. Evans—North American Species of Frullania.
The following statement is quoted from the description by that author of his Frullania saxicola :— Perianth longer than in /. Vir- ginica and more exserted, but angled much in the same manner; however, the angles are never crested, and the ‘style’ or mouth is very different ; (tubular and considerably elongated in #. Virgin- ica).” The perianth is so extremely variable an organ that the dif- ferences brought forward are hardly sufficient to keep the plants dis- tinct, particularly as the characters derived from leaves, underleaves and involucres are almost identical in the two. The short beak of F. saxicola is at first sight a striking peculiarity, but there are inter- mediate grades between it and the typical beak of #. Virginica, while the absence of crests is a rather inconstant feature In his account of 7” Sullivantii, Austin gives no direct comparison with F. Virginica but indicates the following differences in his description : —the larger lobule, the connate bracteole, the fewer keels in the perianth. Differences as great as these may sometimes be found in a single specimen. I have been able to study the types of both of Austin’s species and find no greater differences than those enumer- ated.
10. Frullania Eboracensis Gottsche in Lehmann, Pugillus, viii: 14. 1844. Frullania saxatilis Lindenb., in G. L. et N. Syn. Hep., 424. 1844. Frullania microscypha Tayl., Lond. Jour. Bot., v: 402. 1846. — Frullania leviscypha Tayl.,1. ¢., v: 403. 1846. Frullania nana Tayl., 1. ¢., v: 404. 1846. PuatTE IX. figs. [Te
Dioicous: plants closely appressed to matrix, usually green but often tinged with brown or red; stems irregularly pinnate, some- times flagelliferous ; leaves imbricated, the lobe suborbicular, arch- ing over the stem and cordate at base, rounded at the slightly de- curved apex, entire; lobule galeate, truncate at base, separated from the stem by about one-sixth its width; stylus minute, two or three cells wide at base: underleaves distant, ovate or rhombic-ovate, bifid about one-third with subacute lobes and sinus, entire or obscurely unidentate on the sides: leaf-cells of lobe rather thick-walled with trigones and intermediate thickenings, the latter becoming fewer toward the base: Q inflorescence terminal on the stem or a main branch ; bracts in two or three pairs, unequally bifid, the lobe ovate, rounded at the apex, entire or slightly crenulate toward base; lobule narrower than the lobe, ovate, acute or obtuse, bearing a small tooth- like segment or stylus at about the middle, otherwise subentire ; bracteole free or connate on one side, ovate, bifid one-third or niore,
A. W. Kuans—North American Species of Frullania, 19
‘with acute lobes and sinus, entire or irregularly dentate on the sides : perianth obovate or obcuneate, more or less compressed, abruptly ‘narrowed into a short, broad beak, with a distinct, sometimes two- angled postical keel but without distinct supplementary ridges, _ smooth or slightly roughened on lateral keels, never tuberculate : “a spike oblong, occupying a short lateral branch, bracts in many _ ~pairs. Pa Stem 0°10™™ in diameter, lobes of leaves 0°45™™ long and wide, _ lobules 0:21™™ long and wide ; underleaves 0:20" long, 0°15"™ wide; leaf-cells from edge of lobe 0: 014™™", from middle 0:017™" in Ais ter, and from base 0:030™™ long, 0:017™" wide; bract I, lobe 0-30™™ long, 0°45" wide, lobule 0:60™™ long, 0: 30m wide ; bracteole I, 055™™ long, 0°23" wide; bract II, lobe 0-60™™ ae 0°38™™ wide, lobule 0°-45™™ long, 0:18™" wide; bracteole I, 0:45™™ long, 0°15™™ _ wide ; perianth 1:10™™ long, o-75mm wide. - On trees and rocks ; from Canada to Florida and westward to Minnesota: very common in the mountains and in northern regions. Distributed in Hep. Bor.-Amer. n. 105, in Hep. Amer. v. 27, and in Can. Hep. n. 1. Frullania EHboracensis is characteristically a northern species and reaches the south only as a rarity, whereas the reverse is true for _ Virginica. In a sterile condition the two species sometimes resemble each other so closely that it is difficult if not impossible to tell them _ apart and we must depend upon perianths for differential characters _ which are constant. The perianth of #2 Eboracensis is very variable ~ both in shape and in the character of the postical keel, but it has the - unusual feature among Zrachycolew of being smooth and without ‘supplementary ridges. In some cases, however, there is a slight trace of an antical ridge, although this seems to be an exceptional ‘condition. F. Virginica is of course distinguished by its tuberculate perianth with distinct supplementary ridges.
Suseenus I1Il.— HOMOTROPANTHA Spruce.
Represented by the single species :— ili. Frullania plana Sulliv., Mem. Amer. Acad., new series, iv: 175. 1849. a Puate IX. figs. 12-21. Autoicous: plants growing in wide depressed tufts, green, some- times tinged with brown: stems irregularly pinnate or bipinnate: leaves imbricated, the lobe orbicular, arching over the stem and
PP oncly cordate or auriculate at the base, decurved at the rounded _ apex, entire ; lobule galeate, close to the stem, truncate at base, inflated particularly in upper and outer parts, stylus minute : under-
20 A. W. Evans—North American Species of Frullania.
leaves distant, reniform, cordate at base, bifid about one-fourth, with obtuse or subacute lobes and sinus: leaf-cells of lobe rather thick- walled with conspicuous trigones and intermediate thickenings : 2 inflorescence terminal on a short, simple, lateral branch ; bracts in about three pairs, unequally bifid, the lobe ovate, rounded at the apex, irregularly crenulate, lobule shorter and narrower, ovate, rounded at apex, irregularly crenulate and bearing at or below the middle of the inner edge a tooth-like, often laciniate segment or stylus ; bracteole free from bracts, ovate, deeply bifid with subacute lobes and sinus, the lobes variously laciniate, dentate or crenulate ; perianth about half exserted, oblong or obovate, narrowed into a
short, broad beak, compressed on the sides, with a broad postical
keel and a shallow antical sulcus, smooth: ¢ spike terminal on a short lateral branch, globose, bracts in two or three pairs.
Stems 0°18™" in diameter; lobes of leaves 0°60™" long, 0°75™™ wide, lobules 0°18"™ long and wide; underleaves 0 36"™ long, 0°40™™ wide ; leaf-cells from edge of lobe 0:014™™, from middle 0°019™™, and from base 0°028™™" in diameter; bract I, lobe 1:00" long, 0°65™™ wide, lobule 0°60" long, 0°35"" wide; bracteole I, 0°65™™ long, 0:40™" wide ; bract II, lobe 0:80™™ long, 0°60™™" wide, lobule 0°45™™ long, 0:30"™ wide ; bracteole I, 0°50™™ long, 0:-40™™ wide ; perianth 1:90™™ long, 0°90™™ wide.
On shaded rocks: French Broad River, Tennessee (Sullivant) ; Closter, New Jersey (Austin); Sand Lake, New York (Peck) ; Woodbridge, Connecticut (Evans). Distributed in Muse. Alleg, n. 269 (as # dilatata, var. 3) and in Hep. Bor.-Amer, n. 102.
Frullania plana is by no means a typical Homotropantha but it shows its relationships with this group rather than with Zrachycolea by the union of the following characters: —(1) the broad, cordate underleaves, (2) the autoicous inflorescence, (3) the female flowers
ee Re ee ee ee a ee eee ME en ee om
te Kee ee SE ah
borne on simple lateral branches, and (4) the smooth trigonous —
perianth. The lobule, however, although small for the size of the plant, is never reflexed, as in J! replicata, etc. The present plant is
usually sterile, but its peculiar underleaves will serve to distinguish -
it even in this condition.
Susgenus IV.—THIOPSIELLA Spruce. Key to the Species. Underleaves reflexed, at least toward the apex. Lobes acuminate or acute; underleaves not crispate at base ; bracts subentire ; innermost bracteole connate on both sides. 12, /. Nisquallensis,
a ae os
A, W. Hvans—North American Species of Frullania. 21
Lobes acute or obtuse; underleaves strongly crispate at base ; bracts more or less dentate ; innermost bracteole free.
15. F Tamarisci.
Lobes obtuse or rounded; underleaves not crispate at base ; bracts entire or slightly dentate ; innermost bracteole usually ° free. 14. #, Asagrayana.
Underleaves plane or nearly so. Lobes marked with a distinct line of discolored cells; under- leaves entire on the sides, sometimes auriculate at base; stylus 4 a conspicuous disc-like process; stylus of lobule of bract a distinct segment. 14. Ff Asagrayana.
Lobes marked with a distinct line of discolored cells ; under- leaves often unidentate on the sides, never auriculate at base ; stylus small or minute ; stylus of lobule of bract not distinct, replaced by a cluster of fine laciniz or cilia.
13. 2 Franciscana.
Lobes with or without scattered discolored cells ; underleaves entire on the sides; stylus minute ; stylus of lobule of bract a distinct segment. 16. 4. Californica.
12. Frullania Nisquallensis Sulliv., Mem. Amer. Acad., new series, iv: 175. 1849. PLATE X.
Dioicous: plants robust, growing in broad, depressed tufts, red- dish-brown, usually tinged with yellow or green: stems mostly bipinnate : leaves imbricated, the lobe ovate, arching over the stem and cordate at base, strongly reflexed at the acute or acuminate apex, margin entire; lobule separated from the stem by about its own width, oblong-clavate ; stylus minute and subulate or sometimes a small disc-like process : underleaves distant or contiguous, orbicular or reniform, strongly reflexed, at least at the apex, bifid about one- fourth with obtuse lobes and sinus, auriculate at base: leaf-cells of
lobe rather thick-walled, trigones and intermediate thickenings becoming prominent towards the middle ; discolored cells usually absent, sometimes occurring on the leaves of ultimate branchlets : 9 inflorescence terminal on a short, lateral branch ; bracts in about three pairs, deeply and unequally bifid, the lobe ovate, acuminate, entire, sinuous or very sparingly dentate on the margin, lobule subulate, often distorted or uncinate at the acuminate apex, revolute on the margins, bearing on the inner side towards the base an indis-
29 A. W. Evans—North American Species of Frullania.
tinct laciniate lobe or cluster of cilia (stylus) ; bracteole (at least the innermost one) connate on both sides with bracts, ovate, bifid one- third or more with narrow subulate, acuminate lobes and narrow sinus, margins revolute and entire above, dentate or ciliate toward base ; perianth exserted one-third or more, ovate or oblong, gradu- ally narrowed into a short broad beak, concave antically, compressed on the sides and deeply one-keeled postically : antheridial spike oval, occupying a short lateral branch, bracts in several pairs. |
Stems 0°30™" in diameter; lobes of leaves 1:20" long, 0°85™™ wide, lobules 0:20" long, 0°15" wide; underleaves 0°68" long,
oes Sw nine Br eet
0°85™" wide; lobes of branch-leaves 0°60" long, 0°35™™ wide; ~ branch-underleaves 0°25"" long and wide ; leaf-cells from edge of —
lobe 0:014™™, from middle 0:023™™ in diameter, and from base 0:035™™ long, 0:023"™ wide ; bract I, lobe 2:00™™ long, 0°75™™ wide; lobule 0°75™™ long, 0°15™™ wide; bracteole I, 1°35™™ long, 0°60™™ wide ; bract II, lobe 1:25™™ long, 0°65™™ wide, lobule 0;55™™ long, 0:10™™ wide; bracteole II, 0:85™" long, 0°30™ wide ; perianth 2°50™™ long, 1:00™™ wide.
On rocks and trees, from Alaska to northern California. Distrib- uted in Can. Hep. 2.3 (as /) Asagrayana, var.), n. J (as #. Asagray- ana, var. Californica), and n. 5 (in part).
The determination of this species is based on Sullivant’s descrip- tion and on drawings in his herbarium: the type specimen was sent by him to Gottsche, together with the rest of the Wilkes’ hepatics, and is presumably in Berlin. #7 Wisqguallensis has frequently been confused with 4! Tamarisci.
.
13. Frullania Asagrayana Mont., Ann. des Sc. Nat., II. xviii: 14. 1842 (footnote). PLATE XI.
Dioicous : plants growing in depressed or pendulous tufts, reddish-
brown or more rarely paler and greenish: stems once or twice pin-~
nate: leaves imbricated, the lobes ovate, arching over the stem and cordate at base, rounded or obtuse at the decurved apex, entire; lobule separated from the stem by about half its width, obovoid- clavate, contracted toward base; stylus a suborbicular disc-like process bearing on its margin one or two minute cilia or run- ning out into. one or two acute points: underleaves distant, orbic- ular-ovate, plane or rarely reflexed at the apex, bifid less than half with obtuse lobes and sinus, sometimes appendiculate or slightly auriculate at base: leaf-cells of lobe thick-walled, trigones and inter- mediate thickenings becoming more conspicuous toward the middle
See
2 v
A, W. Hvans—North American Species of Frullania. 2
and base; discolored cells usually forming a long, distinct, median line, rarely obsolete: ¢ inflorescence terminal on a short branch; bracts in two or three pairs, bifid to or beyond the middle, the lobe ovate, acute, entire or sparingly dentate, becoming broader and more obtuse away from the perianth, lobule narrowly ovate or lanceolate, acuminate, usually revolute on the margins, bearing at the base on the inner edge a more or less distinct, variously toothed or laciniate segment or stylus, otherwise entire; bracteole free or connate on one side, ovate, bifid to or beyond the middle with subulate acuminate
lobes and pointed sinus, margin usually bearing at the base on each
side a variously toothed or laciniate segment, otherwise entire ; per- ianth exserted beyond the middle, oval or obovate, narrowed into a rather short beak, somewhat compressed on the sides and with a deep postical keel, smooth: ¢ spike oval, occupying a short lateral branch, bracts in several pairs (six to ten).
Stems 0°15™™ in diameter; lobes of leaves 0°70™™ long, 0°50™™ wide; lobules 0°25"™ long, 0°17"" wide ; underleaves 0°35™™ long
_ and wide ; lobes of branch leaves 0°50"™ long, 0°35"" wide ; branch- _ underleaves 0°25™" long, 0:°15™" wide; leaf-cells at edge of lobe
0:014™™, in the middle 0:017™™ in diameter and at the base 0:028™™ long, 0°018™™ wide ; bracts I, lobe 1°50™™ long, 0°70" wide, lobule 0°85™" long, 0°25™™ wide; bracteole I, 1:20™™ long, 0°50™™ wide ; bract II, lobe 0°95™™ long, 0°60™" wide ; lobule 0°50™™ long, 0°15™™
wide; bracteole II, 0°70™™ long, 0°25"™ wide ; perianth 1°85™™ long,
0°90"™™ wide.
On rocks, on bark of trees, or pendulous from small branches : from Newfoundland to Georgia and west to Wisconsin. Common in the Eastern States, especially in hilly or mountainous regions.
‘Distributed in Muse. Alleg. ». 266, in Hep. Bor.-Amer. 7». 107, in
_ Hep. Amer. 7. 7, and in Can. Hep. x. 2.
14, Frullania Tamarisci (L.) Dumort., Recueil d’Obs. sur les Jung., 13. 1835. Jungermannia Tamarisci L., Species plantarum, 1134. 1753 (Ed. I).
Frullania major Raddi, Mem. di Matem. e di Fiscia aan Soe. Ital. della Sei. (Mo- dena), xviii: 20, pl. 2. 1820. Jubula Tamarisci Dumort., Comm. Bot., 112. 1822. Puate XII. figs. 1-10. _ Dioicous : plants growing in depressed tufts, reddish-brown, rarely tinged with green : stems mostly bipinnate : leaves imbricated, the lobe ovate-orbicular, arching over the stem and deeply cordate at
24 A. W. Evans—North American Species of Frullania.
base, obtuse, apiculate or acute at the decurved apex (on ultimate branches sometime acuminate); lobule separated from the stem by about half its width, subparallel with the stem, short-clavate, con- tracted toward base; stylus a small disc-like or strongly crispate
rahe
4
4
process: underleaves distant, orbicular, strongly reflexed at apex and —
usually on the sides, bifid about one-sixth with a broad, shallow sinus and obtuse or apiculate lobes, crispate-auriculate at base: leaf-cells
of lobe thick-walled, trigones and intermediate thickenings becoming ~
more pronounced on passing inward and toward the base; discolored cells usually indistinct, either scattered or in a short median line : 2 terminal on a short branch; bracts in three or four pairs unequally
bifid, the lobe ovate, acute, irregularly dentate or crenate, especially
in the upper part, lobule lanceolate, acuminate, revolute on the mar- gins, bearing at the base on the inner side a cluster of fine cilia, otherwise subentire: bracteole free from the bracts, deeply bifid with ovate-lanceolate, acute, irregularly dentate or laciniate lobes and narrow sinus, margins ciliate at base, perianth exserted one-third or more, oblong, narrowed into a short beak, postically strongly one- keeled, smooth: ¢ spike oval, borne on a short lateral branch, bracts in several pairs.
Stems 0°20™™ in diameter; lobes of leaves 0:80"™ long, 0°75™™ wide, lobules 0°25" long, 0°10 wide; underleaves 0°45™™ long, 0-40™™ wide; lobes of branch-leaves 0°35™™ long, 0°25™™ wide ; branch-under- leaves 0°18™™ long, 0°15™" wide; leaf-cells from edge of lobe 0°012™™, from middle 0:019™™ in diameter, and from base 0°038™™ long, 0:022™™ wide; bract I, lobe 1:20™" long, 0°60"™ wide, lobule 0-75™™ long, 0°30" wide, bracteole I, 0:80™™ long, 0°60™" wide ; bract II, lobe 1:10™" long, 0°55™™ wide, lobule 0°60™™ long, 0:24"™ wide; bracteole IT, 0:90™™ long, 0°45™™ wide; perianth 2°15™™ long, 0:90"™ wide.
On rock and trees ; Miquelon Island (Delamare): Newfoundland
(Waghorne): Blackstone, Rhode Island (Bennett). ‘‘ Vancouver and
Orcas Islands, Lyall. Collected also on the N. W. coast by Menzies and Douglas.”* Apparently rare, but probably more abundant in the far north. ‘
In the absence of fertile American material the above descriptions of involucre and perianth are drawn from Swedish plants collected by Dr. Arnell. The Rhode Island specimens which I have had an opportunity of examining are very fragmentary, but are apparently referable to this species.
* Mitten, Jour. Linn, Soc., viii: 53. 1865. It is probable that these specimens from the Pacific Coast would now be referred to other species. ¢
‘
A. W. Evans—North American Species of Frullania.
b> or
15. Frullania Californica (Aust.).
Frullania Asagrayana, var. Colifornica Aust. in Underwood, Bull. Ill. State Lab, Nat. Hist., ii: 67. 1884 (in part).
Frullania Asagrayana, var. Californica Aust. (emend.), Howe, Erythea, ii: 98. 1894. _ Frullania Asagrayana, var. alsophila Howe, 1. c., ii: 99. 1894.
Frullania Tamarisci Bolander, Catalogue of the Plants growing in the vicinity of San Francisco. 1870 (not (L.) Dum).
PLATE XII, figs. 11-22.
Dioicous : plants closely appressed to matrix or growing in wide depressed tufts, green, varying to brownish-red : stems once to thrice pinnate, often irregularly so: leaves subimbricated, the lobe orbicu- lar, arching over the stem and cordate at base, rounded and more or less decurved at the apex, entire ; lobule separated from the stem by less than half its width, obovate-clavate; stylus a minute subulate or disc-like process : underleaves orbicular, plane or slightly refléxed on one or both sides toward the base, bifid about one-third with obtuse or subacute lobes and sinus, margin entire, sometimes slightly auricu- late at base ; leaf-cells rather thick-walled, trigones inconspicuous and intermediate thickenings scanty ; discolored cells wanting or few in number, scattered or more rarely in a short median line: Q inflores-
_ cence terminal on a short branch ; bracts in two or three pairs, un-
equally bifid, the lobe ovate or ovate-lanceolate, acute, entire, lobule
lanceolate or subulate, acuminate, margin more or less reflexed, bear-
ing on the inner side at the base a laciniate lobe-like segment or
stylus, otherwise entire, bracteole connate on one side with bract,
ovate, bifid to about the middle with lanceolate acuminate lobes and acute sinus, bearing toward the base on each side a distinct usually laciniate or ciliate sezment, otherwise entire ; both bracts and bracte- oles becoming smaller and simpler on receding from the perianth ; perianth about half exserted, compressed on the sides, oval, narrowed into a short beak, deeply one-keeled postically, smooth; ¢@ spike on a short lateral branch, globose, bracts in about two pairs.
Stems 0°15"™" in diameter; lobes of leaves 0°50" long, 0°45™™ wide, lobules 0°17"" long, 0:09" wide ; underleaves 0°22" long, 0°25™" wide; lobes of branch-leaves 0°22" long, 0°17"™ wide ; branch-underleaves 0°12" long, 0:09™" wide ; leaf-cells at edge of lobe 0°010"" in the middle, 0°014"" in diameter, and at the base 0-030™ long, 0:022™" wide ; bract I, lobe 1:40™™ long, 0°55™™" wide, lobule 0°75™ long, 0°25™" wide; bracteole I, 1:00"™ long, 0°50™™ wide ; bract II, lobe 0°90™ long, 0°50" wide, lobule 0°45"™ long,
26 A, W. Evans—North American Species of Frullania.
0:20"™" wide; bracteole II, 0°80™™ long, 0°50™™ wide ; perianth 1°70™™ long, 0°80™™" wide.
On rocks and trees: British Columbia to California. Distributed (as 4, Nisquallensis) in Hep. Bor.-Amer. n, 108 (in part), in Hep. Amer. 2. 102 (as #. Asagrayana, var. Californica) and n. 148 (as F, Nisquallensis), and (also as . Nisquallensis) in Can. Hep. n. 4 (in part).
The specimens distributed in Hep. Bor.-Amer., which we may con- sider the type of Austin’s /. Asagrayana, var. Californica, are a mixture of this and the next species, and it seems allowable to retain the name “ Californica” for the present series of forms,
16. Frullania Franciscana Howe, Erythea, ii: 99. pl. 2. 1894.
Frullania Asagrayana, var Californica Aust. in Underwood, Bull. Illinois State Lab. Nat. Hist , ii: 67. 1884 (in part).
Frulidnia unciflora, var. Californica Gottsche in Bolander: Catalogue of the plants
growing in the vicinity of San Francisco. 1870. PLATE XIII. figs. 1-8.
Dioicous : plants appressed to matrix or growing in wide depres- sed tufts, reddish-brown, varying to greenish ; stems mostly bipin- nate: leaves imbricated, the lobes ovate, arching over the stem and cordate at base, rounded, obtuse or apiculate at the. decurved apex,
entire ; lobule separated from the stem by about its own width, »
short-clavate ; stylus minute: underleaves distant, plane, rhombic- ovate, bifid about one-third with obtuse lobes and narrow sinus, margins usually bluntly unidentate at about the middle, neither auriculate nor appendiculate at base : leaf-cells of lobe rather thick- walled, trigones and intermediate thickenings becoming more con- spicuous toward the middle and the base ; discolored cells usually in a short median line, sometimes obsolete: 9 inflorescence terminal ona
short branch; bracts in about three pairs, unequally bifid, the lobe ~
ovate, acute or acuminate-apiculate, ciliate at antical base, otherwise entire ; lobule ovate or lanceolate, acute, bearing a cluster of cilia toward base, otherwise entire; bracteole connate on one side with bract, ovate, bifid about one half with lanceolate, acuminate lobes and narrow sinus, ciliate at base, otherwise entire; perianth oblong- obovate, abruptly short-rostrate, compressed on sides and with a deep postical keel, smooth.
Stems 0'17"" in diameter: lobes of leaves 0°95™™" long, 0°70™™ wide, lobules 0:25"" long, 0-12™ wide ; underleaves 0:40™™ long, 0°35™™" wide ; lobes of branch-leaves 0°35™™ long, 0°25™™ wide ;
ee ee Oe re oe ee
‘ag TS ata, 5 far,
Retest cos Staab Jk.
Se
A. W. Evans—North American Species of Frullania. 27 branch-underleaves 0:17" long, 0°15™" wide ; leaf-cells at edge of lobe 0:014™ in the middle 0-019™ in diameter and at the base 0:035"™ long, 0°023™™" wide ; bract. I, lobe 1°50™™ long, 0°75™™ wide, lobule 0:60"™ long, 0°25™" wide ; bracteole J, 1:35™" long, 0°60" wide ; bract II, lobe 1:25" long, 0°50™™" wide, lobule 0°40™™ long, 0°20™™ wide ; bracteole IT, 1:00™™ long, 0°50™" wide ; perianth 2:20™™ long, 1:00™™ wide.
On trees ; California. Distributed (as #. Wisquallensis) in Hep. Bor.-Amer. 7. 108 (in part).
The plants of the present group have long been a puzzle to Ameri- can hepaticologists ; most of them are species of wide range, they vary greatly according to environment and are apparently connected with one another by transitional forms. In sterile material, more- over, the essential characters of a species are often so slightly developed as to make determination difficult if not impossible, and the same thing is true, though in a far less degree, of antheridial material. In the eastern parts of the United States the only com- mon representative is Mrullania Asagrayana, a plant which fre- quently assumes forms very unlike the specimens originally described by Montagne: the most important differences brought out in this description between our plant and /. ZYamarisci are in the under- leaves, which are said to be plane, and in the perichetial bracts, which are said to be subentire. Neither of these characters is con- stant ; the underleaves may be reflexed, and the bracts are some- times dentate. The underleaves, nevertheless, do afford us a second and more important distinction in the basal auricles or lobes some- times found in / Asagrayana ; these are never crispate as in the constant and well developed auricles of #. Tamariscz. The stylus of F. Asagrayana is unusually large, being sometimes as long as the lobule. A somewhat similar stylus is sometimes found in other species of this group, butit is always smaller than in /. Asagrayana ; and, in # Tamarisci, it is more or less crispate like the auricles of the underleaves. In /. Asagrayana, finally, the lobules of the peri- cheetial bracts and usually also the bracteoles bear at the base more or less distinct, laciniate segments : these reappear in the western / Californica, but are unlike the equivalent structures found in our other species.
In the far north the closely related #. Zamarisci, which is abun- dant in Europe, apparently becomes more common, sometimes occurring in company with # Asagrayana. The most important points of difference between the two are indicated above, but the
28 A. W. Evans—North American Species of Frullania.
involucres furnish one or two additional ones : in 2 Tamarisci bracts and bracteoles are relatively broader and the distinct stylus of #. Asagrayana is replaced by a cluster of fine cilia. The leaves of F.. Tamarisci, also, are usually somewhat pointed, while the median row of discolored cells is rarely distinct.
As we go westward J Nisquallensis, another close ally of F. Tamarisci, makes its appearance. The leaves of this species are still more sharply pointed, and the reflexed underleaves show basal auricles which are not strongly crispate as in # Tamarisci. The involucres, also, provide us with important differences; in 7 Wis- quallensis, the bracteoles, at least the innermost, are connate on both
sides with the adjacent bracts, and the lobes of the latter are narrow, —
acuminate and subentire.
In addition to #! Nisquallensis, we have in the west the plants which have been known as #! Asagrayana, var. Californica, The confusion in regard to these plants has been partially cleared up by Mr. Howe, but it seems best to go still farther than he has done and to recognize two distinct species, instead of trying to retain one of them as a variety of / Asagrayana. The first of these species, / Franciscana, resembles the eastern plant in its plane underleaves and in the median line of discolored cells in its lobes ; but the latter is a less striking feature than in 1 Asagrayana and the underleaves are different in shape and never auriculate at the base. The stylus, too, is usually reduced to a minute subulate process and, in the perichetial bracts, is replaced by a cluster of cilia, a similar cluster being found also at the antical base of the lobe.
The second of these two species, /! Californica, is usually more slender than any of our other Thiopsielle and its less imbricated leaves give it a somewhat looser appearance. In the involucre, the bracts rapidly increase in size; so that, while the outer ones are often smaller than the corresponding ones in / Asagrayana, the innermost bracts are larger: the bases of bracts and of bracteoles are much as in the eastern species, but the lobes of the bracts are proportionately narrower. The underleaves are variable; in rare cases they are slightly auriculate at the base, while their mar- gins are either plane or slightly reflexed on one or both sides near the base, never at the apex. The stylus of the leaves is minute, very much as in J. Franciscana, from which the present species differs in the usual absence of the median line of discolored cells, in the shape of its underleaves, and in the characters of its perichetial bracts and bracteoles,
A. W. EHvans—North American Species of Frullania. 29 Suscenus V.—DIASTOLOBA Spruce.
Key to the Species.
Bracts and bracteoles (at least those of the innermost row) strikingly dentate or spinose. Lobes of leaves marked by a line of discolored cells; lobules of bracts with a distinct segment or stylus on inner edge. 17. E Selwyniana,
Lobes of leaves without discolored cells; stylus of bracts not distinct. 19. #. Donnellii.
Bracts and bracteoles entire or nearly so. Lobules of leaves parallel with the stem. 18. 4. Kunzet.
Lobules of leaves widely spreading from the stem. 20. # Caroliniana.
17. Frullania Selwyniana Pearson, List of Canadian Hepatic, 7. pl. 7. 1890,
Frullania Sullivantie Aust., Bull. Torrey Bot. Club, iii: 16. 1872 (not # Sulli- vantii Aust.).
Frullania fragilifolia Aust., 1. c., vi: 301. 1879 (not Tayl.).
PuaTe XIII. figs, 9-17.
Autoicous : plants appressed to matrix, reddish-brown or purplish : stems irregularly pinnate: leaves imbricated, the lobe ovate, arching over the stem and cordate at base, somewhat decurved at the rounded apex, entire ; lobule close to the stem and subparallel with it, short- clavate ; stylus minute, subulate ; underleaves distant, rhombic-oval, bifid about one-third with obtuse lobes and sinus, entire or unidentate on the sides; leaf-cells of lobe thick-walled, trigones inconspicuous except in the middle and toward the base, intermediate thickenings scanty, discolored cells in a median line: @ inflorescence terminal on the stem or a main branch; bracts in about three pairs, unequally bifid, the lobe ovate, acute (becoming obtuse as we recede from the perianth), irregularly dentate, lobule narrower than the lobe, ovate, acute, irregularly ciliate-dentate and bearing a distinct, usually dentate segment or stylus below the middle of the inner edge ; bracteole free, broadly ovate, bifid to below the middle, with acute lobes and sinus, irregularly ciliate-dentate (becoming simply dentate on receding from the perianth); perianth about a third exserted, obcuneate, compressed on the sides, and with a strong postical keel, abruptly narrowed into a short, broad beak, minutely setulose at the
30 A, W. Evans—North American Species of Frullania.
mouth: ¢ spike globose, occupying the extremity of a short lateral branch near the perianth, bracts in about two pairs.
Stems 0°13"" in diameter ; lobes of leaves 0°50™™ long, 0°45™™ wide, lobules 0°23™™ long, 0°14"™ wide ; underleaves 0°25"™ long and wide ; lobes of branch-leaves 0°35™" long, 0°30™" wide; branch- underleaves 0°18™" long, 0°14™" wide ; leaf-cells at edge of lobe 0:015™", in the middle 0°018™™ in diameter, at the base 0:030"™ long 0:023™™" wide ; bract J, lobe 1.10™™ long, 0°50™™ wide, lobule 0°95™™ long, 0°40" wide; bracteole I, 0°85™™ long, 0:°70™™ wide; bract II, lobe 0°90"" long, 0°85™" wide, lobule 0°75"™ long, 0°25™™ wide ; bracteole II, 0:70™™ long, 0°40™™ wide; perianth 1:25™™ long, 0:90™™ wide.
On bark of trees (mostly the white cedar); near Urbana, Ohio (Sullivant, Miss Biddlecome) : Campaign County, Ohio (Werner): Ste. Anne’s River, Gaspé, Canada (Macoun). Distributed in Hep. Amer. n. 176.
Austin was apparently familiar with the characters of this distinct little species, for he has described them clearly and fully under his F. Sullivantie ; but, strangely enough, he afterwards considered them too unimportant to separate our plant from the European F. fragilifolia. Ina sterile condition, the two species certainly resem- ble each other closely ; both are reddish in color and branch in about the same way and both show discolored cells in their leaves. In F. fragilifolia, however, the stems are a little more slender than in F, Selwyniana, the underleaves are narrower, and the discolored cells are usually irregularly scattered instead of being arranged in a more ~ or less distinct line or group in the middle of the lobe. When inflo- rescence or perianths are present, /. Selwyniana can be at once dis- tinguished from the European plant by its antheridial spikes borne close to the involucre ; its perianth, too, is less exserted, its inner- most bracts are acute, and its bracteoles are free. . Selwyniana is- our only autoicous species with discolored cells in its leaves. I have retained for this species the name recently given to it by Mr. Pear- son, Austin’s older name being too like 1. Sullivantii.
18. Frullania Kunzei Lehm. et Lindenb. in G. L. et N. Syn. Hep., 449. 1845. Jungermannia Kunzet Lehm., et Lindenb. in Lehmann, Pugillus vi: 50. 1834. Frullania Drummondii Tayl., Lond, Jour. Bot., v: 401. 1846.
PuiaTE XIV. figs. 1-16.
Autoicous: plants closely appressed to matrix, reddish-brown, varying to deep blackish-purple in more exposed situations: sterile
A, W. Evans—North American Species of Frullania, 31
stems regularly once or twice pinnate, fertile stems more irreeularly pinnate: leaves contiguous or imbricated, the lobe orbicular-ovate, arching over the stem but not cordate at base, mostly plane at the rounded apex, entire: lobule close to the stem and subparallel with it, short-clavate ; stylus minute : underleaves distant, obovate, plane, bifid about one third with obtuse or subacute lobes and sinus, entire : leaf-cells of lobe thick-walled with inconspicuous trigones and no intermediate thickenings: @ inflorescence terminal on the stem or a main branch, bracts in three or four pairs, unequally bifid, the lobe ovate or oblong, rounded or obtuse and often apiculate at the apex, entire ; lobule narrower, ovate, subacute, bearing a minute tooth or stylus on the inner edge near the base, otherwise subentire ; brac- teole free or connate on one side, ovate, bitid to about the middle with subacute lobes and sinus, entire or slightly sinuous-dentate ; perianth more than half-exserted, compressed on the sides and el one-keeled postically, oblong, abruptly narrowed into a short beak, setulose at the mouth: ¢ spike globose, terminal on a short branch at some distance from the involucre, bracts in about _ two pairs.
Stems 0°10™™ in diameter ; lobes of leaves 0:°45™™ long, 0°35™™ _ wide, lobules 0°18" long, 0°10" wide; underleaves 0:20"™ long, ~0:17™" wide; lobes of branch-leaves 0°35™" long, 0:25™" wide; branch-underleaves 0:17™™ long, 0°10" wide; leaf-cells from edge g lobe 0°012™", from middle 0:016™™" in diameter, and from base 0-035" long, 0°022™™" wide ; bracts I, lobe 0°85"™™ long, 0°45™™ wide, ‘lobule 0:80™ long 0:30™™ wide ; bracteole I, 0°85™™ long, 0°50™™ wide; bract II, lobe 0°70™™ long, 0°45™™ wide, lobule 0°50" long, ) 0°25"™ wide ; bracteole II, 0-50™™ long, 0°35" wide ; perianth 1:25™™
long, 0°75™™ wide.
On bark of trees, or sometimes creeping over lichens; Florida to Louisiana. Distributed in Hep. Bor.-Amer. 2. 105d, and in Hep. — Amer. ». 101.
_ Frullania Kunzei was first described from Cuban material, but
~ does not seem to have been collected very often in the West Indies ; ‘in our Gulf States, on the contrary, it is apparently not uncommon. It is most closely related to the following species, 7. Donnellii, but also bears some resemblance to /. Selwyniana. From this latter
species it differs in its more regularly pinnate stems, in its narrower, less convex and homogeneous leaves, which are not cordate at the base, in its more exserted perianth with longer setulze at the mouth, and in its entire bracts.
Siar Oy ae
32 A, W. Evans—North American Species of Frullania.
19. Frullania Donnellii Aust., Bull. Torrey Bot. Club, iv: 301. 1879. PLATE XIV, figs. 17-29.
Autoicous: plants closely appressed to matrix, reddish-brown : sterile stems regularly once or twice pinnate, fertile stems more irregularly pinnate: leaves contiguous or imbricated, the lobe orbic- ular-ovate, arching over the stem but not cordate at base, plane or nearly so at the rounded apex, entire ; lobule separated from the stem by a little less than its width, subparallel with it or slightly oblique, short-clavate ; stylus minute: underleaves distant, oblong- obovate, plane, bifid about one third with obtuse or subacute lobes and sinus, entire: leaf-cells of lobe thick-walled with inconspicuous
trigones and no intermediate thickenings: @ inflorescence terminal —
on the stem or a main branch; bracts in two or three pairs, sub- equally bifid, the lobe ovate, acuminate, sharply and irregularly incised-dentate, lobule similar to the lobe but usually still more toothed, stylus not distinct ; bracteole free, bifid to or beyond the middle with segments like the lobules ; bracts and bracteoles becom- ing rapidly smaller and less toothed on receding from the perianth ; perianth about half-exserted, compressed on the sides and strongly one-keeled postically, oblong, abruptly narrowed into a short beak, setulose at the mouth; ¢@ spike globose, occupying a short branch close to the involucre, bracts in about two pairs.
Stems 0°10™™" in diameter; lobes of leaves 0°60" long and wide ; ~
lobules 0:19"" long, 0°12" wide ; underleaves 0°30™" long, 0°25™™ wide; lobes of branch-leaves 0°30™" long, 0°25™" wide ; branch-
underleaves 0°20" long, 0°10™ wide; leaf-cefls at edge of lobe
0:012™", in the middle 0-018™™ in diameter and at base 0°635"™ long, 0°018"" wide; bract I, lobe 0:90™™ long, 0°55™" wide, lobule 0°90"" long, 0°45™" wide; bracteole I, 0:'70™™ long and wide; bract II, lobe 0°75™" long, 0°45™" wide, lobule 0°60™™ long, 0°35™" wide; brac-
©"?
teole ike OOre long, O°5 5mm wide . perianth 1:290™m™ long, 0°70™™ wide.’
’
On trees; East Florida (J. Donnell Smith): Eustis, Florida (Underwood): Monticello, Florida (Lighthipe).
FE. Donnellii seems to be a rare species, but, as sterile material is usually indistinguishable from /. Aunzei, it has probably been over- looked. ‘The main difference between the two species is to be found in the perichtial bracts : in #! Awnzet these are entire or nearly so, while in # Donnellii the innermost ones at least are strongly incised-dentate.,
The first three species which are here placed in Déiastoloba are in some respects intermediate between that subgenus and Thiopsiella, agreeing with the latter mainly in having their lobules parallel with
a. + > aye
A, W. Evans—North American Species of Frullania. 33
the stem. Still, it has seemed better on the whole to group them with & Caroliniana, as they agree with it in the following impor- tant points : (1) the plane underleaves, (2) the autoicous inflorescence, and (3) the position of the 9 flowers.
20. Frullania Caroliniana Sulliv., Muse. Alleg. n. 270. 1846. Amer. Jour, Sci. and Arts, II. i: 74. 1846,
Frullania brunnea Aust., Hep. Bor.-Amer., 7. 105e. 1875 (not Spreng.). PLATE XV.
Autoicous : plants closely appressed to matrix, varying in color from yellowish-green to reddish-brown ; sterile stems mostly bipin- nate, fertile stems more irregularly branched: leaves closely imbri- cated, the lobe ovate, arching over the stem but not cordate at base, rounded and slightly decurved at the apex, entire; lobule distant from the stem and spreading at a wide angle (30°—40°), short-clavate; stylus a small, obliquely spreading triangular process: underleaves
_ contiguous or imbricated, orbicular, plane, bifid about one-half with
obtuse or subacute lobes and sinus, entire or vaguely unidentate on the sides ; branch-underleaves much narrower and with sharper points : leaf-cells of lobes sometimes thick-walled with inconspicuous trigones, sometimes thinner walled with conspicuous trigones and occasional intermediate thickenings: ¢@ inflorescence terminal on the stem or a main branch ; bracts in three or four pairs, unequally bifid, the lobe ovate, obtuse or apiculate at the apex, entire ; lobule shorter and
_ narrower, ovate, subacute, bearing on the inner edge near the base
a distinct tooth-like segment or stylus, otherwise entire ; bracteole free or connate on one side with bract, bifid about one-third with acute lobes and sinus, entire or nearly so; perianth exserted about one-third, obcuneate, abruptly narrowed into a short, setulose beak,
“
_ compressed on the sides and strongly unicarinate postically ; ¢ spike
globose, occupying a short lateral branch near the Q inflorescence, bracts in one or two pairs. Stems 0°10™" in diameter; lobes of leaves 0°70™™ long, 0°80"™
_ wide, lobule 0:20™" long, 0°10" wide; underleaves 0°40™™ long,
0:30"™ wide; lobes of branch-leaves 0°30" long, 0°40™™ wide; branch-
underleaves 0-20™" long and wide; leaf-cells from edge of lobe 0'017™™,
from middle 0:018™" in diameter, and from base 0:038™™ long, 0°023""'
wide; bract I, lobe 1°25™™ long, 0°75"™™" wide, lobule 0°80™™ long, 0-40™" wide; bracteole I, 0-80" long, 0°40" wide, bract II, lobe 0:90" long, 0°57™™ wide, lobule 0°60" long, 0°30" wide, bracteole II, 0°55™" long, 0.35™" wide; perianth 150™™ long, 1:20"™ wide.
Trans. Conn. ACAD., VOL. X. May, 1897. 3 ‘
34 A, W. Evans—North American Species of Frullania.
On trees; Florida to Louisiana. Distributed in Muse. Alleg. n. 270, in Hep. Bor.-Amer. n. 105e (as / brunnea), and in Hep. Amer. n. 84 (as F. brunned).
F. Caroliniana was referred by Austin to J’ brunnea, a plant found in southern Africa, Herr Stephani has kindly sent me a speci- men of this species from the type locality and I find it to differ from our plant in the following points:—the leaves are usually minutely apiculate, especially on the branches; the lobules spread still more widely, often forming more than a right angle with the stem; the stylus is smaller ; the underleaves are sharply spinose-dentate, those of the branches being sometimes almost laciniate. J Caroliniana is easily distinguished from all our other species by the distant and ~ widely spreading lobules of its leaves.
INCOMPLETELY KNOWN SPECIES. 21. Frullania Chilcootiensis Steph., Engler’s bot. Jahrb., viii: 98. 1886.
Dioicous : plants growing on bark with other hepatice, brownish: stems filiform, simple, 3-4™™ long; leaves. remote, the lobe broadly ovate, obtuse, concave, decurved at the apex, entire, arching over the stem; lobule a third as large as the lobe, galeate, constricted near the mouth, close to the stem or often obliquely incumbent ; stylus large, triangular-lanceolate, spreading: underleaves spreading, cuneate- obovate, bifid to about the middle, with a narrow obtuse sinus and connivent, obtuse lobes, angled or obtusely unidentate on the sides ; leaf-cells uniformly thickened, 0°010™" in diameter: perichzetial _ bracts in about two pairs, those of the innermost pair double the size of the stem-leaves, spreading, unequally bifid, the lobe oblong-ovate, obtuse, lobule half as broad as the lobe, acute, with a long cilium or stylus above the middle of the inner edge ; bracteole nearly as large as bracts, oblong, bifid about one-third with an open obtuse sinus and subacute lobes: perianth and ¢@ spike unknown.
On bark; Chilcoot, Alaska (Krause).
I have been unable to obtain specimens of this very minute species and have compiled the above account of the plant from the original description, aided by drawings and notes kindly sent me by Herr Stephani. The plant is apparently a Zrachycolea.
22 Frullania Wrightii Aust., Bull. Torrey Bot. Club, iii: 15, 1872 Dioicous: plants brownish : stems irregularly pinnate : leaves im-
bricated, the lobe orbicular, arching over the stem and cordate at
Or
A. W. Evans—North American Species of Frullania.
base, decurved at the rounded apex, entire ; lobule close to the stem, galeate, inflated, sometimes more distant and explanate; stylus minute, subulate: underleaves orbicular-obovate, bifid about one- fourth with obtuse or subacute lobes and sinus, entire or repand-den- tate on the margins; leaf-cells of lobe rather thick-walled with con- spicuous trigones and intermediate thickenings; @ inflorescence ter- minal on the stem or a main branch ; bracts in two or three pairs, unequally bifid, the lobe ovate, rounded at the apex, entire, lobule shorter and narrower, acute, bearing a small tooth or stylus on the inner edge below the middle, otherwise entire, bracteole free from the bracts, narrowly ovate, bifid about one-third with acute lobes and ‘sinus, subdentate toward base; perianth and ¢ plant not seen.
New Mexico (Wright).
Frullania Wrightii, known only from its type specimens, is appar- ently a form of Ff riparia. It is impossible to be sure of this, how- ever, until better developed plants are found, as there are slight differences between the two in areolation, underleaves, and involucre.
In conclusion I would express my thanks to Professor Underwood for the loan of his valuable collection of Frudlaniv, to Herr Stephani for helpful notes, drawings and specimens, to Dr. Robinson for ac- cess to several of Taylor’s and Sullivant’s types, and to Mr. Howe and others for specimens.
Yale University.
36 A, W. Evans—North American Species of Frullania.
EXPLANATION OF PLATES.
Each species is represented in natural size and with enlarged details: in Plates I and IX-XIV, the leaf-cells are enlarged 255, and the other details 28 diameters; in Plates II-VIII and XV, the leaf-cells are enlarged 290, and the other details 32 diameters.
PLATE I.
Frullania arietina Tayl.—Fig. 1, Plant, natural size.—Fig. 2. Part of plant, postical view, showing perianth and hypogynous perigonial bracts.—Fig. 3. Leaf-cells from middle of lobe.—Fig. 4. Bracts and connate bracteole I.—Fig. 5. Bracts and connate bracteole II.—Fig. 6. Transverse section of perianth. All the figures. _ from Florida specimens collected by Mr. Smith.
Frullania Oakesiana Aust.—Fig. 7. Plant, natural size.—Fig. 8. Part of plant, postical view, showing perianth and andrcecium.—Fig. 9. Part of stem with leaf, antical view.—Fig. 10. Leaf-cells from middle of lobe.—Fig. 11. Bract and connate bracteole I.—Fig. 12. Bract I.—Fig. 13. Bract Il.—Fig. 14. Bracteole II.—Fig. 15. Transverse section of perianth. All the figures from specimens collected by the author on Carter Dome, New Hampshire.
PLATE II.
Frullania Bolanderi Aust.—Fig. 1 Plants, natural size.—Fig. 2. Part of plant, postical view, showing perianth.—Fig. 3. Part of plant, postical view.—Fig. 4. Part of branch, ending in a flagellum.—Fig. 5. Part of stem, antical view.—Fig. 6. Leaf. cells from middle of lobe.—Fig. 7. Bracts with connate bracteole .—Fig. 8. Bract and connate bracteole I.—Fig. 9. Same, from another involucre.—Fig. 10. Bract and connate bracteole II, from same involucre as Fig. 9.—Fig. 1]. Bract and connate bracteole I.—Fig. 12. Bract I.—Fig. 13. Bract and connate bracteole II. —Fig. 14. Bract Il.—Fig. 15. Transverse section of perianth. Figs. 1-3 and 5— 8 from Californian specimens collected by Mr. Howe; Figs. 4, 9 and 10 from British Columbian specimens collected by Professor Macoun; and Figs. 11-15 from Californian specimens collected by Bolander.
PLATE Ill. :
Frullania inflata Gottsche.—Fig. 1. Plant, natural size.—Fig. 2. Part of plant, postical view, showing perianth and andreecium,—Fig. 3. Part of plant, postical view.— Fig. 4. Part of stem, antical view.—Fig. 5. Leaf-cells from middle of lobe.—Figs. 6, 7. Bracts IL—Fig. 8. Bracteole I.—Fig. 9. Bract II.—Fig. 10. Bracteole 11.— Figs. 11, 12. Transverse sections of perianths.—Fig. 13. Bract I.—Fig. 14. Bracteole I.—Fig. 15. Bract II.—Fig. 16. Bracteole IIl.—Figs. 17, 18. Transverse sections of perianths. Figs. 1, 3, 4, and 6-12 from specimens collected by Dr. Underwood at Austin, Texas; Figs. 2, 5, and 13-18 from Hep. Amer. n. 68.
PLATE LV.
Frullania Cataline Fivanus.—Fig. 1. Plants, natural size.—Fig. 2. Part of plant, postical view, showing perianth and androecium,—Fig. 3. Part of plant, postical view.—
A. W. Evans—North American Species of Frullania. 3
Fig. 4. Part of stem, antical view.—Fig. 5. Leaf-cells from middle of lobe —Fig, 6. Bract and connate bracteole I.—Fig 7. Bract I.—Fig. 8. Bract and connate bracteole II.—Fig. 9. Bract IIl.—Fig. 10. Bract and connate bracteole III.—Figs. 11, 12. Transverse sections of perianth. All the figures from the type specimens.
PLATE V.
Frullania riparia Hampe.—Fig. 1. Plants, natural size —Fig. 2. Part of plant, postical view, showing ¢ inflorescence.—Figs. 3, 4. Parts of plants, postical view.—Fig. 5. Part of stem, antical view.—Fig. 6. Leaf-cells from middle of lobe.—Figs. 7, 8. Bracts II.—Fig. 9. Bracteole II.—Fig. 10. Bract III.—Fig. 11. Bracteole III. Figs. 1, 4, and 5 from specimens collected by the author at Trumbull, Connecticut ; Figs. 2 and 6-11 from specimens collected by Dr. Underwood at Greencastle, In- diana; and Fig, 3 from specimens collected by Mr. Holzinger near Washington.
PLATE VI.
Frullania squarrosa (R. Bl et Nees) Dumort.—Fig. 1. Plant, natural size.—Fig. 2.
Part of plant, postical view, showing perianth —Fig. 3. Part of plant, postical view, showing young ¢ inflorescence.—Fig. 4. Part of stem antical view.—Fig. 5. Leaf-cells from middle of lobe.—Fig. 6. Bracts and connate bracteole I.—Fig. 7. Bract and connate bracteole I—Fig. 8. Bract and connate bracteole Il.—Fig. 9. Transverse section of perianth. Figs. 1, 2, and 5-9 from Florida specimens collected by Dr. Underwood; Figs. 3 and 4 from Tennessee specimens collected by Mr. Ruth.
Puate VII.
Frullania Brittonie Evans.—Fig. 1. Plants, natural size.—Fig. 2. Part of plant, pos- tical view, showing perianth.—Fig. 3. Part of plant, postical view, showing young andreecium.—Fig. 4. Part of stem, antical view.—Fig. 5. Leaf-cells from middle of lobe.—Figs. 6, 7. Bracts I.—Fig. 8. Bracteole IL.—Fig. 9. Bract II.—Fig. 10. Bracteole II.—Fig. 11. Bracteole I, from another involucre.—Fig. 12. Transverse section of perianth. All the figures, from specimens collected by the author at Meriden, Connecticut.
Frullania dilatata (L.) Dumort.—Fig. 13. Bract I.—Fig. 14. Bracteole I.—Fig. 15. Bracteole II. All the figures from Swedish specimens collected by Dr. Arnell.
PLATE VIII.
Frullania Virginica Gottsche.—Fig. 1. Plants, natural size.—Fig. 2. Part of plant, postical view, showing perianth.—Figs. 3, 4. Parts of plants, postical view.— Fig. 5. Part of stem, antical view.—Fig. 6. Leaf-cells from middle of lobe.—Fig. 7. Bracts and connate bracteole I.—Figs. 8, 9. Bracts II.—Fig. 10, Bracteole II. —Fig. 11. Bract III.—Fig. 12. Bracteole III.—Fig. 13. Bracts and connate bracteole I, from another involucre.—Fig. 14. Transverse section of perianth.—
_ Fig. 15. Bract 1—Fig. 16. Bracteole I.—Fig. 17. Bract Il.—Fig. 18. Bracteole II.—Fig. 19. Transverse section of perianth. Figs. 1-3 and 5-14 from specimens collected at Auburn, Alabama, by Dr. Underwood; Figs. 4 and 15-19 from speci- mens collected at Wilmington, Delaware, by Mr. Commons.
38 A, W. Evans—North American Species of Frullania. Y/
PLATE IX.
Frullania Eboracensis Gottsche —Fig. 1. Plants, natural size.—Fig. 2. Part of plant, postical view, showing perianth; the innermost bract on the left is abnormal in having its stylus about as large as its lobule.—Fig. 3. Part of stem, antical view. —Fig. 4. Leaf-cells from middle of lobe.—Fig. 5. Bracteole I.—Fig. 6. Transverse section of perianth.—Fig. 7. Bract and connate bracteole I.—Fig. 8. Bract I.— Fig. 9. Bract If.—Fig. 10. Bracteole Il-—Fig. 11. Transverse section of peri- anth. Figs. 1-3 from specimens collected by Dr. Jelliffe at Lake George, New York; Figs. 4-6 from specimens collected by Miss Vail in the Catskills, New York; Figs. 7-11 from specimens collected by Mr. Commons at Wilmington, Delaware.
Frullania plana Sulliv.—Fig. 12. Plaut, natural size.—Fig. 13. Part of plant, postical view, showing perianth and androecium.—Fig. 14, Part of stem, antical view.— - Fig. 15. Leaf-cells from middle of lobe— Figs. 16, 17. Bracts I.—Fig. 18. Bracteole 1.—Fig. 19. Bract IJ.—Fig. 20. Bracteole II.—Fig. 21. Transverse section of perianth. All the figures from Hep. Bor.-Amer. 7. 102.
PLATE X. ;
Frullania Nisquallensis Sulliv.—Fig. 1. Plant, natural size.—Fig. 2. Part of plant, postieal view, showing perianth.—Fig. 3. Part of stem, postical view, underleayes dissected away.—Fig. 4. Part of stem, antical view.—Fig. 5. Leaf-cells from mid- dle of lobe.-—Fig. 6. Bracts and connate bracteole I.—Fig. 7. Bract Il.—Fig. 8. Bracteole II.—Fig. 9. Bract III.—Fig. 10. Bracteole I1].—Fig. 11. Transverse section of perianth. All the figures from specimens collected at Anacortes, Washington, by Mrs. F. K. Sears.
PLATE XI.
Frullania Asagrayana Mont.—Fig. 1. Plant, natural size.—Fig. 2. Part of plant, postical view, showing perianth.—Fig. 3. Part of plant, antical view, showing ~ androecium.—Fig, 4. Part of stem, postical view, underleaves dissetted away.— Fig. 5. Leaf-cells from middle of lobe.—Fig. 6. Bract I.—Fig. 7. Bracteole I.— Fig. 8. Bract II.—Fig. 9. Bracteole II].—Fig. 10. Bract III.—Fig. 11. Bracteole IlI.—Fig. 12. Transverse section of perianth.—Figs. 13, 14. Bracts I.—Fig. 15. Bracteole I.—Figs, 16, 17. Bracts II.—Fig. 18. Bracteole IIl.—Fig. 19. Subinyo-_ lucral leafi—Fig. 20. Subinvolucral underleaf.—Fig. 21. Bract and connate bracteole I.—Fig. 22. Bract I. Figs. 1-12 from specimens collected by Mrs. Britton on White Top, Virginia; Figs. 13-20 from specimens collected by Mr. Waite at Washington, D. C.; Figs. 21 and 22 from specimens collected by Mr. Ruth at Knoxville, Tennessee.
PLATE XII,
Frullania Tamarisci (L.) Dumort.—Fig. 1. Plant, natural size.—Fig. 2. Part of plant, postical view.—Fig. 3. Part of stem, antical view.—Fig. 4. Leaf-cells from middle of lobe.—Fig. 5. Bract I.—Fig. 6. Bracteole I—Fig. 7. Bract II.—Fig. 8 Bracteole Il.—Fig, 9. Bract III.—Fig. 10. Bracteole III. Figs. 1-4 from New foundland specimens collected by Mr. Waghorne ; Figs, 5-10 from Swedish speci- mens collected by Dr. Arnell.
teh eae Bile alte
or
A, W. Evans—North American Species of Frullania. 39
Frullania Californica (Aust.) Evans.—Fig. 11. Plant, natural size.—Fig. 12. Part of
plant, postical view, showing perianth.—Fig. 13. Part of stem, antical view.— Fig. 14. Leaf-cells from middle of lobe.—Fig. 15. Bract and connate bracteole I. —Fig. 16. Bract I.—Figs. 17, 18. Bracts Il.—Fig. 19. Bracteole II.—Fig. 20. Bract I11.—Fig. 21. Bracteole III.—Fig. 22. Transverse section of perianth. All the figures from specimens collected at Seattle, Washington, by Mr. Piper.
PLATE XIII.
Frullania Franciscana Howe.—Fig. 1. Plant, natural size.—Fig. 2. Part of plant,
postical view, showing perianth.—Fig. 3. Part of stem, antical view.—Fig. 4. Leaf-cells from middle of leaf.—Fig. 5. Bract and connate bracteole I.—Fig. 6. Bract I.—Fig. 7. Bract I].—Fig. 8. Transverse section of perianth. All the figures from Californian specimens collected by Mr. Howe.
Frullania Selwyniana Pearson.—Fig. 9. Plants, natural size.—Fig. 10. Part of plant,
postical view, showing perianth and andrcecium.—Fig. 11. Part of plant, postical view.—Fig. 12. Part of stem, antical view.—Fig. 13. Leaf-cells from middle of lobe.—Fig. 14. Bract I.—Fig. 15. Bracteole I.—Fig. 16. Bract II.—Fig. 17. Transverse section of perianth. All the figures from Ohio specimens: Figs. 9, 10 and 14-17 from specimens collected by Mr. Wilcox; the others from speci- mens collected by Mr. Werner.
PLATE XIV.
Frullania Kunzei Lehm. et Lindenb.—Fig. 1. Plant, natural size.—Fig. 2. Part of plant,
postical view, showing perianth, androecia, and 2 inflorescence.—Fig. 3. Part of stem, antical view.—Fig. 4. Leaf-cells from middle of lobe.—Figs. 5, 6. Bracts I.
—Fig. 7. Bracteole I.—Figs. 8, 9. Bracts II.—Fig. 10. Bracteole Il —Fig. 1}. - Bract 11J.—Fig. 12. Bracteole I1I.—Fig. 13. Bract and connate bracteole I.—
Fig. 14. Bract I.—Fig. 15. Transverse section of perianth.—Fig. 16. Setule from mouth of perianth. All the figures from specimens collected by Dr. Underwood : Figs. 13, 14 and 16 from Florida specimens and the others from specimens collected at Ocean Spring, Mississippi.
Frullania Donnellii Aust.—Fig. 17. Plant, natural size.—Fig. 18. Part of plant, postical
i
view, showing perianth and androecium.—Fig. 19. Part of stem, antical view.— Fig. 20. Leaf-cells from middle of lobe.—Figs. 21, 22. Bracts I—Fig. 23. Bracteole I.— Figs. 24, 25. Bracts II.—Fig. 26. Bracteole I].—Fig. 27. Bract I1I.—Fig. 28. Bracteole III].—Fig. 29. Transverse section of perianth. All the figures from specimens collected at Eustis, Florida, by Dr. Underwood.
PLATE XV.
_ Frullania Caroliniana Sulliv.—Fig. 1. Plant, natural size.—Fig. 2. Part of plant,
postical view, showing perianth and andreecia.—Fig. 3. Part of stem, antical view. Fig. 4. Leaf-cells from middle of lobe.—Fig. 5. Bract and connate bracteole I.— Fig. 6. Bract I.—Figs. 7, 8. Bracts Il.—Fig. 9. Bracteole Il.—Fig. 10. Bract III. —Fig. 11. Bracteole III.—Fig. 12. Transverse section of perianth. All the fig- ures from specimens collected by Mr. Langlois in Louisiana.
sie Fi A
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II.—A Strupy or THE FAMILY PEcTINID#, witH A REVISION oF THE GENERA AND SuBGENERA. (Six pirates.) By A. E. VERRILL.
Tue classification of the Pectinide must, for the present at least, be based mainly upon the characters of the shells, for the soft parts have been carefully studied only in a few of the very numerous spe- cies. There is good reason to believe that in some cases good gen- eric characters may be found in the structure of the foot, but this organ is so contractile that alcoholic specimens give but a poor idea of its form in life. The palpi and gills are known to afford import- ant characters in some species, but they have been studied in only a few genera. (See Plates xx, xx.) But the relations of the shell to the soft parts are so very intimate that the form and structure of the shell may be taken as an expression of the modifications of some of the important internal parts, and therefore must give valid evidence of generic modifications.
In this family the use of shell-characters for generic and subgen- eric groups has this great advantage that it will enable us to classify the vast number of fossil species, which are far more numerous than the living ones, and to compare those of successive geological periods, group by group, with each other and with modern groups. In this way we may be better able to trace the lines of evolution. Until the minor modifications of structure, such as characterize subgenera and “ sections ” of genera in this and other bivalve families, are duly considered, no great progress can be made in the study of their evo- lution.
It is very essential that students of Mollusca should become im- pressed with the idea that even the slightest modification of the form or structures of the shell, if persistent, has its meaning, and some distinct value in progressive evolution, whether we may be able to discover it or not. Even the color is often of protective value. For instance, the reddish and brown colors of Chlamys Islandica matches the colors of the red nullipore that covers the stony bottoms where it is usually found, and C. irradians has generally a gray color, similar to that. of the sandy bottoms, which it mostly fre- quents. Instances of protective coloration in the shells of gastropods are very numerous,
42 A. E. Verrill—Study of the family Pectinide.
In the Pectinidie we can appreciate the value of some of the mod- ifications of the shell, on purely mechanical principles. Others are known to be correlated with the habits of the species. Thus the strong radial ribs or corrugations, found on species living in shallow water, serve to give their shells great strength, while the interlock- ings of the ends of the ribs, either in the form of marginal points or scallops, serve to keep the valves in exact apposition when closed, and therefore compensate for the absence of cardinal teeth. Such corrugations and marginal projections are generally lacking in the deep-sea species that are not exposed to the action of the waves. The special, internal, radial ribs of Amusium and allied genera also serve to greatly strengthen the thin, smooth shells of this group and — enable the valves to resist the action of the powerful adductor mus- cle in the act of sudden contraction for the purpose of swimming. In this group, the very compressed, round and polished shell indi- cates an adaptation to very active swimming habits, for such a shell gives the least friction in the singular manner in which these bivalves swim. Our large native scallop (C. Clintonius or Magellanicus) has a similar form and is remarkable for its swimming powers, even when of large size.
It seems difficult to explain satisfactorily why Amusium, and forms like C. Clintonius, should have a simple, thin. margin, without interlocking points or scallops, and often with the shell incapable of. closing tightly. It is possible that many such simple-edged and gap- ing shells have descended from those with radial ribs and interlock- ing scallops, for many of the ancient mesozoic fossil species are thus | defended. It is true that most (but not all) of the species that have no radial corrugations, and no marginal scallops, are from deep water, where they are not exposed to the rough action of waves and currents. Still they are, even there, exposed to the attacks of various crabs and fishes, against which strong interlocking valves ~ would be an obvious advantage. It seems probable that the increased lightness of the shell, by facilitating rapid swimming, may more than compensate for the loss of the power of passive resistance. This might well be the case whenever their principal enemies are sluggish animals, like drilling gastropods and voracious starfishes, for actively swimming Pectens could easily escape from such enemies, while their heavier and more sendentary relatives, especially those attached either directly or by a strong and persistent byssus, might be unable to escape.
Experience in the extensive cultivation of oysters, on the American
AL. EE. Verrill—Study of the family Pectinide. 43
coast, and in other parts of the world, shows that starfishes and the drilling gastropods are by far the most destructive enemies of those bivalyes. On our shores vast numbers of drilled oyster shells can be found almost everywhere, but drilled shells of our common scallop (Chlamys irradians), which is an active swimmer, are comparatively rare. Therefore it is probable that the gradual loss of radial ribs and corrugations, or their failure to develop in certain genera, is due to natural selection, in consequence of the advantage gained by the lighter shells for swimming purposes, in escaping from these slug- gish enemies.
Concentric ribs and undulations, found on some very thin shells, serve to stiffen and strengthen the shell against transverse strains, but they tend, also, to facilitate the tight closing of the valves at the simple and thin margins, for they permit a certain degree of flexi- bility of the thin shell, parallel with the margin. This kind of clo- sure is very obvious in Propeamusium and Cyclopecten, which include many small, thin, deep-water forms. In some cases the clo- sure is still farther perfected by a flattened or bevelled margin,
Most members of the family,if not all, form a byssus while young, for attachment, but they release themselves very easily and swim actively away. Many large and thick species seem to lose the habit entirely at maturity, and to rest unattached upon the bottom. But some small and delicate species, although capable of active swim- ming, appear to live attached much of the time through life. This is the case with Camptonectes or Palliolum vitrea and some allied deep-water species, which attach themselves to the branches of gor- gonians, corals, and hydroids, and thus gain protection from their enemies. The presence of a byssus is, however, consistent with the most active swimming powers. (See remarks under Cyclopecten, Bp. 71):
The extreme inequality of the valves in typical Pecten (=Janira) is a singular character, of ancient origin, for it was fully, devel- oped in many mesozoic species, closely allied to modern forms. It is the more strange, because, in most of the other groups having unequal valves, it is the under, or right, valve that is the flattest, but in true Pectens the right valve is strongly convex, while the left or upper valve is flat, or even concave externally, and usually shuts inside the margin of the lower valve like a lid. Both valves are thick and strongly ribbed. Probably this shape is advantageous when the shell is resting upon the bottom, with the lower valve partly buried in the sand and gravel around it, but not attached by a byssus, for
44 A, BE. Verrill—Study of the family Pectinide.
then there would be but little surface presented to the waves. The species of this group usually live in rather shallow water, within the limits of wave-action. It is usual for oysters and other attached forms to have the attached valve deepest. When the shells of these one-sided Pectens and others of similar form are dropped into the water they generally sink with the flat valve uppermost, so that this form may be useful in keeping them “right side down,” now that they have acquired a structure that requires them to lie on the right side, but this will not explain the first origin of the form, which was probably due to the gradual loss of swimming habits.
These one-sided Pectens seem to be rather sessile, as compared
with most of the other groups, and certainly the great thickness and —
weight of the shell, and its special form, do not seem to be adapted to active swimming habits. On the other hand, the byssal organs for attachment are not very well developed, so that the adult shells probably rest upon the bottom unattached, and move about by means of the foot.
That they do not have the swimming habit well developed is also indicated by the unusually tight closure of the valves at the base of the auricles, where the expulsion of the water takes place during the act of swimming, in this family. The right valve is strongly excurved at the byssal notch, and the left valve is strongly bent inward at the corresponding part, so as to fit the marginal notch very completely, leaving at most only a narrow passage for byssal threads. The form of these shells is poorly adapted for swimming, for if cur-
rents of water should be expelled in the usual way, from the sub- |
auricular margins, the currents would naturally be forced upward, and out of the concave lower valve, and thus the reaction would be strongly downward, so that the shell would not be raised from the bottom,
In those species that are able to rise to the surface and swim
actively about, the left or upper valve is always the more convex, and therefore the expelled currents of water must be directed more or less downward, so that the reaction forces the shell upward as well as backward.
In the typical one-sided Pectens the foot is pretty well developed, and there is a strongly marked and usually double scar on the left valve, where the pedal muscle is attached, above the scar of the adductor muscle. It is probable, therefore, that they can use the foot effectively for moving about, even when adult. ;
A, FE. Verrili—Study of the family Pectinide. 45
Remarks on the Ontogeny and Phylogeny of the existing genera of Pectinide.
It is not my intention to discuss the special phylogeny of the fam- ily, as a whole, at this time, for I have not at hand a sufficient num- ber of the palzozoic genera, in good preservation. This subject has been discussed, to some extent, by Dr. Jackson’ and others. It is certain that as we go back to the Paleozoic forms, the Pectinide, Pernide, and other related families gradually converge, and give evidence of a common ancestry. Dr. Jackson has suggested that a Nucula-like genus must have been the common ancestral form from which many families of Bivalvia, including Pectinid, Aviculide, etc., were derived. This view, which has been adopted by. Prof. Hyatt,* seems to me to lack demonstration and to be improbable, for Nucula, although an ancient genus, dating from the Paleozoic, is a rather highly specialized form, as to its foot, palpi, and some other parts, while its hinge-teeth are far more specialized than those of Cu- cullea, and many other early Paleozoic forms. The veliger-stages of Mytilus, Nucula, or of Perna present more nearly the forms and characters which I believe pertained to the earliest bivalves than any that we yet know from adult forms. But the early veliger-shell of Pectinidz is similar, and perhaps nearly as primi- tive in its characters. (See pl. xviii, figs. 1, 12,13.) In the most ancient allied types, we should, therefore, look for thin, delicate, ovate shells, without any differentiated hinge-teeth. A small in- ternal cartilage, or resilium, was probably coexistent with an ex- ternal ligament not differentiated from the general perisostracum, and connected directly with the resilium. The latter is formed by the invaginated cells of the primitive shell-gland of the veliger, and the perisostracum by the cells of the general ectodermic membrane, which is continuous with the shell-gland at first; therefore, the two structures must have been continuous, primarily, and probably of simultaneous formation. The greater part of the early bivalves have the resilium® and ligament both developed, but in many of the more modern genera, and also in some palwozoic genera, one or the
1 Dr. R. T. Jackson, Phylogeny of the Pelecypoda, Mem, Boston Soc, Nat. Hist., iv, No. 8, p. 277. 1890. »? Science, vol. v, p. 166, Jan. 29, 1897.
8 The useful term resilum was proposed by Dr. Dall, for the so-called internal car- tilage of the hinge of bivalves; resilial pit may replace ‘ cartilage pit.”
46 A. E. Verrill—Study of the family Pectinide.
other often becomes obsolete in the adult. The Pectinidz all retain both these structures in a decidedly primitive form. The ligament is thin and but little differentiated, and occupies a straight, narrow marginal groove along the whole of the hinge-line. The resilium is wedge-shaped or triangular, nearly central, with its apex joining the ligament. This seems also to be nearly the condition in the youngest shells of this group that I have been able to examine. The very early stages figured by Dr. Jackson seem to indicate the same thing.
The post-veliger shells, “ nepionic stage,” in all the genera that I have examined, are nearly smooth, and have, at first, only slightly — angulated dorsal margins, indicating the origin of the auricles, but the auricles develop rapidly and soon become sufficiently evi-— dent, the anterior one developing more rapidly and showing a byssal notch very early. In some species, like C. vitrea, the posterior auricle always retains its early undeveloped form.
In most species the form of the shell, characteristic of the family, is developed before the larval shell or “spat” is 1™™ in diameter, When. about 13 to 2™™ in diameter the characteristic sculpture usu- ally begins to appear. In most forms this consists at first of a number of small, straight radial riblets on both valves. These may or may not be accompanied by a peculiar, divergent vermiculation or “camptonectes-sculpture.” When the latter appears at all, it either slightly precedes, or is simultaneous with, the radial riblets.
A few species, like P. simile (PI. xvii, figs. 8, 8a), do not distinctly develop either of these forms of radial sculpture at any stage, but seem to retain through life the simple condition of the smooth or : radially striated larval shel], as well as much of its form, with little alteration. Others retain the ‘“ camptonectes-sculpture” and fine riblets without much change (C. striata). Hence we may conclude that in such groups as Hyalopecten, Palliolum, and Camptonectes we have survivals of very primitive or archaic Pectinide. This is also™ indicated by the very feeble differentiation of the posterior auricle, so noticeable in C. vitrea (Pl. xviii, fig. 6, 7, 10, 11) and in P. simile (Pl. xvii, fig. 8). In these species the posterior auricle’ retains the form that it has in the young spat of Chlamys irraatene (Pl. xx, fig. 3), and other more highly specialized species.
It is possible that the power of swimming, so well developed in young Pectinide, was acquired by the early fossil forms, even in the palwozoic ages. Possibly some of the early small forms of this group developed the power of swimming by the sudden closure of the shell before they entirely lost their ciliated velum, so that they
A, E. Verrill—Study of the family Pectinide. 177
may have retained, more completely than at present, a free-swim- ming or pelagic life. There may, very Jikely, have been small forms that retained the velum through life and used the valvular method of swimming when a more rapid motion ‘than the action of cilia could give was required, in order to avoid enemies.
According to Dr. Jackson’s observations on the young of (. érra- dians, the spat, at first, creeps about with its foot before it is able to
swim by the valvular method, but even in that stage the byssal
groove is present.’ (PI. xx, figs. 1, 2.)
In any case, it is probable that the first form of bivalve foot to be
developed, in the later veliger-stages of primitive bivalves, would have been a simple foot adapted to adhesion to floating objects or to stationary alg, etc., and not a foot adapted to creeping over the muddy bottom as Mr. Jackson has assumed, when considering the Nucula-like forms as the most primitive of bivalves. _ We may suppose that the earliest form of adhesion was temporary, and merely for the purpose of rest during the veliger condition, and it may have been effected by means of the mere surface adhesion of a little specialized, soft, fleshy or tongue-shaped foot, aided, perhaps, by a secretion of mucus from the surface. Such a mode of adhesion to objects is common among planarians, small nemerteans, annelids, and the young forms of many groups, at the present time.
From such a primitive adhesive foot the transition to a larger foot with more specialized cells situated in a groove for the secretion of stronger byssus-like threads of mucus would have been easy.
Such threads of adhesive mucus are formed by the foot glands of many land slugs and by certain marine species at the present time (e. g. Litiopa bombyx, a small gastropod that attaches itself to float-
ing sargassum in this way).
From this structure of foot the transition would have been easy
1 The form of this foot is like that of Mytilide, in which the foot is used for climb-
_ ing about and forming a byssus.
Dr. Jackson states that his youngest spat were not attached by a byssus, but crept
about by means of the long, ligulate, grooved foot, and seemed incapable of swim-
ming. He also observed that the spat could use the marginal tentacles for creeping about and clinging to objects. Those spat that I have observed (apparently quite as young) were capable of attaching themselves by a byssus, and when slightly older were seen to swim. When kept in stili water in vessels, such spat may be much
_ more inactive than when living in open waters, in coustant motion. The form of the
foot of the young spat, as described by Dr. Jackson, is better adapted for climbing over and adhering to sea-weeds than to creeping on the bottom, and requires far less
_ Specialization than does a disk-hke or flattened muscular foot for creeping purposes.
48 A. BE. Verrill—Study of the family Pectinide.
to a larger foot, accompanied by a muscular development for creeping about, and the formation of a definite byssal gland and groove for more secure, but temporary, attachment.
But the power of forming such a byssus does not imply a loss of swimming habits, for we find that, at the present time, many of
those species that can swim most rapidly have also the power of
forming a byssus very quickly when they wish to rest, by attaching themselves to seaweeds, etc. (See notes on Cyclopecten orbicularis, p. 72.) These two coincident habits are particularly noteworthy in the case of the smaller and more active forms of Pectinide, such as Palliolum and Cyclopecten, and in the young of the larger forms,
such as C, irradians. This fact tends to confirm the conclusion
that the early Pectinide had similar habits.’ The development of large, strong, or thick, ribbed and fluted shells took place later in geological time and was undoubtedly accompanied by more or less loss of swimming powers, just as the young of such species at the pres- ent time lose more or less of their swimming habits as they grow older and deyelop thicker and strongly ribbed shells. ‘his loss of swimming power may, or may not, be accompanied by a loss of the power of forming a byssus. In some cases, like Hinnites and Hemi- pecten, it is followed by a permanent attachment of the shell toa solid object. The true Pectens seem to lose the byssal organs
when adult, and to depend upon the weight and the form of the —
shell for safety. In general, those species that are best specialized for swimming
have a broadly rounded, symmetrical, and compressed shell, fre-_
quently with thin, nearly smooth valves, but generally strengthened by corrugations, undulations, external radial ribs, or internal lire or flutings.
Species that swim but little when adult often have a high and narrow form, with the auricles oblique and usually unequal, and the byssal notch is often highly developed, while the shell itself may become oblique and unsymmetrical, or heavy and thick, with strong ribs and grooves.
Most species swim well when quite small, but many lose this
‘The existence of a small gaping of the margins below the auricles at each end, seems to be due to the swimming habit, for the jets of matter are mainly ejected
from these places (see description by Dr. Jackson). But it is difficult to ascertain ~
whether the fossil species were gaping at these places or not. In many hving species the gaping is very slight, but in those that are active swimmers it is often considera- ble. (See C, opercularis, C, Clintonius, Amusium.)
"
ts
: mw Botce rd ee ree
ars
A, E. Verrill—Study of the family Pectinida. 49
power, more or less, when adult, especially in the case of the one- sided Pectens and those species that have very oblique shells, like certain species of Chlamys, for example C. madreporarum, which lives, when adult, in cavities between the branches of corals from which it cannot escape. This shell becomes very oblique and has very unequal auricles. HWinnites has gone farther in this direction, and has become attached and irregular when adult.
C. Clintonius, when it becomes very large, seems to be nearly sedentary and is often heavily covered with barnacles, ascidians, bryozoa, etc., but I have never found it with a byssus when adult. It probably retains more or less power of swimming through life, and seems to be migratory in its habits, changing its station accord- ing to the seasons. The same is true of Chlamys irradians and its
allies. But C. Clintonius, like Amusium, has the foot rather large
and divided at the top into two lobes by a deep fissure, so that the lobes can be spread apart in the form of a terminal disk. It is prob- ably used as a pushing organ.’ By means of this organ these species can probably push themselves slowly about when necessary to change their positions.
The teleological reasons for the development of ribs, flutings, and other forms of sculpture on the shells have been discussed elsewhere (pp. 42, 43). It is only necessary to state here that as the strongly ribbed and fluted conditions naturally and necessarily succeed the simple and fine ribbed stages, during growth, so in geological time
the strongly ribbed genera succeeded the simpler and thinner forms,
though many of the latter persist at the present time. But still, strongly ribbed forms of Pecten, Chlamys, Neithea, etc., had already become well-developed in mesozoic times, and Lyropecten, remarka- ble for the strength of the ribs, appeared early in tertiary time. But some of the modern, thin, smooth forms have probably descended from ribbed species, by the gradual reduction of the ribs.’ It is doubtful whether any generic or subgeneric group of Pectinide has been evolved since the Eocene. Even in the Cretaceous, nearly all the existing generic and sectional groups were in existence, together with a few that are now extinct.
1 Dr. Dall has described this organ as a terminal ‘‘sucker,” but it is doubtful if it can be used for adhesion.
2 This appears to be the case with Placopecten and Lissopecten, in which the ribs have become nearly obsolete, though they are in other respects nearly allied to Chlamys.
Trans. Conn. ACAD., Vou. X. JUNE, 1897. 4 bs
50 A, E. Verrill—Study of the family Pectinide.
In many respects, Amusiuvm seems to be one of the most special- ized groups. It may have descended from ribbed species of earlier ages. When adult the byssal organs are obsolete and the highly modified foot, with its large, terminal, concave disk, serves as a pushing organ, and perhaps for creeping about. The auricles and byssal notch are degenerate in form, while the internal strengthening ribs or lire are peculiar, secondarily acquired features, due to the special development of the thin shell for active swimming through life.
Synopsis of the principal characters available for the classification of Pectinide.
Shell.—The form may be broadly rounded, or high and narrow ; oblique, or upright; the texture may be thin and hyaline, or thick and opaque. Sometimes a somewhat prismatic or partly pearly structure appears on the inside, but the inner surface is generally porcellaneous. 5
The valves may be nearly equal, or one valve (either right or left) may be less convex, or even flat; they may gape widely at both ends, or close almost completely; the margin may be simple and thin and meet evenly, or the edge of the lower valve may bend up against the upper, or it may be bevelled ; more frequently the margins are scalloped and interlock. In some cases (Hinnites) the right valve becomes cemented to foreign objects and irregular in form, when adult.
Auricles—These may be small or large; straight or oblique; —
prominent and angular, or poorly developed and obtuse*; equal or unequal. The ends gape or flare apart, more or less, to contain the pallial tentacles and ocelli, The posterior auricles are sometimes nearly obsolete.
Byssal notch.—This may be deep, or shallow, or even obsolete (as 7 in Amusium and Pallium). As its margin grows it often leaves a “fasciole,” indicated by lines of growth, behind it.
Pectinidial teeth—These may be strong or feeble; few or many ; sometimes they are absent, especially in adult shells of large size. (Pl. xvi, fig. 9, p.)
Hinge-plate.—This may be thick or thin; broad or narrow; plain, or bearing longitudinal ribs, or oblique teeth-like processes (pl. xxi, fig. 4).
Cardinal ribs.—These are longitudinal ribs or folds, either nearly parallel with the hinge-margin or somewhat oblique ; or they may
A. E. Verrili—Study of the family Pectinide, 5]
become nearly transverse and tooth-like, alternating with pits (Pa/- lium).
The number of cardinal ribs may be from one to three on each end of the hinge-plate; most frequently there are one or two, the upper one forming the inner boundary of the ligamental groove, the second one somewhat divergent from the first. (Pl. xvi, fig. 6, ¢.)
Cross lines or incisions.—The upper ribs, and sometimes the other eardinal ribs, are generally crossed by numerous fine transverse grooves or incisions, alternating with ridges of about the same width. ‘These cross-lines may be straight and regular or they may be crooked or vermiculate. They are generally more distinct in the young shell than in the adult (pl. xvi, figs. 6, 9, 12 a, 7; pl. xix, figs. 1, 2,7). In certain extinct early genera they were much more con- spicuous than in any existing forms ( Neithea, Crenipecten).
Auricular crure.—These are divergent raised ribs or faint ridges running along the inner margins of the auricles (pl. xix, figs. 6-9). Sometimes they terminate distally in a rounded denticle (pl. xvi, fig. 9, d); sometimes the denticle alone is present. They are often obso- lete in the thick-shelled species, but are sometimes conspicuous struc-
— tures (Amusium).
Resilium.—This is generally nearly central and triangular or wedge-shaped. The surfaces next the shell are often calcified.
Resilial pit or Chondrophore.—This may be excavated entirely within the outline of the hinge-plate, or it may project considerably below it. In typical Pecten the pits are unlike in the two valves. Bea xvi, figs. 6,9, 7, pl. xxi, figs. 2, 2a, r.)
Ligamental groove.—This is always narrow, submarginal, and ex- tends along the whole length of the hinge-line. The elongation of the auricles serves to give it greater extension and importance; the exterior margin of one valve is often curved inward, partly over the groove. (PI. xvi, figs. 6, 9,7, pl. xxi, fig. 2, 1.)
Muscular scars.—The scars left by the adductor and pedal muscles often show marked differences that are, perhaps, of generic value, but unfortunately they are often very indistinct, and in the smaller species nearly or quite invisible. The scars differ considerably in
the two valves, for the pedal retractors are lacking in the right
valve. The pallial line is very simple. (PI. xxi, figs. 2, 2a.) Internal ribs and lire.—The inner surface of the shell may be per-
_ fectly smooth in some of the small plain species, but in Amusium _ and some other groups special raised radial ribs, often opaque white in
color, are developed independently of any external sculpture (pl. xix,
52 A. EF. Verrill—Study of the family Pectinide.
figs. 8, 9). Generally the inner surface is marked by ribs or flutings corresponding to the larger external grooves, but the internal ribs are usually bicarinate or double, especially near the margin, owing to a special thickening along each of their margins, which renders them more angular than the external gooves (pl. xvi, figs. 7, 10).
External sculpture.—The surface may be smooth, or it may be covered with sculpture of many kinds. It may differ on the two valves, or not. The most common sculpture consists of numerous strong radial ribs and grooves, alternating on the two valves, so that their ends interlock at the margin. The primary ribs as well as the grooves may be covered with smaller ribs, or by scales or spines of various kinds. The primary ribs may not increase in number with — age, but become continually broader (Pecten), or new ribs may be introduced between them, so that the larger ribs become more and more numerous without increasing in size (Chiwmys). Concentric sculpture may be developed on one or both valves, either with or without radial ribs. The whole surface may be evenly cancellated. There may be regular concentric undulations (JZyalopecten). The sur- face may be covered with a fine, divergent, vermiculated structure often described as “camptonectes sculpture” (pl. xvi, fig. 7, and xviii, fig. 14a); this may coexist with radial ribs.
Gills.—There are usually two pairs of normal fillibranchiate gills (see pl. xx, fig. 6, gy), but according to Dr. Dall, in at least one deep- water species (Paramusium Dalli), there is only a single pair of gills.
Foot.—The foot shows considerable variation in form. It gener- ally has a well-developed byssal groove, and usually a more or less developed terminal slit, which is often so large that the end of the foot can be expanded into a disk-like form (pl. xx, fig. 8). The grooved side of the foot is turned obliquely downward to the right (pl. xx, fig. 6, 7). The pedal retractor muscle is usually developed only on the deft side, so that its scar is lacking on the right valve.
Palpi.—These are generally large and broadly triangular organs, strongly fluted on the apposed surfaces. (PI. xxi, figs. 1, 2, d, d’.)
Labial tentacles—These are usually much-branched organs sur- rounding the mouth or situated at the sides and in front of it. They may be few or many. Sometimes they are free, in other cases more or less webbed together, and often more or less attached to the bases of the palpi (pl. xx, fig. 6, and pl. xxi, figs. 1, la, 3, e).
Sewes.—Some species are known to be dicecious, others are known to be moneecious, but most of the genera and species have not been studied with reference to this character.
A, EF. Verrill—Study of the family Pectinide, 58
Pallial eyes and tentacles.—All the larger forms have very numer- ous marginal, pallial tentacles, varying in size and length and gener- ally arranged somewhat in relation to the size of the corresponding radial ribs and grooves of the shell. There is a separate inner row or rows of “ guard tentacles” on a raised inner pallial fold (pl. xx, figs. 5, 6, 7, 8a). ‘The marginal tentacles are accompanied by a series of well-formed pallial eyes, very lustrous while living, and having a crystalline lens. These eyes in the larger species are numerous, and differ in size, the larger ones corresponding to the primary ribs, the smaller ones alternating in pretty regular order, according to the sculpture (pl. xx, figs. 6, 6a). The tentacles and eyes extend all the way around the margin of the mantle, beyond its free portions and even to the end of the auricles, which usually gape at the ends to give room for these organs. The tentacles and eyes are more or less reduced in the anal region. In some of the small deep-sea forms there are but few eyes, and in some cases they are not pigmented (at least in alcoholic specimens).
Remarks on the Nomenclature of Pectinide.
There is still so much diversity of opinion in recent malacological works concerning the nomenclature of the genera and subordinate groups of Pectinide that a brief review of the subject seems war- ranted. In general, the subdivisions here adopted correspond in most cases pretty nearly with those defined by Stoliczka,’ with some additional ones. But as Stoliczka considered that the ante- binomial names of Klein (1753), should take precedence of those given under the binomial rules, there is considerable disparity in the nomenclature. Mr. Dall’ has more recently discussed several of these groups. He followed the more generally adopted rules respect- ing priority of names, and therefore his conclusions were more nearly in accord with those adopted by me.
The most fundamental and impertant question to be settled in the nomenclature of this family is the correct application of the name Pecten to one of the restricted modern genera. The old genus Pecten has been divided by various authors into several genera, sub- genera, and sections, of very unequal value. Many of them were not definitely defined by their authors, and for several no definite
1 Ferd. Stoliczka, Mem. Geol. Survey of India. Cretaceous Pelecypod Fauna of southern India, Vol. iii, pages 423-430, 1871. 2 W. H. Dall, Bull. Mus. Comp. Zool., xii, pages 210-219, 1866.
54 A, E. Verrill—Study of the family Pectinide.
type-species were given. There is, therefore, much diversity of usage regarding their names and limits. The type-species of Pecten itself has not yet been settled. The name is of very ancient origin. It seems to have been first used, under the binomial system, by Miilier in Prod. Zool. Dan., 1776. His first species was P. maxinvus L. The same is true of DaCosta (1778), and Cuvier (1798). H. and A. Adams cited Linné as authority for Pecten, with P. varius as type, but I cannot find any basis for so doing, for Linné never adopted the genus. Fischer and others go back to ancient polyno- mial writers for the name, but the works of such authors should have no influence in determining priority of names under the binomial system. ‘
The determination of the true type of Pecten depends upon what rules of nomenclature one adopts. If we follow the well known and important rule that priority of binomial names does not apply to the names in works earlier than the 10th edition of Linneus, we must treat the names given by Klein as dating only from the time when first adopted by a binomial writer. Much of the confusion and disagreement in the nomenclature of authors is due to the neg- lect of this rule by several prominent writers. By the application of the rule much confusion may be avoided in the future. Klein, him- self, merely adopted the name Pecten, from much earlier writers, in a somewhat restricted or modified sense. His first division of Pecten was made for the one-sided species, and the first species in his list was P. maximus. On a subsequent page, however, he gave the name Vola to another species of the same group. The latter, judged by his own diagnosis and figure, should have beem placed in his first section of Pecten.
The author who first subdivided an old genus has the right, under the rules generally adopted, to assign the old name to either division unless there be very positive evidence that a special type had pre- viously been designated by its originator.
The first binomial writer to subdivide Pecten and restrict the name to a particular type was Bolten, 1798.’ He divided it into
1 As Bolten’s work is rare, I here reproduce an extract from it relating to the
Pectinidi, furnished to me by Dr. Dall. ‘ Bolten in 1798 worked in a very rational manner. He divided the old genus Pecten as follows: ” CHLAMYS. 1. GLABR&. Striate (two names of species, both = P. islandicus). * * Suleate (10 species; citrina, glabra, tranquebarica, gibba, radula). Plicate (15 sp.; cornea,
crocea, rubiginosa are identifiable).
i‘
A. E, Verrili—Study of the family Pectinide. 5:
or
three genera: Amusium (type A. plewronectes) ; Chlamys (type = U. Islandica) ; Pecten (type P. maximus, also dubius, Jacobwa, etc.)
He gave no diagnoses, but cited well known and figured species as types, so that his meaning is clear,
Schumacher, in 1817, undoubtedly ignorant of Bolten’s work, again divided Pecten. He restricted Pecten to the group called Chlamys by Bolten, and for the true Pecten he proposed the name Janira. He used Amustwm for the same type as Bolten, and established another good division under the name Pallium, for P. plica Linné,
The genus Amusium was adopted by Bolten from Klein, and its type has never been in doubt.
The genus Chlamys of Bolten included a large majority of the species included in the genus Pecten by Lamarck and most of the conservative writers. The name was adopted by H. and A. Adams for a small and ill-defined section of Pecten, while most of the more typical forms cited by Bolten were retained in Pecten, so that Pecten of H. and A. Adams (type, P. varius) is practically the same as Chlamys of Bolten.
Fischer, 1887, adopted Chlamys, in its widest sense, to include the greater part of the family Pectinide, such groups as Pseudamusium, Pallium, Lyropecten, Camptonectes, and several others, being regarded as its subgenera or sections. But C. Islandica was cited as the special type of the restricted group.
Stoliczka, 1871, adopted Chlamys in a more restricted sense, with C. bifrons Lam. as the type. He followed H. and A. Adams in adopting Pecten for a very large group, with P. varius Linné as the type, while the true genus Pecten was called Vola (after Klein and H. and A. Adams). It is singular, however, that none of those writers who have adopted Klein’s generic names have referred to the fact that Klein himself placed the one-sided pectens in the first sec- tion of his genus Pecten, and that his first “ species” included
P. maximus! On a subsequent page he gave Vola, with a brief
diagnosis, for a single species of the same group, evidently not real- izing its close affinity to his typical Pecten. Thus Vola was a syn- onym of Pecten even in the work of Klein !
2. ScaBRriuscuLz. Nodosa (1 sp.; P. nodosa). Squamate (13 sp.; P. pallium, sulphurea, porphyrea, aurantia, pusio, varia, lingua-felis, incarnata, pseudamusium, and
_ vitrea are identifiable.)
Rage.
, PECTEN. (P. maxima, dubia, Jacobea, pictus, ziczac.)
AMUSIUM. (P. plewronectes, japonicum, Laurentii.)
56 A. E. Verrill—Study of the family Pectinide.
Synospis of the generic and sectional groups of Pectinide.
After the previous explanations it will, perhaps, be useful to give, in a brief summary, the divisions of the old genus Pecten that seem worthy of recognition, either as genera, or subgenera, or sections, with their original types, so far as they can be fixed. The groups are here arranged in the order of their sequence in date, under the binomial system. Several fossil genera are here included, for com- parison with modern forms, but some fossil groups are omitted for lack of accurate knowledge of their characters.
Pecten Miiller, 1776. Type, P. maximus Linné.
Pecten (1st section) Klein, 1753 + Vola.
Pecten Miiller, Prod. Zool. Dan., 1776 (pars); Pecten Bolten. 1798 (rest.) ; DaCosta, 1778; Cuvier, 1798; Lamarck. Syst., 1801.
Janira Schumacher, 1817: Dall (pars) 1886; Fischer, 1987.
Vola H. and A. Adams (after Klein), 1858; Stoliezka, 1871; Zittel, 1881.
Vola + Janira Chenu, 1862.
Since Bolten, in 1798, definitely restricted the name Pecten to this group, as explained above, his restriction has precedence over that of Schumacher. The shells are generally large and heavy, and the valves are very unequal even when very young. The right valve is strongly convex with a large and much incurved umbo and beak, while the left valve is flat or even concave. It is usually smaller than the right, and shuts closely inside of its scalloped margin, and its umbo is nearly or quite obsolete. The auricles are of moderate size and not oblique, and in the right valve they are strongly convex or excurved in the middle. This valve has a sinuous, excurved bys- sal notch, with obsolete pectinidial teeth. The opposed auricles of the left valve are deeply ineurved to fit closely against the others. The hinge-plate in both valves has usually two or three divergent ribs on each end, of which the innermost is usually the strongest and most divergent. The resilium in the right valve rests on a shelf-like chondrophore inside the hinge-margin. A distinct tooth-like tubercle exists below each auricle within the margin of both valves. The surface of both valves has strong radial ribs interlocking at the mar- gin. Internally there are angular, thickened and fluted radial ribs, opposite the external grooves ; these ribs become more prominent and bicarinate or fluted near the margins.
Some of the species are known to be hermaphrodite. The foot of P. maximus is described as spatulate at the end.
A. FE. Verrill—Study of the family Pectinide. 57
Chenu and some other writers have made two generic divisions of the one-sided Pectens, viz: Vola, type P. maximus L.; and Junira, type P.atavus. The latter is a fossil of cretaceous age. No import- ant diagnostic characters have been pointed out, however. These two names, as originally used, were absolutely synonymous.
This genus occurs in rocks as early as the lower Cretaceous. The extinct group Vetthea is closely allied. It differs only in having conspicuous transverse incisions on the upper cardinal ribs. Pecten has a wide distribution in all tropical and subtropical seas. P. zic- zac L. and P. hemicyclicus Ravenel occur in the West Indies; P. dentatus is found on the Pacific coast of America, from the Gulf of California to South America.
Amusium Bolten. Type, A. plewronectes (Linné).
Amusium Bolten, 1798; Muhlfeldt, 1811; Schumacher, 1817; Woodward, 1866; Dall, 1886.
Amussium (pars) H. and A. Adams, 1858; Stoliczka, op. cit., p. 426, 1871; Fischer, 1887; Zittel, 1881.
Pleuronectia Swain., 1840; Chenu, 1862.
In this very distinct generic group the shell is round, thin, nearly smooth and strongly compressed. ‘The surface is often polished, sometimes lightly radially striated, never strongly ribbed. The mar- gins are simple and thin. The valves may be a little unequal in con- vexity and usually differ in color, and somewhat in sculpture. The valves come together ventrally, but usually gape at both ends. The auricles are small, symmetrical, nearly equilateral, often with lateral erure ; the byssal notch is small or absent, pectinidial teeth nearly or quite abortive. The adult probably has no byssus. Hinge-plate simple. Interior of valves strengthened by a number of raised diver- gent ribs, or lire, independent of any external sculpture. Accord- ing to Dr. Dall, the foot in A. plewroneetes = Mortoni is large, with “a spade-shaped tip and well-developed sucker, with moderate stem.” In A. Dailiit is described by him as having the “ sucker large, hood-shaped, thin-walled and darkly pigmented, with a broad base, abruptly enlarged from a very slender stem.” The stem is strongly grooved, the expanded cord is hollow and forms “ an exag- gerated and efficient sucker.” The ocelli are witbout pigment.
In the latter species there is but a single gill on each side, “ fur- nished with long separate filaments, much as in Dimya.”’
1 Owing to these important differences in the structure of the gills and fool, it seems necessary to place this species in a distinct genus, for which I have proposed the name Paramusium. (See p. 72.)
58 A, E. Verrill—Study of the family Pectinide,
The expanded end of the foot, in this and some other genera, is probably used as a pushing organ rather than a “sucker,” as sup- posed by Dr. Dall. It is analogous to the large pushing disk on the foot of Nucula and Leda. A sucker would not be of much use on the soft mud where many of these species live.
The type of this genus is from the East Indies; other large species, closely allied, are found at Japan (A. Japonicum), and elsewhere.
A species, referred, at first, by Dr. Dall to A. pleuronectes (1.), hut perhaps the A. Mortoni, and A. Dalli were dredged in the West Indies and Gulf of Mexico by the “ Blake” Exp.. The latter ranged from 218 to 1591 fathoms. The former was from 35 to 60 fathoms. Most of the species are from deep water and mud or ooze bottoms.
Species of the genus occur fossil in Cretaceous and Tertiary rocks.
Curiously enough, H. and A. Adams did not give the presence of internal ribs as a character of the genus, but« based it only on the form of the shell. Accordingly, they included our large American species, C. Clintonius or Magellanicus in the genus Amusium, from: its general external resemblance to shells of that group. It is really much more nearly allied to the typical species of Chlamys and Pseu- damusium, so far as the shell is concerned.
Amusium must be restricted to those species having specialized, internal, radial ribs and small auricles.
Chlamys Bolten. Type, C. Islandica (Linné).
Chlamys Bolten, Mus. Bolt., ed. I, p. 165, 1798, restr.; Fischer (pars), 1887.
Pecten (restr.), Schumacher, 1817.
Pecten (pars) and Chlamys (pars), H. and A. Adams, 1858; Chenu, 1862; Zit- tel, 1881.
Pecten (restr.), Stoliczka, 1871.
PLATE XVI. figs. 2-5. Puate XX. fig. 9. _PuaTe XX. fig. 2.
As stated on a previous page, the original type of this genus is identical with P. Zslandicus (Linné). Therefore this should be adopted, without question, as the true type, as has been done by Fischer and others.
The group called Chlamys by Stoliczka is the same as -4quipecten Fischer, 1866 ; it is usually regarded as a mere section of Chlamys. (See p. 67.)
The typical species of Chlamys are high, rounded, somewhat oblique, nearly equivalve shells, with large inequilateral and oblique auricles ; a large byssal notch; and several pectinidial teeth. The surface is strongly radially sculptured, both with primary and with
A. E. Verrill—Study of the family Pectinide. 59
numerous interpolated ribs, increasing in number with age. The ribs are generally crossed by concentric sculpture, often forming rough scale-like projections. The margins are scalloped and the shell closes rather tightly except at the byssal area. The inner sur- face has ribs and double flutings, corresponding to the external grooves and radii. The hinge-plate generally has two slightly divergent ribs on each end. For some account of the anatomy of C. Islandica, see p. 72. Some of the species are known to have sep- arate sexes.
The genus is world-wide in distribution. It appeared early in mesozoic geological time (Triassic) and is common in Cretaceous and Tertiary rocks.
Among the American species of Chlamys are the following, be- longing to the typical or restricted group: C. Zslandica Linné, from the northern coasts and fishing banks; C. ornata (L.), from the southern coasts of the United States and the West Indies; C. exas- perata (Sow.), West Indies; C. costellata Verrill and Bush, from deep water; C. Benedicti V. and B., from deep water off the east- ern coast of the United States and in the West Indies; C. phrygia (Dall), Gulf of Mexico and West Indies, 95-127 fath; C. effluens (Dall), 127 fath., off Havana. Several of the common American species belong to the subgenus -dquipecten (see p. 67). Among them are C. irradians, C. dislocata, C. Antillarum, C. glypta (pl. xvi, figs. 7-11), and C. ventricosa.
Pallium Schumacher. Type, P. plica (Linné).
Pallium Schumacher, 1817; H. and A. Adams, 1858; Chenu, 1862; Stoliczka, 1871; Zittel, 1881; Fischer, 1887.
Dentipecten Ruppel, 1835.
PLaTE XXI. fig. 4.
The special feature of this very distinct group is the development of several (usually three) well-marked, nearly transverse, blunt teeth, alternating with distinct pits, on each end of the hinge-plate. The shell is elevated, rather thick, with external, large, obtuse or rounded, radial ribs or corrugations and with internal, angular, dou- ble or bicarinate ribs, opposite the external grooves, near the margin. The auricles are small, but high. The hinge-teeth are marked with distinct cross lines.
Hinnites Defrance, 1821. Type, H. Cortessi Def.
Shell free and much like Chlamys, when young, but later in life it becomes attached by the right valve and irregular. Mr. Dall has
60 A. E. Verrill—Study of the family Pectinide.
described H, Adams, from the West Indies, in deep water. H. pusio is from the European coasts. Other species occur in the East Indies. The type is an extinct tertiary species.
Neithea Drouet, 1824. Type, P. equicostatus Lam.
This group agrees in form and sculpture with typical Pecten. It differs chiefly in having a series of transverse denticles and pits along the dorsal border of the hinge-plate. These seem to be homologous with the much finer transverse incisions and denticles — found in many living Pectinide, but are more highly developed. It — seems to be scarcely more than a subgenus or section of Pecten.
All the species are mesozoic fossils.
Hemipecten Adams and Reeve, 1849. Type, H. Forbesianus Ad. and R.
This group includes species with thin, irregular shells attached by the right valve, like Hinnites, but the attachment is effected mainly by a permanent modified byssus. The posterior auricles are nearly obsolete. The byssal notch becomes irregular and nearly en- closed, as in Anomia. Ido not know any American species.
Aviculopecten McCoy, 1855. Type, A. concavus McCoy.
Shell broad, roundish, more or less inequilateral and oblique, with regular radial sculpture. Auricles unequal, the anterior smaller with a byssal notch. Hinge-plate without a central resilium, but with a ligamental groove along its entire length. The absence of a central resilial pit renders it doubtful whether this genus should be placed — in this family. It may belong rather to Aviculide. .
The genus is confined to the Palzeozoic rocks,
Pseudamusium H. and A. Adams. Type, P. ewoticwm (Chem, Lam.)
Pseudamussium (pars) H. and A. Adams, 1858 (after Klein); Chenu, 1862; Stoliczka ,- 1871; Zittel, 1881; Fischer, 1887; Dall (pars) 1886.
Puate XVII. figs. 8, 8a.
The typical species of this group have nearly smooth, round, sym- metrical, closed shells with well-defined, small, straight, obtuse- angled auricles. The valves are nearly equal, and have nearly sim- ple, even margins. The external sculpture consists of small radial strie or riblets, without strong angular ribs and grooves, and it may differ on the two valves ; the margin is not scalloped, or but faintly so, and there are no definite internal ribs. The hinge-plate usually has but one longitudinal fold on each end; this is feeble and nearly
A. E. Verrill—Study of the family Pectinide. 61
parallel with the marginal ligamental groove. It is usually cross-
incised. The byssal notch is small and the pectinidial teeth are up
to five in number, or sometimes lacking. Some of the species, if not all, show the fine divergent “ camptonectes-seulpture,” on one or both valves, especially when young.
This group was adopted by H. and A, Adams, from Klein, but they gave no adequate definition, and designated no special type. They gave an alphabetical list of twenty-one species. Among these are representatives of several diverse groups (Amusium, Cyclopecten, Chlamys, Palliolum or Camptonectes, etc.). If these incongruous groups be eliminated, those that remain may be referred to the group characterized above, with such species as P. dispar (Lam.) and P. exoticum (Lam.) = pseudamusium (Sowerby) as typical forms, whether they be distinct species or not.
Klein, himself, referred to his genus certain fossil shells which, as he stated, differ from Amustwm only in lacking ribs and grooves. He also cited one of Lister’s species, with his diagnosis, but he gave itno name. His figure (copied from Lister) is a crude representation of a variegated species, like P. exoticwm or P. dispar.
Although I do not, personally, consider Klein’s type as of any importance in the limitation of the group, it may satisfy others to
_ know that his type is the same as that adopted above.
Chenu, 1861, gave as “examples” three species: P. dispar (Lam.); P. pseudamusium (Lam.); P. glaber (Linné). The first two are typical, but the last belongs to another section.
Stoliczka gave a definition of the group, with P. exoticum as the type. He also cites P. corneum, P. hyalinum, P. tigrinum, and P. natans as typical examples. Some of these belong in diverse groups.
Zittel, in 1881, gave P. glabrum and P. hyalinum as types.
From the incongruous species included in this group by H. and A. _ Adams and others, we may well separate those that have internal ribs, and also the thin, delicate, deep-sea species, with unlike _ valves, the right valve usually having strong concentric sculpture.
For this last group I have established the genus Cyclopecten. Of
the American specjes, our common large New England species, P. Clintonius or Magellanicus (Lam.) resembles this subgenus. It is _ similar to the typical species in form of shell and auricles. It differs _ mainly in gaping at both ends and having stronger radial sculpture onthe upper valve. But the character of the shell and the form of _ the foot are so peculiar as to warrant the institution of a special _ genus or subgenus for its reception (see p. 69).
x an
62 A. E. Verrill—Study of the family Pectinide.
Of the European species examined by me, P. similis (pl. xvii, figs. 8, 8a) seems to belong to the restricted group. Allied species occur in the Tertiary and Cretaceous formations.
Camptonectes Meek, 1864. Type, P. lens Sowerby. Camptonectes Stoliczka, 1871; Zittel, 1881 (type, arenatus Goldf.).
Shell subovate, plain, not corrugated, and without strong radial ribs; margin nearly plain. Valves subequal. Auricles unequal ; byssal notch well developed. Surface of the shell covered with fine, — obliquely divergent, curved, crenulated or vermiculated riblets with intervening, narrow, punctate grooves.
The curious vermiculated sculpture is not peculiar to this division, — but is more or less obvious on the shells of some species of Pseuda- musium, and on species of several other groups, both with and with- out radial ribs. It is a structural feature that runs obliquely across the ribs and grooves. Most of the species are mesozoic fossils.
The recent Pecten striatus and P. tigrinus Lam. of Europe, appar- ently belong to this group, and P. Testw might, also, well be referred to it. It is generally regarded as only a section of Pseudamusium.
Entolium Meek, 1864. Type, D. cornutwm Queenst.
Body of shell rounded, not oblique ; valves thin, nearly equal. Sculpture delicate. Auricles well developed, those of the right valve prolonged dorsally beyond the hinge-line in the form of angular lobes. Byssal notch obsolete. Apparently there are no internal ribs — or lire. This genus appears to be allied to Amusiwm, which it — resembles in the form of the disk, shortness of the auricles, absence of byssal notch, and apparently in the texture of the shell. But it differs in lacking the internal lire, and especially in the dorsal pro- longation of the auricles. The last character distinguishes it, also, from Protamusium V., to which it appears to be still more closely related.
Syncyclonema Meek, 1864.
Type=P. rigida Meek and Hall, Mem. Amer. Acad. Arts and Sciences, Boston, vol. Vv, p. 381, pl. i, figs. 4, a, b, c=P. Halli Gabb.
Syncyclonema Meek, Smithsonian Check List Cret. Fossils, p. 31, 1864.
The type of this group is a small, poorly preserved, thin shell, with concentric grooves or undulations on at least the “left or inferior” valve, and with radial striz on the other. The shell is compressed, obovate, higher than long, not oblique, gradually narrowed to the
AGE: Verril—Study of the family Pectinide. 63
v
auricles, which are small and nearly equal, and apparently differ but little in sculpture, but the actual sculpture of the type is uncertain.
Many mesozoic fossil species appear to belong to this group. The type is from the North American cretaceous formation.
Some of the broad fossil species that have been referred to this group appear to belong rather to Protamusium. These have tine concentric grooves and lamelle, and equal auricles, but no radial sculpture.
Several living delicate deep-sea species are perhaps nearly allied to this group.
As the real structure of the type of the genus is still uncertain, I have thought best to make a new genus for the recent forms, under the name of Hyalopecten (see p. 71).
Among the known species are H. undatus V. (pl. xviii, fig. 5) #1. dilectus V. and B., 1. fragilis (Seft.), H. pudicus (Smith).
The group of shells here indicated forms a very definite division, worthy, perhaps, of generic rank. These recent species do not show the camptonectes-sculpture, but both valves are distinctly undulated. Some of these species are otherwise smooth, but others are finely radially striated. They are all hyaline, very thin, and very simple in structure.
b
Pernopecten Winchell, 1865. Type, P. limijormis W.
Pernopecten Winch., Proc. Acad. Nat. Sci., Philad., for 1865, p. 125; Hall, Pal. New York, v, pt. I, sec. IT, Introd., p. Ivii, figs. 1, 2.
The type of this genus has the form of a small round Amusium, or Propeamusium, to which it appears to be closely allied. The
auricles are small, short, subequal, poorly differentiated, much as in
Amusium. Byssal notch slight. There is a well-developed median
_ resilial-pit. The cardinal ribs bear a series of minute transverse
&
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: é
grooves, or “crenulations ” on each end, like those of Huchondria,
ete. There is a well-developed, rib-like auricular crura on each end.
The shell is smooth, or has fine radial striz.
Burlington Sandstone of Iowa.
This genus should probably be regarded as the ancestral form of Amusium and Propeamusium. It was probably a free-swimming species, like the modern genera.
Lyropecten Conrad, 1867. Type, P. nodosus (Linné). Liropecten of several later authors.
Shell large and strong, corrugated, with large, fluted, and usually nodose, primary radial ribs, which do not increase in number, and
64 A. E. Verrill—Study of the family Pectinide.
with coarsely scalloped margins. Valves somewhat unequal. Auri- cles of medium size, unequal. Hinge-plate with several, usually three, oblique, divergent ribs on each end. This is one of the best defined groups, and may be regarded as of generic value. It is allied to Pallium.
Several species occur in the American tertiary deposits. Z. nodosus occurs at Florida and in the West Indies. JZ. subnodosus is from the tropical regions of the Pacific coast of America. JL. corallinoides (D’Orb.) is from West Africa and the Canary Islands; Z. noduliferus (Sow.) is from East Africa.
Euchondria Meek, 1874. Type, Z#. neglecta (Geinitz).
Euchondria Meek, Amer. Journ. Sci., 3d ser., vol. vii, p. 443, 1874; Hall, Pal. of New York, v, Pt. 1, sec. Il, Introduction, p. Ixii, figs. 4, 5, 1885.
The shell of the type of this genus has nearly the form of Cyelo- pecten and Propeamusium, to which it seems closely allied. The auricles are well-developed and angular, subequal. The body of the | shell is well rounded; the sculpture is slight, consisting of concentric¢ lines on the body, but there are radial ridges on the auricles. There is a distinct triangular chondrophore, situated slightly one side of the beak. The hinge-plate has a row of very small, close, incised, transverse pits or grooves on each end. These seem to correspond closely with the grooves, alternating with denticles, on many modern species, and not with ligament pits.
Carboniferous of Illinois.
Propeamusium Gregorio, 1883. Type, P. inequisculpta Tib. = P. fenestratum Forbes. Propeamusium (subgenus) Dall, Bull. Mus. Comp. Zool., xii, p. 210, 1886; Fischer,
1887. PLATE XX. figs, 5-9.
This group is a subdivision of Amusiwm. It includes small, mostly ~ deep-sea species, with rounded thin shells, having the valves unequal in size and sculpture; the lower and flatter one is concentrically grooved, and usually turns up at the thin margin to meet the upper valve, as in Cyclopecten. The upper valve may be cancellated or radially sculptured. When full grown there are several well-formed, raised, internal ribs ; these may be absent in the young.
This division differs from Amusiwn in the sculpture of the valves and in having the auricles and byssal notch well-developed.
The species closely resemble those of Cyclopecten ; the only ob- vious difference in the shell consists in the internal ribs, The species
A. E. Verrill—Study of the family Pectinide. 65
are mostly from the deep sea, and several are from the West Indian area. P. thalassinum (Dall) is the only species taken off our northern coast, and this is not a typical species, as it has but two radial ribs, _ besides the auricular crure. Dr. Dall has recorded the following species from the West Indian region: P. Pourtalesianum Dall, 13 _ to 805 fath.; P. cancellatum (Smith), 13 to 1591 fath.; P. Holmesii
Dall, 100 fath.; P. Sayanum Dall, 16 to 150 fath. Tle also records _P, Alaskensis Dall, from Alaska; P. Hoskynsi (Forbes, non Sars, nec Jeff.) from the East Atlantic and Mediterranean ; P. lucidum (pars) Jeff., East Atlantic and off Brazil, 675 to 1000 fath.
The following additional species, described as species of Amusium, were obtained by the Challenger: P. scitulwm (Smith), 28 fath., off New Guinea; P. TJorresi (S.), 155 fath., off Cape York ; P. propin- quum (8.), 100 fath., off the Azores; P. obliquum (S.), 390 fath., West Indies. P. thalassinuin was taken in 43 and 417 fath.
_ Crenipecten Hall, 1883. Type, C. crenulatus Hall; Devonian. Crenipecten Hall, Pal. of New York, vol. v, pt. I, plates and expl., p. 3, 1883; sect. I, Introd., p. xii, pl. ix.
The shell and auricles are shaped much as in Luchondria. Some of the species show radial sculpture.
The hinge is said to have no median resilial pit, but to have a continuous series of small transverse pits, grooves, or incisions, along the whole of its length. These minute pits are probably of the same nature as those of Huchondria and many modern Pectinide, and not ligament-pits.
Several species have been described from the Chemung and Way- erly formations.
The alleged absence of a resilial pit is the only tangible difference between this and Euchondria.
Palliolum Monterosato, 1884. Typescited: P. Teste Biv, and P. vitrewm (Chem.)
Puate XVIII, figs. 6-14.
This is a group separated from Pseudamusium of H. & A, Adams, _and is scarcely to be distinguished from Camptonectes.
The two species named by its author as types agree in having thin, rounded, nearly equivalve, shells, with the posterior auricle poorly developed, and with fine camptonectes sculpture on both valves, with small radial riblets, and usually with rows of small scales. The “margins are plain and come evenly together, without flattening.
TRANS. Conn. Acap., Vou. X. JUNE, 1897. 5
66 A. EF, Verrill—Study of the family Pectinide.
They are closely allied to P. striatum (pl. xviii, figs. 14, 14a) in form and sculpture and, like the latter, might well be referred to Camp- tonecteés.
The second type-species (P. vitrewm) is rather more distinct, on account of the very slight differentiation of the posterior auricle, which is small and resembles the condition seen in the very young examples of other species. This feature is also present in the other species in a somewhat less degree.
P. vitreum (pl. xviii, figs. 6-13) has a broad, round, thin hyaline shell, with the auricles unequal; the posterior auricle is short and — only slightly differentiated from the shell; it has an obtuse dorsal angle from which the margin slopes to the body of the shell, without a notch or angle. It has a number of sharp, free, pectinidial teeth and a long series of discarded ones above the byssal fasciole. Auric- ular crure are rather prominent. The byssal notch is deep, and the byssus is well developed. It is used to attach the shell to branches of gorgonian corals (Acanella, Primnoa), etc. The “ camptonectes- — sculpture” is strongly marked. The mantle has rather few tentacles and pigmented ocelli.
This arctic and northern European species has been recorded (Voy. Challenger) from the E. Indies, 100-700 fath., and from off Japan, 345 fath., and off Patagonia (140 to 400 fath.) It is common on the | deep-water fishing grounds off Nova Scotia and Newfoundland, in 57 to 400 fathoms, and extends southward in deep water down to 1537 _ fath., off Chesapeake Bay. z
P. Teste is less hyaline, and the auricles are more nearly equal. It — has well-marked camptonectes-sculpture and radial riblets on both valves. ‘There are several pectinidial teeth. According to Jeffreys (Brit. Conch., v, p. 167), the animal of this species has two unequal — rows of slender tentacles, the inner row much the smaller. Ocelli few, in two rows, those in the outer row unequal in size and position ;- those of the inner row are much smaller and more numerous. Foot cylindrical. It swims actively and often attaches itself by a byssus.
It has been brought up attached to a telegraph cable from the depth of 1000 fathoms in the Mediterranean.
My conclusion is that these two species must be placed in the same section with P. striatum and P. tigrinum. Nor can I see any valid — reason why the four should not be placed in Camptonectes.
It is true that the ‘‘ camptonectes sculpture ” is found also in larger — Species of allied groups when young (e. g. P. Clintonius), and also in — certain ribbed species of Chlamys, but in the typical forms referred to Camptonectes it is the predominant sculpture through life,
A, FE. Verrill—Study of the Jamily Pectinide. 67
These may be regarded as primitive and simple forms, from which larger species with stronger ribs have been developed without losing this primitive fine sculpture. But this kind of sculpture depends upon structural features that must be taken into account in the classification of the family. Possibly all true species of Psewdamu- sium have, also, the camptonectes sculpture, but I have not at hand snfficient material to settle this question.
It appears to be lacking in Cyclopecten, Pectinella (typical), Prop- eamusium, Amusium, Paramusium, Pecten, Lyropecten, and Pal- lium. Nor have I observed it in various species of -guipecten examined, It is also absent from many species of Chlamys (restr.)
Zéquipecten Fischer, Manual, 1887. Type, Chiamys opercularis (L.) PLATE XVI. figs. 6-11. PLaTe XX. figs. 1-3, 6, 6a.
Shell broadly rounded, with the valves nearly equal and sym- metrical. Auricles well formed, angular ; byssal notch well-devel- oped. The sculpture consists of a moderate number of large and nearly equal primary radial ribs, which increase in size, but are not much increased in number with age, by the interpolation of new ones. Internal ribs or flutings correspond to external grooves, but each one is bicarinate or double, especially near the margins. Hinge-plate with one or two slightly divergent ribs at each end, often crossed by strong transverse incisions. Pectinidial teeth abortive in the type, but present in most species. Foot of the type-species is subcylin- -drical, well-developed, with a byssal fissure and a terminal deeply bilobed “scoop-shaped” disk, which can be expanded. In our C. irradians (see pl. xx, fig. 6) the foot has a similar structure, but the terminal disk appears to be smaller. In the type there are (t. Jef- freys) 35 to 40 ocelli, and two or three irregular rows of tentacles,
The type-species of this group (C. opercularis) has a well-rounded, ‘searcely oblique shell, with the valves a little unequal and conspicu- ously gaping at both ends, but especially so anteriorly. The auricles are well-developed, angular at both ends, and nearly equal. The byssal notch is large, but the pectinidial teeth are nearly or quite obsolete in the adult. Both valves have about twenty-one large “primary ribs, which are at first narrow and angular, but become
broad and rounded by growth. In the left valve both ribs and “grooves are covered with small radial riblets, decussated by fine concentric lamellae. In the right valve the radial riblets are less distinct, but evident. The interior has about twenty-one bicarinate ‘Tibs or flutings on each valve. The hinge-plate has one broad, low,
¥
68 A. E. Verrill—Study of the family Pectinide.
transverse rib, besides a very narrow one bounding the ligamental groove. In the right valve the margin, above the ligament, is in- curved. The auricular crure are feebly developed, but end in dis- tinct denticles, larger in the right valve.
In most of the species referred to this group the valves gape ae less at the ends than in the type-species.
A large number of shallow-water species from all parts of the world belong to this group. Many of them grow to large size. The following are some of the American species :—
C. irradians (Lam.) (pl. xvi, fig. 6 ; pl. xx, figs. 6, 6a), from Cape Cod to the Gulf of Mexico; C. dislocata (Say), Cape Hatteras to the West Indies; C. Antillarum (Recl.), West Indies; C. nucleus (Born.), West Indies; C. glypta Ver. (pl. xvi, figs. 7-11), off the Eastern coast of U.S. and in the West Indies, 69 to 200 fath.; CG. ventricosa (Lam.), Pacific coast of tropical America ; C. purpurata (Lam.), Pacific coast of South America; C. caurina (Gould), coast of California, ete.
Pectinella, gen. nov. Type, P. Sigsbei Dall.
Shell small, thin, swollen, nearly smooth, with convex and slightly unequal valves. Auricles very unequal, oblique, the anterior larger, with a deep byssal notch in the right valve, but without pectinidial teeth ; posterior auricle small. The surface is smooth except for fine lines of growth. Camptonectes sculpture is not present. The texture is not hyaline.
The only known species is Pectinella Sigsbet (Dall), Bull. Mus. ' Comp. Zool., xii, p. 223, pl. iv, fig. 2, 1886. It was taken by the Blake Exped. ‘A the West Indies, in 158 fathoms.
This form differs so much from all the other divisions of Pecti- nid that it seems necessary to form a new genus for it. In its swollen form, it approaches some of the species of Zéimopsis and allied forms.
Lissopecten, new subgenus of Chlamys. Type, L. hyalinus (Poli).
Shell slightly inequivalve, broadly rounded, not oblique, thin, translucent, nearly smooth. The external sculpture consists of faint, nearly obsolete radial ridges and obscure riblets, but one or both auricles may have a more or less cancellated sculpture. The interior sculpture consists of very distinct, simple raised ribs. Auricles angular, well developed. Byssal notch deep. Pectinidial teeth prominent. Margin not scalloped, nearly plain and simple.
A. E. Verrili— Study of the family Pectinide. 69
Although this group agrees with Amusiwm in having internal ribs without corresponding external grooves, it seems to be allied rather to Chlamys. It way be regarded as a division of the latter in which the external radial ribs have degenerated.
It includes C. hyalinus (Poli), from the Mediterranean, and several
other similar small species.
Leptopecten, new subgenus of Chlamys. Type, 0. Monotimeris (Conrad).
Shell thin, translucent, oblique, broadly rounded, with strong, rounded radial ridges or folds, like corrugations, which appear in reverse on the interior surface. The internal ribs are not angulated by a deposit of shell, nor distinctly thickened. Margin with broad scallops. The exterior surface is covered with fine divergent camp- tonectes sculpture, both on the ribs and intervals. The ribs do not increase in number with age, but become broader and more flattened. Auricles large and broad, thin, corrugated. Byssal notch large and deep. Pectinidial teeth prominent. Hinge-plate thin and but little differentiated. Cardinal ridge thin and small, close to the liga- ment, crossed by fine incisions. The resilial pit is small, but projects beyond the thin hinge-plate in the left valve.
This is a peculiar group, remarkable for its thin but strongly cor- rugated oblique shells, with fine camptonectes sculpture.
C. Monotimeris (Con.), from the California coast, is the only Species studied.
Placopecten, new subgenus of Chlamys. Type, P. Clintonius (Say). Puate XVII. figs. 1-7. Puate XX. figs. 7, 8, 8a. PLATE XXL. figs. 1, la, 2, 2a.
Shell large, compressed, broadly rounded, rather thin, with simple sharp edges, meeting evenly ventrally, but gaping considerably at both ends, especially when adult (pl. xvii, fig. 5). Valves only slightly unequal in form, the right one being a little flatter, but they differ in color and somewhat in sculpture, the right one being
smoother and paler. Both have fine radial lines or riblets, and they
rT § ¥
7
3 ~
A
x *
have vermiculated divergent riblets when young. Auricles small, symmetrical, nearly equal. Byssal notch small, simple. Pectinidial teeth generally obsolete, except when young. No internal ribs. Inner surface often with more or less pearly luster and a crystalline
structure. Hinge-plate with two feeble, slightly divergent ribs on
each end, crossed by fine transverse incisions. The foot (pl. xx, fig. 8) is well developed, oblique, slightly narrowed distally and enlarged at the end, where it is divided into two lobes by a rather deep,
70 A. EF. Verrill—Study of the family Pectinide.
oblique, longitudinal fissure, so that the lobes can be spread apart or closed, at will, thus resembling somewhat the foot of Ledida, Toward the base, on the anterior side, there is also a short, deep byssal slit, terminating at a prominent tubercle about the middle of the front side.
Cyclopecten, gen. nov. Types, Pecten pustulosus Verrill; P. imbrifer Loven. PLATE XVI. fig. 1. PLate XIX. figs. 1-4.
Shells thin, rounded, scarcely oblique, with symmetrical auricles and simple margins. The two valves are unlike in sculpture. The right valve is a little flattened and upturned at the flexible margin, — so as to fit tightly against the upper valve. The thin lower valve has, in the typical species, regular, thin, elevated, concentric lamelle, —
Figure 1.— Cyclopecten pustulosus V.; a, left, 6. right valve, natural size.
which aid in the adaptation of the edge to that of the upper valve ; the margin is usually flattened or bevelled. The upper (left) valve is radially sculptured, rarely smooth ; it usually has radial rows of arched scales, pustules, or points, and also concentric raised lines ; it is sometimes cancellated. No radial ribs, nor interlocking points at the margin. Auricles well developed, subequal, angulated and well- defined at both ends; byssal notch well defined ; few or no pectini- dial teeth. Cardinal folds single, rather feebly developed, often cross-lined. Eyes few. Byssus small, and of few threads.
The species of this group have usually been referred to Pseuda- musium, but they differ widely from the typical forms of that group, such as P. exoticwm, P. dispar, etc., in which the valves are of nearly equal size, with simple edges that come evenly together without flex- ure of the lower one, and in which the auricles are small and nearly equal.
This genus includes a large number of small species, mostly from deep water. Among these are the following: C. imbrifer (Lov.), northern coasts of Europe; C. pustulosus (Ver.) (cut, fig. 1), (pl. xix, figs. 3,4); C. subimbrifer (V. and B., see p. 84), 121 fath.; C. leptaleus (Ver.), 142 fath.; C. nanus V. and B.; (pl. xvi, figs. 12- 12c¢), the last four are from deep water off the eastern coast of the
A. EF. Verrill—Study of the JSamily Pectinida. 71 United States; C. reticulus (Dall); C. simplex Ver. (pl. xvi, fig. 1, xix, figs. 1, 2); and C. Culebrensis (Smith), 390 fath., are from the West Indies; C. Murrayi (Smith), 1400 fath., off Australia; @. clathratus (Mart.), 120 fath.; C. subhyalinus (Smith), 400 fath.; and C. distinctus (Smith), 100 fath., from the Antarctic regions ; CU. Ker- madeciensis (Smith), 600 fath., off Kermadee I.
C. orbicularis (Sowerby), which occurs on the west coast of Africa, living among, and usually attached to, floating fucoids (Sar- gassum, etc.), near the shore, appears to belong to this genus. It has concentric sculpture on both valves; that on the left forms raised seale-like lamellz. The shell is hyaline. The valves close tightly by the upturning of the edge of the right one. According to Dr. Charbonnier (Journ. de Conch., ser. ITI, vol. iv, p. 261) this. species swims about very actively, but attaches itself very firmly and quickly (in 15 minutes), to floating alge by a byssus of several threads. When at the bottom of the glass vessel, it creeps about by means of its foot.
Hyalopecten, gen. nov. Type, H. wndatus V. PLATE XVIII. fig. 5.
Shell compressed, thin, hyaline. Valves nearly equal, with con- centric undulations or corrugations, affecting the entire thickness; margins simple; sculpture none, or consisting of fine radial lines on one or both valves, without camptonectes sculpture. Hinge-plate thin and nearly plain; auricles well developed, unequal; byssal notch distinct.
The possible relations of this group to Syneyclonema were dis- cussed on page 63.
The species known to me are as follows: 7. dilectus V. and B., from 1813 fath., off Martha’s Vineyard; H. fragilis (Jeff.), from northern Europe and the Arctic Ocean, and off the U.S. coast, in 578 to 1525 fath.; HZ. wndatus Ver., off the U.S. coast, in 1423 fath.; and H. pudicus Smith, off Marion I., in 1375 fath.
Protamusium, gen. nov. Type P. demissum (Phil.).
Body of shell disk-like, nearly circular, and compressed, valves thin, with fine regular concentric grooves and fine raised lamell. Auricles short, but distinct, angular, not oblique, nearly equal, not prolonged dorsally ; no byssal notch.
This division is proposed for certain mesozoic shells that closely resemble Amusium in form, but appear to be entirely destitute of the internal radial ribs, so characteristic of the latter. It differs
72 A, E. Verrill—Study of the family Pectinide.
from Entoliwm in not having the auricles strongly produced dorsally, though their distal angles are often a little prominent. The type (P. demissum) is from the Jurassic formation of Germany, as is also P. disciforme (brown Jura).
Various other allied species have been described from the Jurassiag
A considerable number of species that have been described as Amusium, from the Jurassic and Cretaceous, apparently belong to. this group, for they show no trace of internal lire. Among these are the following Cretaceous species: LP. membranaceum (Nils.), from Europe and India ; P. ilustre Stol., from India ; P. sedeatellum (Stol.), from India; P. obovatwm (Stol. as Syncyclonema), from India. ;
Paramusium, gen nov. Type, Amusivm Dalli Smith.
Shell thin, rounded, much compressed ; valves nearly equai ; sculp- ture nearly obsolete, different on the two valves; the lower valve has concentric undulations. Auricles very small, equal. Byssal notch and pectinidial teeth obsolete. The shell has a prismatic structure. Internal lire and auricular crure well developed.
A single pair of gills, with long, simple, separate filaments (t. Dall). The foot is slender, with a byssal groove; the end is much enlarged, with an oblique, expanded, concave terminal disk, striated within. No labial palpi. Ocelli without pigment. The. structure of the animal, as described by Dr. Dall, is very different from that of typical Amusitum. P. Dalli ranges from 218 to 1591 fathoms, from the Gulf of Mexico to Barbados. Another similar — species, P. meridionale (Smith), was taken by the Challenger Exp., off Brazil.
Descriptions of new species and descriptive notes on others, Chlamys Islandica (Chemn.)
Ostrea Islandica Miller, Zool. Dan. Prod., No. 2990, 1776. Fabricius, Fauna Gr6nl., p. 415, 1780. 3
Pecten Islandicus Chemn., Conch., vii, p. 304, pl. 65, figs. 615, 616, 1784, Lamarck, Anim. sans vert., ed. 2, vol. vii, p. 145. Hanley, Ree. Biv., p. 284. Gould. Invert. Mass., ed. 1, p. 133, fig. 87; ed 2, p. 198, fig. 495. Verrill, Invert. Vineyard Sd., ete., p. 402. G. O. Sars, Mollusca Reg. Arcticee Norvegize, p. 16, pl. ii, fig. 2, 1878.
Pecten Pealei Conrad, Amer. Mar. Conch., p. 12, pl. ii, fig. 2, 1831.
Chlamys Islandica Fischer, Man. Conch.
PLATE XVI. figs. 2-5b. Puate XX. fig. 9. Puare XXI. fig. 2) :
In this species the labial palpi (pl. xxi, fig. 3) are broad, triangular, — with the distal end acute, strongly flattened on the apposed surfaces.
A, HE. Verrill—Study of the family Pectinida.
~J
2 v
The bases of the outer ones are broadly connected with the mantle
lining, and thus enclose the anterior ends of the gills.
Around the mouth are about ten much-branched, unequal, brown, labial tentacles, which are extensively webbed together. Of these, three are at the posterior side of the mouth, one large median one, much divided, and one much smaller one on each side; a large much-divided lateral one lies on each side, at the base of the inner _palpus; a still larger and more branched one lies at the base of each outer palpus; three small ones united to the others by a web, lie in front of the mouth. The foot (pl. xx, fig. 9) in alcohol is oblique, _ stout at base, suddenly contracted at the distal third and again slightly enlarged at the blunt tip. Its anterior face is turned to the _ right and has a large byssal groove extending to the distal constric- tion; at the tip there is a very small deep slit, of which the sides ean be somewhat expanded ; this slit is entirely separate from the _ byssal groove. In the breeding season the abdomen is prominent and crowded with eggs; it projects downward, and bends abruptly _ backward, terminating. in a small, sharp papilla. Gills four, of the normal fillibranchiate structure. Pallial tentacles very numerous and unequal, the outer ones smaller and much crowded. Ocelli con- 3 Spicuous; there are generally three between every two of the
_ primary tentacles.
. Young specimens (pl. xvi) when 4™™ long and 4.5™™ high, show an g irregular “camptonectes sculpture,” together with small and simple _ radial ribs. The byssal notch is broad and angular, but rounded at : bottom ; three pectinidial teeth are developed.
Those that are 8" long and 9°5™™ high have four acute pecti- nidial teeth, and a broad notch, rounded within. The ribs have become stronger and the edges of the shell are scalloped. The valves gap slightly at the anterior end and at the ends of the auri- eles. Near the margin, especially of the left valve, concentric 7 _ sculpture appears, and by crossing the ribs produces a scaly appear- ance.
Var. insculpta, nov.
A variety is occasionally taken on our northern coasts, which has amore elaborate sculpture than usual. The concentric and diver- gent lamine and smaller radial riblets cross each other in such a way ‘that a peculiar decussated sculpture is formed between the primary ribs on the early part of the shell, while on the older parts the inter- _ spaces are covered with elevated scales. The surface rises into six
74 A. E. Verrill—Study of the family Pectinide.
or eight broad rounded radial ridges covered, like the interspaces, with rough radial ribs. The sculpture of this variety is figured on pl. xvi, figs. 4-5b. This variety is connected with the common form by intermediate specimens.
This species is found at and just below low-water mark, as well as in deeper water, down to 179 fath., in the Bay of Fundy and north-
ward to Greenland and Iceland, ete. It is common and of large size -
on the fishing banks off Nova Scotia and Newfoundland. Further south it occurs in 20 to 100 fathoms, as far south as Cape Cod. Dead shells were taken off Martha’s Vineyard in 69 to 194 fath.
It is found as a Post-pliocene fossil in Maine, New Brunswick, Canada, Labrador, Greenland and Northern Europe.
Chlamys Benedicti Verrill and Bush, sp. nov.
Shell small, higher than long, with the posterior auricle much
longer than the anterior, with a deep byssal notch in the right valve. —
The dorsal margin is straight and only slightly oblique ; the anterior auricle, in the right valve, is decidedly angular, with its outer end slightly incurved and serrated by the terminations of the radial ribs, The posterior auricle is considerably prolonged and angulated at the upper corner, obtusely rounded at the end and deeply notched where it joins the main shell; it has four strongly marked radiating ribs, besides the dorso-marginal fold ; below these there is a slightly con- cave space corresponding to the byssal notch. On the body of the shell there are six or seven sharp serrations along the lower margin of the notch. In the upper valve the posterior auricle is broad and decidedly angular, the dorsal and outer margin forming less than a right angle ; its surface is covered with about five or six radiating ribs decussated by more numerous and finer concentric raised lines, the anterior and posterior margins of the body of the shell slope about equally and form an acute angle ; the ventral margin forms a regular semicircular curve; its entire surface on both valves is
covered by strongly raised, rather close radiating ribs, separated by
rather wider deep grooves. The interspaces are decussated by regu- lar raised concentric lines ; these are scarcely apparent on the ribs except on very young shells, but there are rather strong elevated spine-like points, especially near the margins, arranged along the ribs in pretty regular concentric lines. These become higher and more pointed anteriorly, and are frequently nearly obsolete in the middle portion of the lower valve; in that case the ribs appear nearly smooth and rounded. ‘The ribs project at the margin as blunt points,
-
A. EF. Verrill—Study of the family Pectinide. rE
Gr
or serrations. On the inner surface there are radial grooves corre- sponding to the external ribs. The hinge-margin is thin, with a slender submarginal ligamentary groove and a small triangular resilial pit in the center. The color is variable. The single valve from station 2571 is uniform lemon-color ; those from the other locality are chestnut-brown and reddish, variegated with paler, and sometimes with white blotches,
Length of largest specimen, 55" ; height, 6™™ ; length of dorsal margin, 4™™.
Off Martha’s Vineyard, in 1356 fath., dead ; West Indies, in 25 to 72 fath., living.
This species is allied to C. varia of Europe, but when compared with the young of that species, of the same size, the radial ribs are found to be fewer and coarser, and there are other differences which render it probable that they are distinct species. The ribs are stronger and fewer than in ©. Jslandica, and the auricles are differ- ent in shape. It is probable, however, that it grows to a much larger size than any of the specimens obtained. It may possibly prove to be the young of some known West Indian species, but does not agree with any known to us.
Chlamys costellata Verrill and Bush, sp, nov.!
Shell small, thin, translucent bluish white, covered on both sides with continuous, elevated, and somewhat thickened, well separated radiating riblets, of which there are more than thirty on our largest example. Length of the shell considerably less than the height. Dorsal hinge-margin elongated, especially on the anterior end. In the right valve the anterior auricle is considerably elongated, obtusely rounded or subtruncate at the end, with a wide, angular byssal notch beneath it, and a broad, smooth, angular area next to the body of the shell, above which there are three well marked, angular, radial ridges, separated by wider concave interspaces. Posterior auricle small, triangular, the outer end convex, forming a little more than a right angle and with the posterior margin nearly straight and without any distinct notch.
The dorsal margins of the body of the shell are nearly straight and diverge at less than a right angle. The ventral margin is pretty _evenly rounded, but a little produced in the middle. The beak is
1 Figured in an unpublished paper sent to Proc. U. S. Nat. Mus. several months ago.
76 A, BE. Verrill—Study of the family Pectinide.
small, acute, appressed and does not project beyond the hinge- margin.
The radial ribs are very distinct and clean cut, thickened and rounded at the summit, and separated by nearly smooth intervals two or three times as broad as the ribs themselves.. The width of the ribs increases regularly from near the umbos to the margin. A few intermediate ridges commence near the margin. :
The left valve is badly broken. It is, however, somewhat more convex than the other, and the radial ribs are crossed by numerous concentric striations, giving them a finely crenulated or beaded appearance, The anterior auricle is broad-triangular, the outer end slightly rounded, and with a slight incurved notch below. It has © about six small radial ribs, similar to those of the body of the shell.
Raised lines of growth occur at irregular intervals. Internal sur- face is smooth and lustrous, and shows the grooves corresponding to the external ribs, and also a very distinct microscopic structure, but is destitute of special radial lire.
Internally, the hinge-plate is narrow, thin, with a sharply impressed submarginal groove on eachend. The resilial pit is excavated in the margin of the hinge itself, and the anterior auricle has internal grooves corresponding to the external ribs.
Length of the largest examples, 6" ; height, 6-5™™.
Off the coast of Newfoundland in 67 to 72 fathoms.
Chlamys (A@quipecten) glypta Verrill. Pecten glyptus Verrill, Trans. Conn. Acad., v, p. 580, 1882. Dall, Proc. U. S. Nat. Mus., xii, p. 248, pl. viii, figs. 2, 3, 1889. ze Pecten Tryoni Dall, Bullet. Mus. Comp. Zool., xviii, p. 438, 1887 (t. Dall). PuLateE XVI. figs. 7-11.
When young this species has strong, ‘well-defined, angular radial ribs of nearly uniform size. In the old shells the grooves are occupied by several small ribs, and a secondary rib develops on each side of the keel of the primary rays; the ribs are all crossed by rather strong concentric sculpture (fig. 8, 11) which is sometimes so coarse as to give both the ribs and grooves a rough appearance, There are about three small, free pectinidial teeth in one of our specimens, but Dr. Dall states that they are absent in his specimen (P. Tryoni). Wt is allied to C. opercularis L., of Europe, and to C. purpurata of the west coast of South America,
Off the eastern coast of the United States in 69 to 156 fathoms. Off North Carolina in 124 fathoms (Dall). This has been taken only in small numbers, and mostly dead and broken.
if A. £. Verrili—Study of the family Pectinide. 7
~I
Chlamys (A®quipecten) irradians (Lam.)
a Pecten irradians Lam., Anim. sans vert., ed. 1, 1819; ed. 2, vol. vii, p 143. Gould Invert. Mass., ed. 2, p. 199, fig. 496. Veerrill, Invert. Vineyard Sd., ete., p. 401 [695], pl. xxxii, fig. 238. Rathbun, Fishing Industries of the U. States, sect. 1, vol. i. p. 509, pl. 255, fig. 8; Ingersoll, op. cit., sect. 5, vol. ii, pp. 565-581, 1887. Jackson, Phylogeny of the Pelecypoda, Mem. Boston Soc. Nat. Hist, vol. iv, pp. 333-350, cut 37, pl. xxvii, fig. 9, pl. xxviii, figs. 1-10, 12, 13, 1890 (young).
Pecten concentricus Say, Journ. Acad. Nat. Sci. Philad., vol. ii, p. 259, 1822.
PLATE XVI. fig. 6. PLATE XVIII. figs. 1-5. Puare XX. figs. 1-4, €, 6a.
Some of the young stages of this species have been figured by Mr. Jackson in the work quoted above, and several of his figures have been reproduced on our plate xx (figs. 1-5), We also give new figures of some of the more advanced stages on plate xviii, as well as a figure of the nuclear shell (fig. 1). These figures illustrate well the changes that the young shells undergo. Fig. 2 of pl. xviii shows three successive stages as indicated by prominent lines of growth. Even in the latest of these stages the posterior auricle is but little differentiated, but pectinidial teeth are already developed. This figure also illustrates well the origin of the radial sculpture. No camptonectes sculpture has been noticed at any stage. Radial ribs begin to appear on the right valve when 2™™ in diameter, and on the left valve when about 1°5"™.
The transverse incisions of the hinge-plate are usually very dis- tinct in shells less than 20™™" in diameter, and often persist in the adult. Occasionally examples are found that show them with unusual distinctness. A specimen in which they are very well developed is figured on pl. xvi, fig.6. The valves gape a little below the auricles and at the ends of the auricles, to which the tentacles and ocelli extend.
The foot (pl. xx, fig. 6) in alcohol is obliquely turned to the right; it is somewhat enlarged at base, with a deep byssal slit extending about half its length, beyond which it is contracted somewhat, and slightly enlarged at the end, which is divided into two lobes by a short median groove. The labial palpi are broadly triangular, strongly grooved on the apposed surfaces. The oral tentacles are large and consist of numerous contorted lamellae, much webbed together, and united with the bases of the labial palpi. There appear to be two (or two groups) on each side and somewhat in front of the mouth. The gills are large and of the type usual in this family. The pallial tentacles are very numerous and very unequal ‘in several rows. Two or more of the larger ones correspond to
78 A. E. Verrili—Study of the family Pectinide.
each rib, while the outer ones are simall and much crowded. The ocelli are numerous, unequal in size, the larger ones-at first alternate evidently with the primary tentacles and are opposite the external sulci. The ocelli are very brilliant in life.
This species is abundant in shallow water, especially of bays and sounds, from Cape Cod to Florida. It is used extensively as food, The adductor muscle is the only part utilized for this purpose.
Chlamys (Placopecten) Clintonius (Say) Ver. See p. 69.
Pecten Magellanicus ? Gmelin, Syst. Nat., p. 3317, 1788, (a bad and misleading name if applied to this species). Lamarck, Anim. sans vert., ed. 2, vii, p. 134. Gould, Invert. Mass., ed. i, p. 132. Dall, Bull. Mus. Comp. Zool., p. 216, 1886. a
Pecten Clintonius Say, Journ. Acad. Nat. Sci. Philad., iv, p. 124, pl. 9, fig. 2, 1824. Verrill, Trans. Conn. Acad. Sci., vi, I, p. 261, 1884. Rep. U.S. Com. Fish and Fish- eries, for 1883, p. 577.
Pecten tenuicostatus Mighels and Adams, Proc. Boston Soc. Nat. Hist., i, p. 49, 1841; Boston Journ. Nat. Hist., vol. iv, p. 41, pl. 4, fig., 7, 1842 (the young of the smooth variety). Gould, op. cit, ed. 2, p. 196, fig. 494. Verrill, Invert. of Vineyard Sound, ete., p. [696] 402, 1893.
Pecten princeps Emmons, Rep. N. C. Geol. Survey for 1858, p. 280, fig. 198 (fossil form).
Amussium Magellanicum H. and A. Adams, Genera Moll., ii, p. 555, 1858.
Pecten (Pseudamusium) Milleri Dall, Bull. U. States Nat. Mus., No. 37, p. 34, 1889 (the young).
Puate XVII. figs. 1-7. PLATE XX. figs. 1-Sa. PuLatTE XXI. figs. I-1la, 2, 2a.
When very young this species is nearly smooth on both sides, but when about 3-4™™ in length, it develops small, regular, raised ribs over the whole surface of the upper valve, and usually at both ends of the lower one, with intervening camptonectes sculpture. (Pl. Xvii.)
These small ribs increase in number, but not much in size, until the shell is 2 inches or more in diameter. After that size, in the _ greater number of shells of the northern variety, they decrease in size till the upper valve becomes nearly smooth, or has only linear riblets. But in some northern examples and in many of those taken in deep water south of Long Island, the small ribs continue regularly over the whole surface of the upper valve, and are more or less roughened by the raised edges of small concentric lamelle or lines of growth, sometimes becoming more or less finely cancellated. There are no corresponding internal ribs, except in extreme exam- ples of this variety, near the margin, and the edge of the shell is usually only slightly crenulated by the riblets, while the edge of the lower valve is essentially plain and sharp. (PI. xvii. figs. 5-7.) -
A, FE, Verrill—Study of the Jamily Pectinide. 79
This ribbed variety agrees with the fossil form described by Say as Pecten Clintonius. For the northern, nearly smooth form, the name given by Mighels (tenuicostata) may be retained as a varietal name,
This shell, when full grown, has the margins gaping considerably at both ends, below the auricles, much as in Amusium. But the anterior sutural line is less sinuous than in most species, and the bys- sal notch is small and not excurved.
__ The gaping is less marked in the young shells, but is evident even in those of small size (pl. xvii, fig. 5). The young shells show three to five pectinidial teeth, but these are usually obsolete in the adult. Transverse incisions on the hinge-plate are evident in the young shells.
The muscular and pallial scars are rather complex and unlike in the two valves (pl. xxi, figs. 2, 2a), but part of them are usually only faintly marked, even in large specimens. The internal surface has a peculiar subnacreous luster and a crystalline structure, somewhat like that often shown on frosted window glass, or that on the sur- face of tin-plate, after heating.
The large shells, about 6 inches in diameter, still retain the habit of swimming, though often partly covered with barnacles, hydroids, bryozoa, sponges, etc., but they doubtless swim much less actively than do the young ones, which are very lively. Still, I have often seen the large ones leap out of buckets of sea-water in which they had been placed for conveyance. The adults apparently do not ordi- _narily form a byssus, but there is no evidence that they are unable to do so, if necessary.
The foot of this species (pl. xx, fig. 8) has been described on page 69. The palpi (pl. xxi, figs. 1, 1a) are large and broad, triangular, broadly attached at the bases which run back so as to embrace the _ anterior ends of the gills; their apposed surfaces are strongly trans- _ versely ribbed.
_ The oral tentacles are very large and complex, arborescently much branched, so that when the branches are contracted in alcohol they "appear somewhat like the heads of caulitlowers. The branches are short and crowded and more or less webbed together, while the lat- ) eral tentacles are attached by webs to the bases of the palpi. There are five groups or clusters of these tentacles; the two larger pairs are lateral and anterior to the mouth; the odd one, which is sim- ilar but rather smaller, is in the median line behind the mouth.
The pallial Be acles and ocelli are very numerous (pl. xx, figs. e c7, 8a). The ocelli differ more or less in size ; they are separated by
GUA thie Veronese: iene be pinks avid
80 A. E. Verrill—Study of the family Pectinide.
one to three larger, and numerous smaller tentacles. The latter dif- fer greatly in size; the outer ones are small, very numerous, and much crowded. The guard-tentacles (fig. 8a) are shorter, conical, — and alternate in two or more rows.
This large species occurs abundantly in many localities off the coast of the United States, north of Cape Hatteras, from just below low-water mark to 60 fathoms, and is sometimes taken below 100 fathoms. It is used as an article of food to a considerable extent in New England.
Hyalopecten dilectus Verrill and Bush, sp. nov.
Shell small, thin, fragile, strongly undulated, slightly oblique, with the ventral margin broadly rounded ; dorsal margin straight.
In the right valve, the anterior auricle is elongated, with a deep angular notch beneath ; the posterior auricle is shorter, with a prominent dorsal angle, which is less than a right angle, owing to the emargination of the posterior end ; in the left valve the anterior auricle is broad, with its posterior angle nearly rectilinear, and it forms a right angle with the dorsal margin; the posterior end has a slightly prominent angle and a posterior emargination in both valves. The anterior auricle is marked by several fine, rough, radial ridges, which are more numerous and stronger on the left valve. The beaks are a little prominent and project somewhat above the dorsal margin. |
The surface of both valves is covered with broad and rather regu- lar undulations, most prominent on the left valve ; the undulations are crossed by regular, well-spaced, thin, raised radial lines, becom- ing finer and more crowded at the ends of the shell ; they are nearly obsolete in the right valve, being indicated only by microscopic striz.
The interior is strongly undulated and marked by very distinct radial grooves in the left valve, and by faint ones in the right valve. Resilium small, central ; color, dirty white.
Length, 8™™" ; height, the same.
One living specimen (No. 52,539), from station 2570, off Martha’s Vineyard, in 1813 fathoms, 1885.
This species is figured in an article by Verrill and Bush, sent to the Proceedings of the U. 8. National Museum, several months ago, but not yet published.
It is closely allied to P. fragilis of Jeffreys, and resembles pretty nearly his figure (Proceedings Zodlogical Society of London, plate 45, figure 1), which probably represents a species distinct from the original type described by him. It may be identical with our shell.
A, E. Verrill—Study of the family Pectinide. 81
Our shell differs decidedly from the original description of P. fragilis. Moreover we have obtained from several stations a shell of similar size, which appears to be the true fragilis, as it agrees closely with the original description. It also closely resem- bles P. pudicus, described by Mr. Smith, from east of Marion Island, in 1375 fathoms. (Chall. Exp.)
We have but one specimen, which is somewhat broken at the margins.
Hyalopecten fragilis (Jeffreys) Verrill.
Pecten fragilis Jefi., Ann. and Mag. Nat. Hist, 1876, p. 424; Proc. Zool. Soc., London, 1879, p. 561, pars (not the fig., pl. 45, fig. 1). Verrill, Trans. Conn. Acad,, vi, p. 232, 1885; Expl. by the Albatross, p. 577, 1885.
This is one of the most simple shells known to me in this family.
The shell is very thin, hyaline, distinctly undulated, but ‘not otherwise sculptured. No camptonectes sculpture is visible. The edges are very thin, apparently not bevelled. The hinge-plate is thin and delicate, without cross-lines, and with a single faint sub- marginal rib, parallel with the margin ; ligament very thin ; resil- ium very small, in a triangular pit; no auricular crure. The ante- rior auricle is well developed, with a deep byssal notch, but without pectinidial teeth ; the posterior auricle is undeveloped. Accord- ing to Friele there are no visible ocelli. This species, which we consider the true P. fragilis Jeffreys, was taken at the following stations :
Station 2115, in 843 fathoms; 2215, in 578 fath. ; 222], in 1525 fath. ; 2234, in 816 fath. ; 2710, in 984 fath. It occurs off the Euro- pean coasts and northward to the Arctic Ocean, in 656 to 1750 fath.
Pseudamusium simile (Laskey).
Pecten similis Lask., Mem. Wern. Soc., i, p. 387, pl. viii, fig. 8, 1811. Forbes and Hanley, Brit. Moll., ii, p. 293, pl. lii, fig. 6, pl.S, fig. 1, animal. Jeffreys, Brit. Conch., ii, p. 71; v, pl. xxiii, fig. 5.
PLATE XVII. figs. 8, 8a.
The shell in this small species is thin, translucent, nearly smooth, symmetrical, broadly rounded, longer than high. The valves are only slightly unequal ; the right valve is a little flattened, and its edge turns up a little so as to fit tightly against the edge of the
TRANS. Conn. Acan., VOL. X. JUNE, 1897. 6
82 A, E. Verrill—Study of the family Pectinide.
upper valve. The sculpture, when any is present, is nearly the same on both valves.
The auricles are straight and rather short, the posterior one is broad and obtuse-angled. The byssal notch is small and the pecti- nidial teeth are obsolete, or nearly so. There are no internal ribs; the inner surface is slightly pearly. The hinge-plate has only one longitudinal rib, just below the ligament ; it is crossed by numerous fine, transverse incisions, often more developed than in allied species. The sculpture, when evident, consists of fine lines of growth, and sometimes of very delicate, straight, radial riblets, without campto- nectes sculpture. 4
According to Jeffreys, the ocelli are few in number, about six or eight in the front row and about twice as many in the second row. It swims about for a long time and then quickly anchors itself by a small byssus.
It occurs in 15 to 200 fathoms on the northern European coasts.
Camptonectes Grceenlandica (Sow.) Verrill.
Pecten Grenlandicus Sowerby, Thes. Conch., p. 57, pl. 13, fig. 40. G. O. Sars, Moll. Reg. Arct. Norveg., p. 23, pl. 2, figs. 4, a-c, 1878.
The shell is rounded, inequivalve, very thin, hyaline, nearly smooth, often with a violet iridescence when fresh. The left valve
is covered, even from the nucleus, with fine microscopic campto-
nectes sculpture, in the form of thin, raised, divergent riblets, more or less irregular and wavy, most visible by translucency. The left valve sometimes has, also, fine radial striz and delicate lines of growth. The margins are thin and smooth, that of the right valve turns up a little against the other, which is larger, and the valves close very tightly, so that anteriorly there is scarcely any visible gape, even at the byssal notch, or at the end of the auricle. The byssal notch is well-marked and the pectinidial teeth are small and few. The byssus is probably very slender.
The auricles are not oblique and are nearly equal. The hinge- plate is very thin ; the single longitudinal ridge is scarcely visible.
A row of six or seven ocelli can be seen through the shell in alco- holic specimens.
Off Newfoundland, in 130 to 224 fathoms. Off northern Europe. and in the Arctic Ocean.
A, EF. Verrill—Study of the family Pectinide. $3
Cyclopecten pustulosus Verrill. (See page 70, fig. 1.)
Pecten pustulosus Verrill, Amer. Journal Science, vol. v, p. 14,1873; Trans. Conn. Acad., vol. iii, p. 50; vol. v, p. 581, pl. 42, figs. 22, 22a; vi, p. 261; Expl. by the Albatross, p. 577, pl. xxxi, figs. 142, a, b, 1885.
Pecten (Pseudamusium) imbrifer (pars) Dall, Bull. Mus. Comp. Zool., xii, p. 220, 1886 (not the figures), (non Loven).
PLATE XIX. figs. 3, 4.
In this species the ligament is thin; there is a narrow, simple, cardinal ridge, with faint tranverse denticulations and strie. The chondrophore is small, excavated in the thickened margin of the hinge-plate in both valves. There are no auricular crure.
The nucleus projects above the hinge-margin in the upper valve, but not in the lower. The posterior auricle is small in both valves, but has a prominent outer angle. The byssal notch is small and narrow, with its margin incurved or sinuous ; there are no pecti- nidial teeth. The valves close pretty tightly, leaving only a slight subauricular slit. The inside of the valves often has a subnacreous luster. There is no flattened submarginal area in either valve.
This species has been referred to Propeamusium Hoskynsi by Jeffreys, and to C. imbrifer by Dall. It never has internal ribs, like the former, which it resembles in sculpture. From the latter, as originally described by Loven, and re-described and figured by G. O. Sars, it differs especially in the character of the ornamentation of the left valve.
The European form has the vesicles much less crowded in each radial row, subconical and mucronate, while in ours they are usually closely crowded and often even in contact in the radial rows ; their form is either rounded or elliptical, with the longest diameter in the direction of the concentric lines, and the summit is evenly rounded, showing no tendency to the subconical or mucronate form. When perfect they resemble small blisters with the surface roughened or minutely granular under the microscope ; when broken or worn off, as frequently happens, the basal part remains in the form of semi- circular or semi-elliptical, imbricated, arched scales, usually consid- erably elevated above the surface and connected by very delicate concentric raised lines. The anterior auricle of the left valve is roughened by the close, elevated, concentric lines and by from four to six well-marked radiating ridges or ribs, upon which the concen- _ tric lines form regular elevated arched projections, often so crowded as to be imbricated ; in some young examples, like the one figured, the concentric lines on the auricle are less crowded, and only two or
84 A. EF. Verrill—Study of the family Pectinide.
three of the radial ribs are developed ; in such examples the vesi- cles on the body ot the shell are relatively fewer, larger, and more rounded and much less crowded in the radial series. In some spec- imens the posterior margin below the auricle is nearly smooth or marked only by the fine lines of growth, but in others, especially the larger specimens, this region is covered by rather sharp granules, some of which, toward the ventral margin, change to pointed scales in crowded radial rows. The raised concentric lines on the right valve are generally more or less appressed or sometimes imbricated ; toward the ventral margin some of them show very fine microscopic granulations, which are much less distinct than in P. imbrifer, as figured by G. O. Sars.
Off the eastern coast of the United States, and northward to Newfoundland, in 99 to 547 fathoms.
This species is evidently distinct from that figured by Dr. Dall (Blake Mollusca, plate 4, figs. 4a, 4b), under the name of P. imbrifer. His figured specimen apparently belongs to our C. subimbrifer.
Cyclopecten subimbrifer Verrill and Bush, sp. nov.
Pecten Hoskynsi Verrill, Trans. Conn. Acad., vol. v, p. 581, pl. xliv, fig. 11, 1882 (non Forbes).
Shell small, inequivalve, white or grayish white, translucent, length and height nearly equal. Dorsal margin straight ; anterior auricle in the left valve rather large and broad, the outer end obtusely rounded and covered with small, close radial ribs and crowded con- centric ridges ; posterior auricle much smaller, with one to three faint radial ridges and many concentric raised lines; outer end forming less than a right angle, with a slight, incurved notch below. In the right valve the anterior auricle has a similar radial sculpture and the byssal notch is rather deep and narrow.
The dorsal outlines of the body of the shell form rather less than a right angle; the ventral margin forms nearly a semicircle, and forms obtuse angles where it meets the dorsal outlines. Umbos a little prominent, with beaks small, acute, smooth, and projecting beyond the margin of the hinge. The surface of the left valve is covered with slightly raised concentric lines, which are raised into small arched scales ; these are often semicircular, but more frequently somewhat angulated or V-shaped ; they are usually separated by intervals about equal to their breadth, These scales are arranged in about 40 or more radial rows and decrease regularly in size to the umbo, where they are replaced by thin and slightly raised radial
A. E. Verrill—Study of the family Pectinide. 85
lines, crossing the stronger and more elevated concentric lines, but not rising into points.
The posterior dorsal area, below the auricle, is nearly smooth, except for the fine lines of growth, but sometimes shows minute granules. The right valve, which is smaller than the left, is covered by fine, thin, close, concentric raised lines, which sometimes show microscopic striations.
The anterior auricle is decussated by six to eight or more, small radial ridges, which are crossed by the raised concentric lines ; the latter rise into sharp scales at the dorsal margin ; the small posterior auricle has fine concentric lines and only two or three faint radial ridges.
Off the eastern coast of the United States ; 121 to 312 fath.
The figures of this and various other species were forwarded, sev- eral months ago, to the U. 8. National Museum, to illustrate an arti- cle in its Proceedings.
Cyclopecten leptaleus Verrill.
Pecten leptaleus Verrill, Trans. Conn. Acad., vol. vi, p. 232, 1884; Expl. by the Albatross, p. 577 [75], 1855.
Dr. Dall has expressed a doubt as to whether this species is distinct from P. imbrifer. In addition to the original description, it should be stated that the concentric lines are somewhat thickened and elevated, even where thinnest, and that the beaded character is quite unlike anything found in P. imbrifer or allied species. The beads are closely arranged, elliptical in form, and most elevated at the center, the elevation being often greater than the diameter ; their summits are smooth or glossy, so that when viewed from above, under a lens, each often appears to have a central cavity. The radial lines are comparatively very thin and delicate and not visible except when considerably magnified. The beaks are more acute than in P. imbrifer, and the nucleus is smaller and smoother.
Off the eastern coast of the U. States; in 142 fathoms off Cape Hatteras.
Cyclopecten nanus Verrill and Bush, sp. nov.’ Puate XVI. figs. 12—12c. Shell small, the breadth and height about equal; the valves are nearly equal in size and convexity. Dorsal hinge-margin rather long
1 Described and figured in an unpublished paper sent to Proc. U.S. Nat. Mus.
86 A. E. Verrili—Study of the family Pectinide.
and straight; auricles relatively large and broad, both ends sub- truncated or a little convex in the left valve, and forming nearly a right angle with the dorsal margin ; anteriorly not well differentiated from the body of the shell. In the right valve the anterior auricle is narrow and somewhat more elongated and obtusely rounded at the end, with a sharp, angular, byssal notch beneath it and separated from the body of the shell by a narrow groove. .
The dorsal margins of the body of the shell are nearly straight and form rather more than a right angle. Ventral margin broadly rounded, nearly semi-circular, forming a very obtusely rounded angle where it joins the dorsal margins. Umbos a little prominent, with a small, smooth, rather acute, incurved beak, which usually projects a little above the hinge-margin.
The surface of the left valve is everywhere thickly covered with fine, almost microscopic, radiating striz, which become a little more
distinct on the anterior auricle ; on some parts of the shell very thin,
slightly raised, concentric lamellz or lines of growth are often dis- tinct, especially on the anterior auricle, where they become closer and more regular ; in crossing the radial striations they produce a microscopic decussation, which is often quite regular. The sculpture on the posterior auricle, though finer, is similar, but in many speci- mens the surface is nearly smooth or marked only by very fine radial strie. The body of the shell of the right valve is smooth, except for very fine concentric lines; on the anterior auricle are three to six or more distinct radial ridges, which are roughened by conspicuous lines of growth; the margin below the byssal notch is entire; the posterior auricle is nearly smooth. .
The internal hinge-plate is thin in the middle, but relatively broad on each auricle, and is crossed by numerous fine, well marked, trans- verse incisions ; these are much more conspicuous than in most of the related species, whether young or old. The resilial pit is small, rounded, and situated just under the beak. There are no internal lire. The inner surface is smooth and glossy, although in fresh specimens the external radiating lines show through by translucency.
The ground-color of the right valve is yellowish or grayish white, with more or less numerous light yellowish brown, or reddish brown spots and blotches, and sometimes with irregular patches of opaque white; right valve white, sometimes with a few yellowish-brown spots. Some specimens are nearly destitute of spots.
The right valve is less convex than the left, and its ventral edge does not quite reach that of the opposite valve; the umbo is less
a Tee |
a
ro
A. E. Verrili—Study of the family Pectinide. 87
prominent ; the beak is less acute and scarcely projects beyond, and often falls short of the hinge-margin, but the inequality is less marked than in most of the allied species.
Length of one of the largest specimens 7™; height 6"™™; dorsal hinge-margin 4™™.
It was taken in considerable numbers. It is so distinct from all the other species of our coast that a detailed comparison is unneces- sary.
Off the eastern coast of the U. States, opposite Chesapeake Bay and Cape Hatteras, in 43 to 132 fathoms. Although very small, this species seems to be adult.
Cyclopecten simplex Verrill, sp. nov. Puate XVI. fig. 1: Puare XIX. figs. 1, 2.
Shell well rounded, thin, compressed, hyaline. Auricles large, prominent at both ends, unequal, in the right valve the anterior end is considerably prolonged, with a deep byssal notch, and the poste- rior end is less prolonged with a rather deep emargination or sinus ; in the left valve both ends are shorter and angulated. The left valve is nearly smooth; the right valve is covered with fine, crowded, concentric, incised lines, and has faint radii on the anterior auricle. Resilial pit small. Transverse incisions of the hinge are fine and vermiculated or irregular.
Height, 4-4" ; breadth about the same. West Indies, U.S. Fish Com.
Propeamusium thalassinum (Dall) Verrill.
Pecten fenestratus Verrill, Proc. U. States Nat. Mus., ili, p 403, 1881 (non Forbes).
Amusium fenestratum Verrill, these Trans., vy, p. 582, 1882.
Pecten (Pseudamusium) thalassinus Dall, Bulletin Mus. Comp. Zool., Blake Exp., Pelecypoda, vol. xii, p. 221, 1886.
Puate XIX. figs. 5-7. This species has been very well described by Dr. Dall’, but has not been figured. The larger specimens and some of the young not more than 4 or 5™™ in diameter have an internal, raised, opaque white, radial rib, on each end, below the auricles, within the concavity of the shell, and a
_~ 1Dr. Dall, in the work quoted, has called the upper valve the right and the lower
' fa!
valve the left, and consequently has reversed the anterior and posterior ends, in the descriptions of this and several other species of Pectinid. He informs me that this was done inadvertently, and should be corrected.
88 A. EF. Verrill—Study of the family Pectinide.
similar but smaller one, on each end or on the posterior only, above the auricular ridge, forming “ auricular crur.”
In many of the younger shells one or both of the internal ribs may be lacking, as in other species of Propeamusium.
The cardinal ridges are rather broad, flat, and crossed by numerous very small transverse incisions and denticles. These are more strongly developed than in many allied species, but less so than in - Cyclopecten nanus V. and B., and several other species examined. Some examples of Chlamys irradians excel in this respect (see pl. xvi, fig. 6).
The resilial pit is peculiar, for in the upper or left valve it pro- — jects distinctly beyond the hinge-plate, as a spoon-shaped process, but in the lower valve it is excavated in the sunken and oblique median notch of the hinge margin.
The byssal notch is broad and shallow, its margin not incuryed, and without any pectinidial teeth. The auricles are very unequal; — the posterior one has a prominent angle.
The upper valve is elegantly cancellated, and often mottled with yellowish white on a reddish ground-color. The lower or right valve is white and covered with strong, even concentric ridges or lamine. The lower valve has a marginal flattened area which fits closely against a similar, flattened, submarginal area of the upper valve.
Those in alcohol had the soft parts poorly preserved, but there were about six rather distant, black ocelli visible through the trans- lucent shell.
Off the eastern coast of the United States in 43-317 fathoms, West Indies in 84 to 450 fathoms, Blake Exped.
This species is here referred to Propeamusium on account of the two internal lire or ribs within the body of the shell, for the pres- ence of such lire is the distinctive character of this genus. In other respects it agrees about as well with Cyclopecten. These two groups are very much alike in form, sculpture, and the unlikeness of the two valves. The present species is, therefore, intermediate between the typical forms of the two groups.
Analytical Key to the genera of Pectinide.
The following table is not strictly a natural one, as to the sequence of the genera, but is as nearly natural as could be made consistently with convenience of use. :
A. FE. Verrill—Study of the family Pectinide. 89
I.—Hinge-plate with a central resilial pit.
A,—Hinge-plate without lateral series of marginal pits, resembling resilial pits, but often with small transverse incisions.
A.—Shell with very unequal valves: the right, or lower, valve very convex with a strongly incurved beak; left valve nearly or quite flat, shutting closely inside of the edge of the right valve. Both valves with strong primary radial ribs and internal liree; edges scalloped. Animal not adapted for swimming.
a. Hinge with the cardinal ribs plain or nearly so. Pecten (restr.) aa. Hinge with the cardinal ribs strongly transversely incised or pitted. Neithea (sub-gen.)
AA.—Shell with the valves not very unequal; the upper or left valve the most con- vex, when any difference exists. Sculpture various. Animal adapted for swim- ming, at least when young.
B.—Shell internally fluted or smooth, without special radial ribs developed independ- ently of external sculpture.
C.—Hinge-plate with several large distinct, nearly transverse, tooth-like processes.
4 Shell with large external and internal radial ribs. Byssal notch obsolete.
: Pallium. CC.—Hinge-plate without transverse tooth-like processes. Sculpture various. D.—Hinge-plate with more than two cardinal ridges, either side of the resilial pit,
the lower ones divergent. Radial ribs large and more or less nodose. <A dis-
4 tinct byssal notch. Lyropecten.
DD.—Hinge-plate with only one or two cardinal ridges, the upper one parallel with
the dorsal margin and bounding the ligamental groove; the second, when
a present, more or less oblique. Sculpture various. _ E.—Shell, when adult, permanently attached by the right valve, which is more or less h distorted. A deep byssal notch. b. Right valve directly attached by a shelly deposit. Hinnites. ___bb. Right valve attached by a modified byssus. Hemipecten. _ EE.—Shell free through life, or only temporarily attached by a byssus of thread-like 2 fibers. Byssal notch usually present.
_ F.—One or both valves with external primary ribs or riblets.
» External ribs small, nearly obsolete on right valve. No internal lire. Edges of valves not scalloped nor strongly crenulated. Placopecten. 7 aa. External ribs and internal lire: strong; edges of valves scalloped. Byssal notch
a well developed. __¢, Primary radial ribs and internal lire simple, not increasing with age by forking; shell scarcely oblique. Aquiphien, cc. Primary ribs and lire of various sizes, increasing in numbers with | by forking, or by the interpolation of new ones; auricles or shell usually oblique. a Chlamys (restr.) aaa. External and internal ribs simple, formed by corrugations of the shell; internal ribs not thickened. Leptopecten.
_ FF.—Both valves destitute of strong external ribs and internal thickened lira; edges
____ either smooth or slightly crenulated.
_ G.—Anterior auricle well developed, with a distinct byssal notch.
d. Valves with the edges simple or slightly crenulated, equal, meeting nearly evenly.
90 A. EF. Verrill—Study of the family Pectinide.
&
. Shell swollen, valves strongly convex, nearly smooth, not hyaline; auricles unequal, oblique. Pectinella,
ee. Shell compressed, valves little convex, often hyaline.
. Both valves undulated, and with fine radial sculpture or smooth. Texture. hyaline. Hyalopecten.,
jf. Valves not undulated, sculpture none, or of fine, radial riblets, or cancellated on
one or both valves, or vermiculated.
g. Shell nearly smooth, or with small radial riblets; divergent vermiculated sculp-
ture may be present, at least when young. Pseudamusium (sens. restr.) gg. Shell more or less hyaline or translucent; vermiculated divergent sculpture con- spicuous when adult, either with or without radial riblets or rows of scales ; auri- cles unequal. Camptonectes. Patliolum. dd. Valves with the edges smooth and unlike, that of the lower one flattened or bevelled and shutting against the upper one, seulpture on the lower valve con- sists of concentric raised lines or riblets; on the upper valve of radial riblets, or strize, lines of scales, or cancellations. Cyclopecten. GG.—Anterior auricle without a byssal notch. Shell thin, rounded, symmetrical. | h, Auricles of one valve, prolonged dorsally in distinct angles or points. Hntolium. . hh. Auricles not prolonged dorsally; sculpture radial on one valve. Syncyclonema. hhh. Auricles nearly equal, angular, not distinctly prolonged dorsally. Shell very thin, with fine concentric sculpture. Protamusiwum.
BB.—Shel!l with special internal, radial ribs, independent of external sculpture. Smooth or with delicate sculpture externally; not ribbed. Edges of valves not scalloped. Body of shell usually broadly rounded, not oblique.
7. Shell nearly closed at both ends; texture hyaline; auricles both well developed;
byssal notch and pectinidial teeth present.
j. Valves nearly equal; lower valve not concentrically sculptured; both valves
with fine radial lines; internal ribs numerous, simple, not differing from the shell in texture. Lissopecten. jj. Valves unequal; lower valve strongly concentrically sculptured > upper valve . with radial lines, or rows of scales, or cancellated; internal ribs few, well differ- entiated from the shell. Propeamusium.
zi. Shell large, thin, round and flat, gaping at both ends, usually smooth and polished, but not hyaline; auricles feebly developed; pectinidial teeth and byssal notch obsolete. Internal ribs strong, usually opaque white.
k, Two pairs of gills. Amusium. kk. One pair of gills. . Paramusium. AA,—Hinge-plate with a series of definite pits, resembling resilial pits, on each side
of the central pit.
i. Auricles small, not well defined. Shell smooth or with fine radial sculpture. —
Pernopecten.
Wl. Auricles well developed, angular. Sculpture concentric. Euchondria. Il.—Hinge-plate without a central resilial pit.
m. Shell nearly symmetrical. A series of small pits, resembling resilial pits, along
« ss,
the hinge-margin. Crenipecten. mm. Shell oblique, inequilateral, auricles unequal, ligament confined to a groove. Aviculopecten.
eT Ee ae hee Se a ae ee el re
A. EF. Verrill—Study of the family Pectinide. 91
List of genera, subgenera, and species discussed.
Synonyms are generally omitted. The few included are enclosed in parentheses. Numbers refer to pages; those where special descriptions are given are printed in italics.
Pecten (typical), 48, 44, 53, 54, 56. P. maximus, 54, 55, 56; P. dentatus, 57; P. hemicyclicus, 57 ; P. dubius, 55; P. Jacobeus, 56; P. pictus, 55; P. ziczac, 55; P. atavus, 57.
Neithea, sub. gen., 60. P. equicostatus, 60.
Lyropecten, 49, 63. L. nodosus, 63; L. subnodosus, 64; L. corallinoides, 64; L. noduliferus, 64.
Pallium, 51, 59. fe plica, 59.
Hinnites, 48, 59. H. Cortessi, 59; H. Adamsi, 60; H. pusio, 60.
Hemipecten, 48, 60. H. Forbesianus, 60.
Chlamys (typical), 55, 38. C. Islandica, 54, 55, 58, 72, 73; var. insculpta, 73 ; C. ornata, 59; C. exasperata, 59 ; C. costellata, 59,75 ; C. Benedicti, 59, 74; C. phrygia, 59; C. effluens, 58; C. madreporarum, 49; C. varia, 75.
Aiquipecten (sub. gen.), 59, 67. C. opercularis, 67; C. irradians, 41, 47, 59, 68, 76 and 77; C. dislo- eata, 59,68; C. glypta, 59, 68, 76; C. Antillarum, 59, 68; C. nucleus, 68; C. ventricosa, 59, 68; C. purpurata, 68; C. caurina, 68.
Leptopecten (sub. gen.), 69. C. Monotimeris, 69.
Lissopecten (sub. gen.), 49, 68. C. hyalina, 69.
Placopecten (sub. gen.), 61, 69. P. Clintonius, 42, 49, 58, 61, 69, 78; (P. magellanica, 78); var. tenuicostata, 79.
Camptonectes, 62.
» C. lens, 62; C. Grenlandica, 82.
Palliolum, sub. gen. or section, 65. P. Teste, 62, 65, 66; P. striatum, 62, 66; P. tigrinum, 62, 66; P vitreum, 43, 46, 65, 66.
92 A. E. Verrill—Study of the family Pectinide.
Pseudamusium (restr.), 60. P. exoticum, 60; P. simile, 46, 87; P. dispar, 61; P. pseuda- musium, 61; (P. glabrum, 61; P. hyalinum, 61; P. corneum, 1; P. natans, 61; P. tigrinum, 61). Pectinella, 68. P. Sigsbei, 68. Hyalopecten, 63, 71. H. undatus, 63, 71; H. dilectus, 62, 71, 80; H. fragilis, 63, 71, 80, 81; H. pudicus, 63, 71, 81. Syncyclonema, 62. P. rigida, 62. Cyclopecten, 61, 70. C. pustulosus, 70, 82; C. imbrifer, 70, 83; C. subimbrifer, 70, 84; C. leptaleus, 70, 85; C. nanus, 70, 85, C. reticulus, 71; C. simplex, 70; C. Culebrensis, 70; C. Murrayi, 71; © clathratus, 71 ; C. subhyalinus, 71; C. distinctus, 71; C. Ker- madeciensis, 71; C. orbicularis, p. 71. | Propeamusium, 64. P. inequisculpta, 64; P. thalassinum, 65, 87, P. Pourtalesia num, 65; P. cancellataen 65; P. Sayanum, 65; P. Holmesii, 65; P. Alaskensis, 65 ; P. Hoskynsi, 65; P. lucidum, 65; P. scitulum, 65; P. Torresi, 65; P. propinquum, 65 ; P. obliquum, 65. Amusium, 42, 49, 55, 37. A. pleuronectes, 55, 58; A. Mortoni, 57, 58; A. Japonicum, 58 ; A. Laurentii, 55; (A. Dalli, 57, 58). Paramusium, 52, 57, 72. P. Dalli, 52,72. Protamusiunm 62, 71, 73. P. demissum, 72; P. disciforme, 72; P. membranaceum, 72; P. illustre, 72 ; P. suleatellum, 72; P. obovatum, 72. Entolium, 62. E. cornutum, 62. Pernopecten, 63. P. limiformis, 63. Euchondria, 64. E. neglecta, 64. Orenipecten, 65. C. crenulatus, 65. Aviculopecten, 60. A. concavus, 60.
A, E. Verrill—Study of the family Pectinide. 93
EXPLANATION OF PLATES.
All the drawings on the following plates were made from nature by Mr. A. Hyatt Verrill, except figs. 5 and 6 of pl. xviii, and fig. 7 of pl. xix, which were drawn by J. H. Emerton, and figures 1-5 of pl. xix, which are copied from Dr. R, T. Jackson.
PLATE XVI. Figure 1.— Cyclopecten simplex V., sp. nov. Right valve, x 15.
_ Figure 2.— Chlamys Islandica, young. Left valve, x 8.
Figure 3.—The same, somewhat older. Right valve, x about 5 diameters.
Figure 4.—The same, var. insculpta, portion of the surface, near the margin. x 15.
Figure 5, 5a—The same variety. Portions of the sculpture of the disk of a young specimen, x 20.
Figure 5).—The same specimen. Portion of the sculpture nearer the umbo, x 40.
Figure 6.—Chlamys irradians. Part of hinge to show the transverse incisions (7),
_ the ligament (7), the resilial pit (vr), and the cardinal ribs (f), x 4.
Figure 7.—Chlamys glypta V. Inside of left valve, x 3.
Figure 8.—The same. Upper part of right valve of a larger specimen; p, pectini- dial teeth, x 4. ;
Figure 9.—The same. Hinge of a small specimen, right valve; c, crural rib; 4, crural denticle; J, ligament; 7, cardinal ribs and transverse incisions ; p, pectini- dial teeth and byssal notch; 7, resilial pit, x 4.
Figure 10.—The same. Two of the internal bicarinate radial ribs, at the margin, x 4.
Figure 11.—The same. Sculpture of a large specimen close to the margin, x 8.
Figure 12.— Cyclopecten parvus V. and Bush. Right valve, x 8.
Figure 12a.—The same. Hinge of left valve, x 15.
Figure 12b.—The same. Sculpture of the left valve near the umbo, x 100.
Figure 12c.—The same. Sculpture of the left valve near the margin, x 100.
Pratt X VIL.
Figure 1.—Chlamys (Placopecten) Clintonius, young, left valve, x 8. Figure 2.—The same, somewhat older, right valve, x 6.
Figure 3.—The same, slightly older, interior of right valve, x 6. _ Figure 4.—The same. Hinge of right valve of a more mature example, x 3. _ Figure 5.—The same, young. End view of posterior end of a specimen showing
unusually large riblets. To illustrate extent of gaping and inequality of the valves, x 9.
Figure 6.—The same. A portion of the sculpture of the left valve of an unusually
strongly sculptured example. x 15. Figure 7.—The same. Portion of the sculpture from near the margin of the left valve of another young specimen, showing the camptonectes sculpture, x 24; Ta,
‘y * 7b, other parts of the same valve more enlarged.
Figure 8.—Pseudamusium simile. Left valve, x 6. Figure 8a4.—The same. Right valve, x 6.
94 A, E. Verrill—Study of the family Pectinide.
PLATE XVIII.
Figure 1.—Chlamys irradians. Nuclear region of a young specimen showing prodissoconch, or veliger shell, x 60.
Figure 2.—The same. Right side of a very young shell, x 15.
Figure 3.—The same. Right side of a somewhat larger specimen, showing twe stages of growth, x 6. :
Figure 4.—The same. Left side of a similar young specimen, x 6.
Figure 5.—Hyalopecten uwndatus. Left valve of the type-specimen, x14. By J. Emerton. (See Trans. Conn. Acad., vi, p. 444.)
Figure 6.—Camptonectes (Palliolum) vitrea. Right side of a mature specimen, J. H. Emerton.
Figure 7.—The same. Hinge of right valve, x 6.
Figure 8.—The same. Hinge of left valve of the same specimen, x 6.
Figure 9, 9a.—The same. Sculpture of the left valve of a perfect specimen, x 27.
Figure 10.—The same. Right side of a very young specimen, x 16.
Figure 11.—The same. Left side of a somewhat larger example, x 12.
Figures 12, 12a.—Dorsal views of the right and left valves of the nuclear shell of} aL small specimen, x 60. ;
Figure 13.—The same. Side view of the nuclear shell, x 60.
Figure 14.— Camptonectes (Palliolum) striata. Inside of right valve, x 6.
Figure 14a.—The same. Sculpture of the left valve, x 24.
PLATE XIX.
Figure 1.— Cyclopecten simplex Ver. (sp. noy.). Interior of left valve, x 15.
Figure 2.—The same. Interior of right valve, x 15.
Figure 3.—Cyclopecten pustulosus. Right side of a young specimen, showing the nuclear shell, x 12. ;
Figure 4.—The same. Left side of a slightly older specimen, x 12.
Figure 5.—Propeamusium thalassinum. Left side, x 12.
Figure 6.—The same. Interior of a right valve, x 9.
Figure 7.—The same. Hinge of left valve of the same example, x 9.
Figure 8.—Propeamusium inequisculpta. Interior of a left valve, x 8.
Figure 8a.—The same. Sculpture of the left valve, x 15. -
Figure 9.—The same. Interior of a right valve of a young specimen showing the beginning of the internal ribs, x 9.
PLATE XX.
Figure 1.—Chlamys irradians. Very young spat, much enlarged; p, nuclear or veli- ger shell; f, foot; g, rudimentary gill; ”, heart; ad, adductor muscle; a, anus; m, mantle; e, ocelli; ¢, tentacles. After Jackson. Figure 2.—The same. Left side of a very young living specimen, much enlarged; /, foot; «, an opening through which water was ejected, showing an unusual pro- trusion of the mantle border (perhaps accidental), Figure 3.—The same, Right side of a very young shell, much enlarged; n, byssal notch and first pectinidial tooth; p, p, prodissoconch or nuclear shell,
. ‘ wy sie oe . , i ofa . ‘ ‘ r i ne ee Ne ee ee ee a ied, . oe Re eH a EY RE RE LE ES os re Se le re ee ASE ae ame
lr Yas a
a a
A. FE. Verrill—Study of the family Pectinide. 95
re 4.—The same. Diagramatic view of gill-filament of adult; a, direct border; a’, reflected border; s, septum uniting the two parts of about every fifteenth filament; bl, blade.
Figure 5.—The same. Diagram of section through shell (s) and adductor muscle (ad); 1, resilium; m, mantle; mw, guard or inner reflected edge of mantle; gf, guard tentacles; #, marginal or pallial tentacles; ¢, ocelli.
e 6.—The same. View of the interior when the shell and mantles are spread open; about twice natural size; a, mouth; b, rectum and anus; ¢, abdominal mass; d, labial palpi; ¢, oral tentacles; /, foot; g, inner gill, showing reflexed margin; m, inner surface of mantle; m’, its reflexed inner margin or guard, and _ guard tentacles. From an alcoholic specimen.
Figure 6a.—The same. Portion of the mantle margin more enlarged: 0, o', primary and secondary ocelli; ¢, ¢, primary and secondary tentacles.
Figure 7.—Chlamys (Placopecten) Clintonius, Front view of a small living and active specimen about one-halfnatural size; m, inner mantle margin or guard.
‘Figure 8.—The same. Foot of a larger alcoholic specimen, x 2; r, byssal groove; 5, terminal bilobed disk.
Figure 8a.—The same. Portion of the mantle-margin of an alcoholic specimen, x 4; m, inner surface of mantle, showing blood spaces and nerves; m’, reflexed inner margin; m’’, guard tentacles; ¢, marginal pallial tentacles; 9, ocelli.
Figure 9.—Chlamys Islandica. The foot of an alcoholic specimen, x 6; 7, byssal groove; s, terminal slit or groove.
PrArTe XX.
sure 1.—Chlamys (Placopecten) Clintonius. Part of animal of an alcoholic speci- ‘men, x 6; a, mouth; e, oral tentacles; d,d’, outer and inner labial palpi; g, ante- rior end of gill.
a gure la.—The same, a part of one of the oral tentacles, more enlarged, to show mode of branching.
Figures 2, 2a.—The same. Right and left valves showing muscular scars, # natural ' size; a, a, scars of adductor; a’, portion of adductor with finer fibers; 8, pallial line; c, row of small pallial scars, of irregular occurrence; d, d, more or less lobed dorsal margin of adductor scar ; the pedal muscle is attached obliquely to the adductor muscle of the left valve at d, in fig. 2a; ¢, e’, special pallial muscles below the auricles (subauricular muscles), situated in the region where the pallial guard become narrow and the guard-tentacles obsolete; , byssal notch ; I, ligament; ¢, transverse cardinal rib; r, resilium.
Figure 3.—Chlamys Islandica. Part of animal, x6; a, mouth; e, oral tentacles; d, d’, outer and inner labial palpi; g, anterior end of gill; 7, byssal groove of foot; 8, terminal slit of foot.
gure 4.—Pallium plica. Hinge of right valve, x 5.
ADDENDA AND ERRATUM.
The two following groups were omitted from their proper places in the list of divisions that have been proposed :
Leptochrondria Bittner, 1891. Type, P. (Z.) @olicus from the Trias of Asia Minor. Proposed as a subgenus of Pecten. Pm Deitopecten Morris, 1892. Type, D. Illawarensis M., from the Permo-carboniferous. of Queensland. Described as intermediate between Pecten and Aviculopecten.
Page 78, line 22. For Mulleri read striatus. .
III.—ReEvision oF THE Marine GAsTROPODS REFERRED TO CycLos- TREMA, ADEORBIS, VITRINELLA, AND RELATED GENERA ; WITH DE- SCRIPTIONS OF SOME NEW GENERA AND SPECIES BELONGING TO THE ATLANTIC Fauna oF America. By KarHarinr JEANNETTE BUsH.
In studying the descriptions and figures of the many species of marine gastropods from various and widely separated localities, which have been referred to Cyclostrema, Adeorbis, Vitrinella, and related genera, I soon found that there was great variation in their form, texture and sculpture. It seemed to me that the most satis- factory and permanent results toward eliminating this confusion would be gained by publishing the original descriptions of the various genera, with their types, for convenience in deciding the relations of the species already known.
In the present article, therefore, I have given descriptions of the genera, arranged chronologically, with lists of the species belonging to the marine fauna of eastern America, which have been correctly or incorrectly referred to them, together with several new genera and species. When possible, I have also given figures of the type species of each genus.
In carrying on these investigations I have been greatly aided by Professor A. E. Verrill, of Yale University, Dr. W. H. Dall, of the U. S. National Museum, and Mr. E. A. Smith, of the British Museum.
Cyclostrema Marryatt, 1818. Type, C. cancellata Marryatt. West Indies?
PLATE XXII. figs. 4, 4a.
“ Oyclostrema.”’
“ Character genericus.” _ “Testa depressa, perspectivo-umbilicata ; apertura circularis.”
“©. cancellata Marryatt.”
“Tab. x, Figs. 3, 4.”
“©. testa alba, lineis longitudinalibus et transversis elevatis decussantibus, inde cancellata. Habitat ut
“ Apertura labiis cancellatis, cancellis tranversim striatis.”
“T found this beautiful little shell among a collection of chiefly West Indian Shells. According to the Linnean system, it would come under the genus Ziwrbo,” etc., etc. Trans. Linn. Soc. London, xii, p. 338, 1818.
Trans. Conn. Acap., Vou. X. JULY, 1897.
7
98 K. J. Bush—Marine Gastropods referred to
These inadequate descriptions and want of knowledge of the operculum, animal and odontophore of the type, together with the — fact that Marryatt stated that Helix depressa and Helia serpuloides were referable to the same genus, have doubtless led to the greater part of the confusion into which the genus has fallen.
As its true position cannot at present be decided, it seems best to follow the authors who have studied similar species. §S. P. Wood-— ward (1851-6) placed the genus in the family Turbinide, with Liotia Gray (1850), and Collonia Gray (1850), as subgenera of Delphinula (Roissy) Lam. The figure he gives as cancellata, however, does not agree very closely with those of Marryatt and the locality of — the species is given as the Philippines. H. and A. Adams (1858) placed it next Liotia in the family Liotiinze and added two sub- genera, Cynisca H. and A. Adams and Serpularia Romer, 1843 (? Spira Brown, 1838). See pages 107,108. The original figure that is given as cancellata, however, represents a very different species, as do also those given by Chenu in his Manual, and by A. Adams in — Sowerby’s Thes. Conch., copied by Tryon.
The following extract from a letter from Mr. E. A. Smith, of the British Museum, under the date of May 7th, corroborates this opinion.
“The shell figured by H. and A. Adams (Gen. Rec. Moll., pl. xlv, — f. 6a. and in Sowerby’s Thesaurus, iii, pl. 255, figs. 5, 6), in my opinion is perfectly distinct from Marryatt’s Cyclostrema cancellata, and I have long ago noted this.
Kiener’s Delphinula cancellata (Icon. Coq. Viv., p. 10, pl. tv, f. 10) = Kieneri Phil. is, I consider, Marryatt’s species. It is curious that both employed the same specific name. I do not know that the shell figured by Adams has been renamed,” ete.
The only reference that I have found regarding this difference is_ in Tryon’s Manual, x, p. 89, 1888, where it is suggested that the Philippine specimens, as figured by Sowerby, may prove to be iden- tical with Cyclostrema eburnea Nevill, from the Bay of Bengal. The figures 27-30, as given on pl. 31, look very unlike, and I very much doubt the accuracy of such a combination. For the species figured as C. cancellata by H. and A, Adams and copied by Chenu, I propose the specific name pseuwdocancellata, I very seriously ques- tion its rightful reference to the genus Cyclostrema, however, but — such a question must be left to the future study of the authentic specimens. Whether or not the description given by A. Adams, P. Z. 8., p. 41, February, 1850, refers to the same species that is
Cyclostrema, Adeorbis, Vitrinella, and related genera. 99
figured, is also an open question. Tryon’s copies of: Sowerby’s fig- ures do not agree perfectly with the figure in Gen. Rec. Moll., and I am unable to consult the original ones.
The numerous small, rather thin, nearly smooth species from deep- water which have been referred to Cyclostrema by several authors, belong to several quite distinct genera. These have few convex whorls, forming a moderately elevated spire; the aperture circular, not modified, as in affine Verrill = proxima Tryon, or nearly circu- lar and modified on the body-whorl, as in Dalli Verrill; the peritreme simple, entire, or more or less modified ; and the umbilicus deep, but varying from one of moderate size to a scarcely percepti- ble chink.
Much careful study of the other numerous and varied forms, which have also been erroneously referred to the genus, is necessary before
_ their true position can be determined.
The family name Cyclostrematide, constituted by Fischer, should now be restricted to forms like the true C. cancellata Marryatt, and perhaps may prove to be closely related to, or synonymous with, Liotiine (Adams and Chenu), Liotiide (Tryon), or Delphinulide (Fischer and Dall).
The following extract from a letter from Dr. Dall, of the U.S. National Museum, under the date of April 14th, shows the possible relation of the genus.
“JT had never looked up Marryatt’s original figures before, but took this occasion for doing so. What he had was, it seems to me, a Solariorbis (as I have been calling them) very much like some I have been describing from the southern Miocene and Pliocene,” ete.
List of species belonging to the marine fauna of eastern America which have been referred to Cyclostrema :
Cyclostrema cancellata Marryatt, C. Schrammii Fischer, C. Beauii (Fischer) Tryon = VU. bicarinatum Guppy, C. angulata A. Adams, (@. Dalli Verrill (non C. fulgidum (Jeffreys, 1883) Dall), and ay ornatum Verrill, C. cingulatum Verrill = C. Verrilli Tryon, (. affine Verrill = C. prowima Tryon (non C. trochoides _ (Jeffreys MSS.) Sars.), C. diaphanum Verrill, C. valvatoides Jeffreys, C. (Granigyra) limata Dall, C. turbinwm Dall, C. pom- pviolye Dall, C. cistronium Dall, C. granulum Dall, C. cancellatum _ (Jeffreys) Dall (non Marryatt), C. tuberculosu (dV’Orbigny) Tryon, C. excavata Watson = C. subexcavata Tryon, C. diaphana (d’Orb.) Poulsen (non Verrill).
100 K. J. Bush—Marine Gastropods referred to
Delphinoidea Brown, 1827. Type, D. serpuloides (Montagu). Devonshire coast, England.
PLATE XXII. figs. 1-10.
“Spire depressed, surface smooth, divested of spinous processes ; aperture orbicular, or nearly so, and not enveloping the body volu- tion.” Ill. Cat. Gt. Britain, p. 19, 1838? (2d ed.)
This genus has been considered by most authors as a synonym of
Cyclostrema, but the very small, nearly smooth species referred to it by Brown are very unlike the highly sculptured type of that genus. Even as restricted and described in the 2d ed. this is a hetero-
genous genus without a designated type, with two subdivisions —
(1.—Volutions Dextral. IJ.—Volutions Sinistral), containing in all four described and figured species, in the following order :
1. D. unispiralis (Montagu). 2. D. depressa (Montagu). 3. D. serpuloides (Montagu), and 4. D. resupinata (Montagu). The last being the only representative of the second subdivision.
These with several other of Montagu’s species of Helix were included under Delphinoidea in the 1st ed., 1827. The first species,
unispiralis, is described by Brown as follows :
“Shell glossy white and opaque, with one volution, umbilicate on both sides ; aperture circular. Diameter scarce a line.”
“ Found at Sandwich, and is very rare.”
This was without doubt a veliger shell, the true specific relations
of which it is impossible to determine without comparing it with similar species from the same region.
Fleming, in 1828’ (Hist. Brit. An.), constituted the genus Skenea and referred depressa Montagu, and serpuloides Montagu, to it. In the 2d ed. Brown restored these two species to his genus Delphi- noidea and gave references to Fleming’s article. Helix depressus Montagu is now considered the same as Skenea planorbis (Fabricius),” which stands as the type of Skenea. See plate xxiii, figs. 5, 8, 8a.
1 Gray, P. Z.8., 1847, p. 152, gives 1824.
® Skenea planorbis (Fabricius) is common on the rocky shores of our eastern coast from Long Island to Greenland. It is a minute shell without sculpture, covered with a conspicuous amber or delicate horn-colored epidermis, of about three well-rounded whorls, with deep sutures, coiled nearly in the same plane so that the spire is but
little elevated. The umbilicus is large, revealing all the whorls, with rounded walls. |
Aperture circular; peritreme simple, continuous. Operculum thin, of a delicate horn-
color, circular, with central nucleus of about six whorls defined by an indistinct spiral
line. Radula with a series of seven unequal, distinctly serrate, curved teeth in each
row. For figures see G. O. Sars, Moll. Reg. Arct. Norv., pl. vi, f. 15; pl. xviii, f. 23, 1878.
Cyclostrema, Adeorbis, Vitrinella, and related genera. 101
The animal of serpuloides was carefully studied and described by Wm. Clark in 1855 (Brit. Mar. Test. Moll.), and the species was placed by him in the genus 7rochus, A figure of it was given by H. and A. Adams in 1858 (Gen. Rec. Moll.), under the name Cyclos- trema serpuloides, but these authors stated that “Should the smaller British species require to be separated from the more typical forms they will take the name of Delphinoidea Brown.” Jeffreys, in 1865 (Brit. Cone., iii), described and figured it as Oyclostrema serpu- loides because “ Delphinoidea is both superfluous and heterogenous.” This decided statement has doubtless been the cause of much of the misunderstanding of Cyclostrema of more recent authors. G. O. Sars in 1878 (Reg. Moll. Arc. Nory.), restored the species to Delphin- oidea, but described and figured the young of Margarita helicina (Fab.) as serpuloides (Supl., p. 346).
Delphinoidea is unquestionably a very unfortunate selection for a generic name, as it is used to designate an ordinal group of Dol- phins, ete.
Delphinoidea Brown (sens. restr.).
Shell small, white, consisting of a few convex whorls coiled nearly in the same plane so that the spire is but little raised ; suture deep ; umbilicus rather large, deep, with rounded walls, showing all the whorls, and not defined by a carina; aperture oblique, nearly cir- cular, slightly angulated above, not modified by the body-whorl ; peritreme simple, thin, entire, but slightly attached ; columellar edge very slightly or not at all flattened.
Type, D. serpuloides (Montagu). Described and figured by Jef- ireys, I; C., ili, p. 290, pl. vii, f. 3.
Specimens (No. 9450) from Guernsey, England, in the Yale Museum, presented by the Rev. Canon A. M. Norman, measure about 1°5"™ in diameter. They are rather thin, white, somewhat shining, of about three convex whorls, forming a very low spire with deep sutures. Very fine microscopic, raised, revolving lines commence on the periphery of the whorls ; these become more conspicuous and widely separated on the base and umbilical region. The aperture is nearly circular, with a slight angle where it touches the body-whorl.
_ The peritreme is simple, continuous, but along the columellar mar- gin the edge seems slightly flattened. Some specimens have a deli- cate raised line just within the aperture.
102 KK. J. Bush—Marine Gastropods referred to
HT, serpuloides Montagu is given by Fischer as an example of the section Daronia under his Cyclostrematide. I place it with the Vitrinellide.
The species described and figured by G. O. Sars (p. 345, pl. 34, figs. 6a-6d), as Cyclostrema areolatum, seems to me a true Delphi- noided.
Adeorbis 8. V. Wood, 1842. Type, A. subcarinatus (Montagu). South coast of Devon. F
PLATE XXII. figs. 5, 9.
Mr. Searles V. Wood, in 1842, in his “ Catalogue of the Crag Mollusca,” published in the Annals and Magazine of Natural His- tory, London, ix, proposed the genus Adeorbis for a group of small shells which he characterized as follows :
“ Whorls subdiscoidal, volutions few, peritreme sharp, inner lip sinuous, umbilicus large and deep.”
He failed to mention a type species as such, but described and figured A. striatus Wood, alone of the five species which he referred to the genus, only naming the other four, all of which are new, except A. subcarinatus (Montagu), which is placed as fourth in the list. In 1848, in his ‘ Monograph on the Crag Mollusca,” he redefined the genus as follows : ;
“Shell generally small, suborbicular, depressed, with a few nearly discoidal and rapidly increasing volutions, umbilicus large and deep ; peritreme entire and nearly continuous, slightly interrupted by the previous volution, deeply sinuated on the inner side, having’a minute or incipient sinus at the upper part of the aperture near the junction of body-whorl.”
Although comparing the genus with Skenea he placed it next to Margarita, Ue again failed to mention a type species, but placed - A. striatus Wood first, as before, and described and tigured four other species, omitting A. subimbricatus, which in 1842 he included in the genus with a mark of doubt, and added A. pulchralis Wood, which in 1842 he had identified as Margarita helicina Wood. He also mentioned that his A. striatus is probably the same as Valvata striata of Philippi.
Mr. 8. P. Woodward in his “ Manual of Conchology,” published in 1851-6, defined Adeorbis and mentioned and figured as the type, A, subcarinatus (Montagu), placing it next to Cyclostrema, as a sub- genus of Delphinula.
Hf. and A, Adams, in 1858, in their “Genera of Recent Mollusca,”
Cyclostrema, Adeorbis, Vitrinella, and related genera. 103
defined the genus and figured A. subearinatus (Mont.) as an exam- ple, placing it next to Cyclostrema.
Chenu, in 1859, defined the genus, mentioning and figuring A. subcarinatus (Montagu) as an example, adding also figures of A. striatus Wood.
Mr. Jeffreys, in 1865, seems to have been the first author to make any generic distinction between the several species of Wood. In “ British Conchology,” vol. iii, p. 315, he described as Trochus Duminyi Requien, a recent shell found at Bonegal Bay, identifying it as Delphinula Duminyi Requien, 1848 = Adeorbis striatus Wood, 1848 = Valvata? striata Philippi, 1836, and mentioning A. supran- itidus Wood, and A. tricarinatus Wood, as fossil varieties. The specific name striata had been preoccupied by Trochus striata Linné. He made a special section under the genus Zrochus for the reception
of such forms, as “ they are very distinct from the typical Adeorbis subcarinatus (Montagu), the operculum of which is paucispiral and horny, with lateral nucleus.” The section is defined as follows :
“C. Very small, circular, nearly flat-spired with exceedingly wide and open umbilicus. Circulus.”” See p. 110.
He mentioned that the animal of Duminyi is unknown, but de- scribed the operculum as follows :
“Operculum circular with about a dozen volutions which wind spirally and gradually and converge to the centre.” A very poor figure of the species is given in vol. -v, pl. 62, fig. 5.
Fischer, in his ‘“‘ Manuel de Conchyliologie,” defined Adeorbis, and stated that the operculum is horny, spiral and excentric, giving and figuring A. subcarinatus as the type.
Tryon, in his “Manual of Conchology,” 1883, defined the genus, mentioning and figuring A. swbcarinatus (Mont.) as an example, but stated that the operculum is “ shelly, subspiral.”
There is little doubt that Wood intended A. striatus to stand as the type, but, as he included three quite distinct forms in the genus and failed to mention a type species, according to definite rules of nomenclature, the type given by the author who first separates the species of the genus has to stand as the one to be adopted.
A, subcarinatus (Mont.), of which Ihave several examples (No. 9428) before me, collected at Guernsey, England, and presented to the Museum by the Rey. Canon A. M. Norman, is a very small (the larg- est is about 2°5™" in diameter), moderately thick, white shell, of about three abruptly enlarging whorls, so coiled that the suture ends at the periphery of the preceding whorl. The whorls are well
104 K. J. Bush—Marine Gastropods referred to
rounded above but decidedly angular below, with the base flat and appearing somewhat concave. The umbilical region is of consider- able size, with a comparatively small but deep opening, which is not in the center of the base, owing to the abrupt enlargement of the whorls. The aperture is large, very oblique, somewhat angular, so that when the shell rests on the base the spire is considerably tilted. The sculpture consists, on the body-whorl, of four to six unevenly elevated, unequally separated carinz, which are crossed by very con-— spicuous, elevated, irregular, oblique, somewhat sinuous lines in the direction of the lines of growth ; these considerably roughen the entire surface from the suture over the base well up into the umbil- -
icus. The suture is defined by a much roughened carina which rests _
well up on the preceding whorl. The nucleus is regularly coiled, smooth and shining. Some specimens are very glassy and some are stained: by oxide of iron.
Nothing seemed to be known of the animal or operculum until — 1865, when Mr. Jeffreys described the latter (B. C., vol. ili, p. 317) as “paucispiral, horny, with lateral nucleus,” and suggested the near relation of the genus to Solarium. In 1869 (B. C., vol. v, p. 216), he mentioned a specimen as having been found with the animal which is of a very red color. In 1885 (‘‘ Proceedings of the Zodlogical Society ” of London, for January, p. 40), he again mentioned the animal and referred to the descriptions given by Mr. Duprey in the “ Annals and Magazine of Natural History,” London, for October, 1876. Mr. Duprey stated that he had had several animals alive for some time and had studied their habits and characters with great care. The following is an abstract of his description :
Animal white, with a pinkish hue, semi-transparent, easily con- tained in the shell. Snout rather long, extensile, cloven at the tip. Tentacles long, extensile, blunt, diverging. Eyes very small, at out- ward base and a little behind the tentacles. Foot slightly notched in front, square behind. Gill comblike, on right side of body.
Unfortunately the odontophore of the type has never been studied, so that the true relation of the genus is still doubtful.
List of species belonging to the marine fauna of eastern America which have been referred to Adeorbis -
Adeorbis Beauii Fischer, A. Orbignyi Fischer, A. Adams Fischer, A. lirata (Verrill) Dall, A. supranitidus Wood = A. trilix (Bush) Dall, A. striatus Wood = sp.?, A. elegans A. Adams, dA. (Clathrella) naticoides Dall, A. nautiliformis Holmes = Cochlio- lepis parasitica Stimpson, A. olivaceus (Verrill) Dall, A. costulata- (Moller) Verrill = Mélleria costulata (Moller) Jeffreys, A. trornatum (VOrb.), A. eyclostomoides Pfr. |
Cyclostrema, Adeorbis, Vitrinella, and related genera. 105
Separatista Gray, 1847 (not described), A. Adams, 1850. Type, S. separatista (Chem.) Dillw. = S. Chemnitaii A. Adams. Philippines.
“Shell orbicular, somewhat discoid, the first whorls contiguous, the last disunited ; aperture wide-spreading, angulated ; umbilicus large, infundibuliform, the whorls visible within as far as the apex.” P. Z. S.,"p. 45, 1850.
Two species were described, S. Grayii Ad., from the Cape of Good Hope; and S. Chemnitzii Ad., from the Philippines, which was figured by H. and A. Adams, 1858, and is considered as the type of the genus.
The following extract from a letter from Dr. Dall, under the date of April 23d, explains the apparent confusion in the names which have been applied to the type:
“Gray’s Separatista was founded on Turbo separatista of Chemn. (x, figs. 1589-90), which was named S. chemnitzii by Adams (1850), who adds a second species 8S. Grayi. Gray (P. Z.S., 1847, p. 136) cites Turbo helicinus Gmel., as a name for the former, but this was a lapsus penne for T. helicoides Gmelin, who had, however, another T. helicoides, so that this one will retain the name of (Zurbo) separa- tista applied to it by Dillwyn (1817), to which Adams’ name will fall in synonymy.”
As nothing is known of the operculum or soft parts, the true posi- tion is very doubtful. H. and A. Adams placed it in the subfamily Rapanine ; Fischer, as a questionable subgenus of Trichotropis, but Mr. Dall, who states that: he has examined the type in the British Museum, placed it in the family ? Adeorbide. He constituted a new section Haloceras' for Separatista cingulata (V.) Dall, the young or immature form of which was described by Prof. Verrill as Cithna cingulata. The family Adeorbidz can, however, only be applicable to the genus Adeorbis and related forms, whose true posi- tion is undeterminable until the odontophore of the type, swbcarin- atus, can be studied.
Vitrinella C. B. Adams, 1850. Type, V. helicoidea C. B. Adams. Jamaica. PLATE XXIII. figs. 9, 9a.
“Shell turbiniform, vitreous, minute, with a large orbicular aper- ture, either umbilicated or with the umbilical region deeply and widely indented.”
1 Bull. Mus. Comp. Zodl., xviii, p. 277, 1889.
106 K. J. Bush—Marine Gastropods referred to
“The form of the aperture would place these shells in Turbo or Margarita. The want of an umbilicus excludes three of the species from the latter. The operculum is unknown ; but as it is extremely improbable that thin, vitreous, almost transparent shells should have solid calcareous opercula, we may assume it to be horny, which will exclude the species from Turbo, regarding this genus as best charac- terized by the operculum. The most widely umbilicated species approximates in form to Skenea, and might be mistaken for a depressed umbilicated species of Helix. In texture, the nearest approach among the kindred genera is in Margarita arctica — Leach,” etc. Monograph of Vitrinella, Amherst, Mass., Feb., 1850. —
The above quotation shows the extended sense in which Mr, Adams intended his genus, Vitrinella, to be taken, and he after- wards added several species from Panama.
The original list included five new species from Jamaica, without mentioning any as a type, given in the following order: V. hyalina, V. interrupta, V. megastoma, V. tincta and V. helicoidea. None of these were figured until 1888, when figures of a species, named V. helicoidea, made from a specimen, were published in Tryon’s — Manual, x, pl. 34, figs. 40-41.
P. P. Carpenter, in 1855-57 (Mazatlan Mollusca, p. 236), men- — tioned the differences in the form of the many species referred to this genus by C. B. Adams and himself, but did not restrict it by naming any one of the above species as a type, and only stated that ““megastoma is an Ethalia” and “ that the indented species accord better with Hthalia and Teinostoma.” He also stated “there are specimens of V. tincta, V. interrupta, V. valvatoides, Teinostoma minuta, and Lthalia megastoma in the Cumingian Collection.” H. and A Adams, in 1858 (Gen. Rec. Moll.), were the next authors to define the genus, but gave a figure of valvatoides for an example ;_ this species, however, not being in the original list, cannot stand as the type.
I propose V. helicoidea as the type and restrict the genus to small, more or less hyaline, low-spired shells of few convex whorls, having a moderate-sized, deep umbilicus; nearly circular, oblique aperture, with simple more or less continuous peritreme, modified on the body-whorl into a more or less conspicuous glaze, which may be absent in the young; columellar margin often flattened in the — adult, having the appearance of being thickened, and angulated at the lower, outer edge.
I have a group of such shells from shallow-water, off Cape Hat- -
Cyclostrema, Adeorbis, Vitrinella, and related genera. 107
teras, N. C., represented by V. helicoidea C. B. Adams, V. Tryoni Bush, V. diaphana (@Orb.), and V. carinata (d’Orb.)
As the family names Cyclostrematide and Adeorbida, if used at all, are applicable to other relations, I introduce Vitrinellide to distin- guish this group and extend it to include all small, more or less hya- line, non-nacreous species, varying in form from those having a low, little raised spire and large umbilicus, like Cireulus, to the higher spired, smaller umbilicated species, like Zissospira, and those with closed umbilicus, like Zharsiella. Taking the form of the umbilicus, the aperture, and peritreme, as the principal distinguishing charac- ters of the several genera, such as :
Delphinoidea Brown, 1827, Vitrinella C. B. Adams, 1850, Cireu- lus Jeffreys, 1865, Ganesa Jeffreys, 1883, Granigyra Dall, 1889, Tharsiella Bush, 1897, Lissospira Bush, 1897, Leptogyra Bush, 1897.
The correctness of such a grouping depends entirely on the future study of the opercula, animals, and odontophores.
List of species belonging to the marine fauna of eastern America which have been referred to Vitrinella:
Vintrinella hyalina C. B. Adams, V. interrupta C. B. Adams, V. megastoma C. B. Adams, V. tincta C. B. Adams, V. helicoidea C. B. Adams, V. striata (dVOrbigny) Tryon, V. anomala (dOrbigny) Tryon, V. (Episcynia ?) multicarinata (Stimpson MSS.) Dall.
Cynisca H. and A. Ad., 1858. Type, C. granulata A. Ad. Philippines.
“ Shell depressly turbinate ; umbilicus wide and deep, perspective, surrounded by a spiral callus; whorls with transverse, granular ribs ; aperture circular; inner lip straight, outer lip thickened, lirate within, continued posteriorly on the penultimate whorl beyond the inner lip.” Gen. Rec. Moll., p. 406, 1858.
Type, C. granulata, not figured.
In Ann. and Mag. Nat. Hist., vol. viii, 1861, A. Adams, under the genus Cynisca, stated: ‘In our Gen. Rec. Moll., vol. i, p. 406, my brother and myself proposed a subgenus of Cyclostrema under the name Cynisca. This was founded ona shell in Mr. Cumings’ collection, which I described as Cyclostrema granulata (P. Z. S., 1853). The great peculiarity consists in the aperture, which is something like that of Stoastoma, the inner lip being straight and i outer lip being continued. posteriorly on the penultimate whorl beyond the inner lip.”
He added two species, Delphinula australis Kiener, which has more recently been referred to the subgenus Liotina Munier-Chal-
108 K. J. Bush—Marine Gastropods referred to
mas, 1877, as its only living representative, and described the new — species Japonica, which in 1863 he stated was the Liotia pilula Dunker, and transferred it to the genus Collonia Gray.
I very much doubt the near relation of this genus to Cyclostrema. The peculiarity of the aperture is much like that seen in Leuco- rhynchia Crosse, 1867, type, L. Caledonica Crosse (Journ. de Conch., Xv, p. 320, pl. 11, f. 4), and Craspedostoma Lindstrém, 1884, type, — CU. elegantulum Lind, (Fischer’s Manuel, p. 831, f. 588).
Morchia A. Adams, 1860, non Albers,! M. Meyer, or Von Martens. Type, M._ obvoluta A. Ad. St. of Korea, 26 fathoms. ’
“ Shell obliquely ovate, widely umbilicate, depressed, convex from above, plane or flattened from beneath ; whorls rapidly increasing, the last dilated, covering all the volutions nearly to the apex. Aperture oblong, oblique, subhorizontal, dilated or expanded below, — angulated above ; peritreme continuous, thickened (énerassato).”
Type, If. obvoluta A. Ad. Figured in Thes. Conch., copied by — Tryon.
“Shell small, opaque, white, angulated at the periphery, orna- mented with crowded incremental striz# ; umbilicus wide, crenulated at the suture.”
“This curious little genus most nearly resembles Zeinostoma, but the base is not covered with a callus, and the mouth is not produced. — The last whorl embraces the others, as it does in Neritula and — Cyclops. Both Mérchia and Teinostoma, however, together with — Vitrinella, are not nacreous,” etc., ete. Ann. Mag. Nat. Hist., v, — p. 801, 1860. P. Z.S., p. 74, 1863.
The affinities of this genus are very doubtful, but the thickened peritreme would prevent its being placed with the Vitrinellide.
Daronia A. Adams, 1861. Type, D. spirula A. Ad. Philippines, Serpularia Rémer, 1843 (? Spiva Brown, 1838), H. and A. Adams, Gen. Ree. Moll., p. 406, 1858.
“Shell orbicular, discoidal, evolute ; spire depressly concave | whorls rounded, disunited ; aperture circular ; peritreme contin-— uous.”
Type, D. spirula A. Ad. Figured P. Z.8., 1850, and Thes. Conch., copied by Tryon, The original figure differs considerably from those in Thes. Conch., as copied by Tryon, q
' Morchia Albers, 1850 = Macrocyclis Beck, 1837, pro parte. -Morchia M.-Meyer 7 1860 = Burtinella Mérch, 1861. (Fischer's Manuel, pp. 456 and 692.)
Cyclostrema, Adeorbis, Vitrinella, and related genera. 109
In 1861 (Ann. Mag. Nat. Hist., vol. viii, p. 244), A. Adams stated: “The very remarkable shell described by me as Cyclostrema spirula (P. Z. S., 1850) is neither a Spira nor a Serpularia and the name of the subgenus may therefore be changed to Daronia.”
I have been unable to consult Sowerby’s Thes. Conch., 1864. Fischer used Daronia as a section of Cyclostrema, but his conception of that genus was in its most extended sense.
Serpularia was used by Munster (1840) for a genus of annelids, and by Rémer (1843) for a group of shells, some of which have been found to be annelid tubes. Fischer, however, retained it as a questionable genus (Manuel, pp. 716, 830), and gave S. centrifuga Rémer, as an example.
Spira Brown, 1838.
“Shell smooth, nearly globular or semiovate ; spire small in pro- portion to the size of the body, and depressed ; aperture enveloping the body. 2d ed. Ill. Cat. Gt. Britain, p. 20.
This genus was proposed by Brown for a group of seven of Mon- tagu’s species of Helix, all of which, with the exception of variegata, he had included in his genus Delphinoidea, 1827. They are doubt- less only larval shells.
Fischer recognized the genus in Journ. de Conch., pp. 45 and 51, 1877, but 1 find no mention of it in his Manuel, nor do I find that it has been recognized by other authors.
Tubiola A. Adams, 1863. Type, 7. cornuella A. Ad. St. of Korea, 63 fathoms.
“Shell somewhat unrolled or loosely coiled; whorls simple, round, with concentric striz ; aperture somewhat circular, peritreme con- tinuous, margin sharp, entire.”
Type, 7. cornuella A. Ad. Figured in Thes. Conch., copied by Tryon. ,
“Shell somewhat discoid, oblong-ovate, thin, yellow-brown, widely umbilicate, apex raised ; whorls 34, rapidly enlarging, round, the last free at the peritreme ; with conspicuous incremental strie ; aperture oblique, transversely ovate ; peritreme continuous, sharp, entire, expanded.” Ann. and Mag. Nat. Hist., v, p. 412, 1860.
The only change that is made in the description of the more adult specimen is in the color, which is given as “ dirty white.”
“Tn this species, which I described from a young individual as a Skenea in Ann. Mag. for 1860, the whorls are disunited, but the volutions are not rolled on the same plane as in Daronia spirula.”
mr 2. S., p. 74, 1863.
110 K. J. Bush—Marine Gastropods referred to
There seems to be considerable doubt in regard to 7. nivea, the other species described by Adams under this genus. In| P. Z. S., for 1850, he described under Cyclostrema, as (Turbo) nivea Chemnitz, a species in the Cumings collection, which he stated — is the Delphinula levis of Kiener; and in 1863, under Zubiola he described as (Zurbo) nivea Chemn., a rare species which is not in the Cumings collection, nor in any other, and is not D. levis Kiener, and “agrees exactly with the original description and figures of Chemnitz.” The xévea of Adams is described as having “the last whorl large, dilated in front, round at the periphery bordering upon or spreading out at the peritreme; aperture oblique, sub-circular, — angulated below,” etc., etc.
As it is impossible for any one to settle such confusion without a careful study of the specimens, I use cornwella as the type of the genus. Zs
Fischer used Zubiola as a section under Cyclostrema, but unfortu- — nately cited serpuloides (Montagu) as an example, and did not men-— tion either of the above species.
The relation of the genus to Mérchia or possibly to Skenea can only be definitely settled by the comparison of authentic types. |
Circulus Jeffreys, 1865. Type, C. Duminyi Requien = striatus Wood = striatus Philippi. Sicily, The Crag, and Ireland. }
Puate XXIII. fig. 11.
“Very small, circular, nearly flat-spired with exceedingly wide and open umbilicus.” ;
“ Operculum circular with about a dozen volutions which wind spirally and gradually and converge to the centre.”
Type, C. Duminyi (R.) Jeffreys.
Described in B. C., iii, p. 315, and poorly figured v, pl. 62, f. 5.
In 1883 (P. Z. S., p. 94), Jeffreys redefined the genus as follows :
« Animal not known.” .
“ Shell coin-shaped or forming a circular, compressed disk, slightly nacreous or pearly ; mouth quadrangular with a continuous peri- stome; umbilicus very wide; operculum multispiral, as in other genera of Trochide.”
Specimens of Dwminyi from both Naples and France, in the U. 8. National Museum, have been loaned me through the courtesy of Dr. Dall. The above statement of their being slightly nacreous or pearly is very misleading and quite erroneous. The interior is very
Cyclostrema, Adeorbis, Vitrinella, and related genera. 111
smooth and shining and not at all pearly as understood in Mar- garita, etc. The mouth cannot be said to be quadrangular; the peritreme is not continuous in the young, and even in the adult is modified on the body-whorl into a very thin glaze. The operculum is very difficult to describe, as the whorls are not clearly defined. It is circular, very thin, of a delicate horn-color, with central nucleus. _ Around the center are two or three evenly separated circles (whether _ spiral or not, I cannot determine), outside of these are irregularly _ disposed wrinkles, with an occasional, very faint, more definite circle. _ Tencountered the same difficulty in studying the operculum of ¢rilix _ (even when mounted in balsam). Five whorls represented by circles could be counted, but their spiral structure was impossible to trace. Fischer placed Circulus as a subgenus of his Solanderia,' 1880, of _ the genus Gibbula Risso, 1826, among the Rhiphidoglossa. The relation of this genus to Vitrinella seems very close, and the i differences, although perfectly apparent when one has several species { from both to study, are difficult to define. It therefore seems best to use Circulus simply as a section of Vitrinella for those species which, like Duminyi, have the more sinuate columellar margin and very wide umbilicus showing all the whorls, It should be redefined as follows :
ae
Ai
_Circulus Jeffreys, 1865 (sens, str.).
Ay oe
Shell small, circular, depressed, not nacreous, of few more or less ~ convex whorls usually more or less grooved and carinated ; aperture
nearly circular, oblique, somewhat angulated below ; peritreme sim- _ ple, more or less continuous, in the adult modified on the body-whorl - into a very thin glaze which is absent in the young; umbilicus _ wide, the reverse of the spire ; operculum thin, light horn-color, 2 with central nucleus, number of whorls doubtful (five or more ?)
; \. Episcynia Mérch, 1875. Type, Z. inornata (d’Orbigny). Island of St Thomas.
“Shell hyaline, the carina with a double series of cilia, apex sim- ple, not inverted.” Synopsis Molluscorum Marinorum Indiarum
_ 1! Solanderia Fischer, 1880 (non Duchassaing and Michelotti, 1846) = Rossiteria _ Brazier, 1895 (type, Zrochus nucleus Philippi). Proc. Linn, Soc. N.S. Wales, ix, 2d
112 EK. J. Bush—Marine Gastropods referred to :
“Shell orbicular, convex, conoid, smooth, shining, white, carinated; spire conic with obtuse apex, of five distinct, convex whorls; um- bilicus small, smooth ; aperture oval. Diam. 3™"; alt. 2™™.”
Hist. L’ile de Cuba, Moll., 11, p. 67; atlas, pl. xix, figs. 25-27, 1853.
Prof. Mérch constituted the above as a subgenus under Archi- teetonica Bolten, 1798 = Solarium Lamarck, 1799.
Prof, Fischer placed it, with a mark of doubt, in the same relation ; but Dr. Dall (Bull. M. C. Z., xviii, pp. 273, 392, 1889) suggested that it might prove to be more closely related to Adeorbis (i. e. Cireulus) or to a Vitrinella (?) like V. multicarinata, which he described as having the epidermis produced into a fringe on the secondary carinz, — above and below the peripheral one. The hairy epidermis would exclude this-species from Vétrinella, p. 105.
Trachysma G. O. Sars, 1878 (Jeffrey's MSS.) Type, 7. delicatum (Philippi). Italian Tertiary and Lofoten Is., 200-300 fathoms.
“Shell globular, rather thin, similar to a Cyclostrema (i. e. Lisso- spira) but more delicate, aperture open or spreading out, peritreme simple, with a very thin, distinct edge.”
Type, Z. delicatum (Philippi) G. O. Sars.
“Shell thin, semipellucid, white, globular with short, little elevated spire, consisting of three and one-half convex whorls, the last large, evenly convex, the base not flattened. Suture deep. Aperture large, _ round-ovate with the outer-lip evenly arched and the columella slightly incurved. Umbilicus deep, not defined from the base (i. e. with rounded walls). Upper part scarcely shining, to the naked eye smooth, but seen under the microscope very finely sculptured, with numerous longitudinal and interrupted spiral lines. Diam. 1:1™™ 5 alt, =”
Var. expansa G. O, Sars.
“Shell more dilated, the last whorl wide and very large, the base slightly flattened, the aperture widely open and expanded in an unusual manner, the lip oblique and irregularly curved. Diam. basis 1-4™™; alt. 1:2™™.” Moll. Reg. Arct. Norv., p. 211, pl. 22) agendas 18d, 1878.
Although Mr. Jeffreys seems to have proposed the name Zrachysma for a species from Lofoten Is. which he identified as the delicatum — of Philippi, I fail to find any reference to such a genus or species in any of his later reports on the “ Porcupine Expedition.” Therefore G. O. Sars should stand as the authority for the genus.
He placed it with the Solariide, but his figures (18a-d) of the
-, , ‘
Cyclostrema, Adeorbis, Vitrinella, and related genera. 113
typical form would place the genus in close relation to Lissospira or possibly Leptogyra, although no epidermis is mentioned ; but the form of the aperture of the variety expansa (figs. 17a-b) is very different and much closer to that of his Adeorbis fragilis (figs. 19a-c), the generic relation of which is very doubtful.
Fischer placed the genus with the Adeorbide with a mark of doubt, while Tryon (Manual, x, p. 13) placed it as a synonym of Archytwa de Costa, 1869, which is said by him to have the same type, A, delicatum of Philippi (p. 87). I have been unable to consult the Ann. Mus. Naples, iii, in which the latter genus is described. Fischer placed this with Solarium with a mark of doubt, but gave A, catenulata de Costa for an example.
Tharsiella’ Bush, 1897. Type, 7. romettensis (Seguenza). Pliocene. Calabria and Sicily. Mediterranean, Bay of Biscay, etc., in 108-1093 fathoms.
“Shell globylar, solid and glossy ; peristome circular and continu- ous, but attached to the pillar on that side: base closed by a pad or thick testaceous layer in the adult, perforated in the young: opercu- lum chitinous or horny, and multispiral.”
“This genus differs from Cyclostrema (i. e. Lissospira) in the peristome being, although continuous, not free or detached from the rest of the shell, and in the umbilicus being closed instead of open in
| the adult,” etc., etc. P. Z.S., p. 92, pl. xix, tig. 7, March, 1883.
Fischer placed this as a subgenus under his Cyclostrema, but Mr. Dall (Bull. Mus. Comp. Zodl., xviii, pp. 261-3, 1889) thought it might prove to be a synonym of, or at most, a feebly characterized section of Hthalia ; but his conception of that genus is in its most extended sense. See p. 116. Moreover, Mr. Jeffreys may have had two forms under the name romettensis, for his figure 7 certainly more closely resembles a Lissospira than an Ethalia, so that I place the genus with the Vitrinellid until some knowledge of the animal and odontophore can definitely decide its position.
The specimen (No. 38244) from off Cape Hatteras, N. C., station 2115, in 843 fathoms, described by Professor Verrill (these Trans.,
vi, p. 201) as Tharsis sp., is unquestionably a small species of Natica.
The aperture is broad-ovate, somewhat produced ; the peritreme is modified on the body-whorl into a rather conspicuous glaze. and the
columellar margin is thickened and reflected over the umbilical chink,
with a slight median expansion.
1 Tharsis Jeffreys, 1883, non Giebel, 1847 (Pisces) ; nec Tharsus Leconte, 1862 (Insecta. )
Trans. Conn. Acap., VoL. X. JuLy, 1897. 8
114 K. J. Bush—Marine Gastropods referred to
Ganesa Jeffreys, 1883. Type, G. nitidiuscula Jeffreys. Off the Hebrides, 570 fathoms. ;
“Shell shaped like a Natica, thin, peristome continuous, free and separate in the young, but united to the periphery in the adult, spire having an oblique axis ; base perforated, not umbilicate ; operculum horny, multispiral.”
“Differs from Tharsis in the obliquity of the spire and perforation of the base at every stage of growth.” P. Z.S., p. 94, March, 1883.
Two very distinct forms were described and figured under this genus, neither of which were mentioned as the type. G. pruinosa, in general appearance, shape of aperture, and peritreme, agrees closely with Cyclostrema (i. e. Lissospira) affine, and differs chiefly in hay- ing a peculiar granular surface. Several species having the same peculiarity have more recently been dredged by the U.S. F. C. at con- siderable depths in southern waters, for which group Mr. Dall consti-. tuted the section Granigyra.
Therefore as the other species, G. nitidiuscula Jeffreys, will stand for the type of the genus, it will be necessary to make some changes in its definition.
G. nitidiuscula Jeffreys.
“Shell differs from G. pruinosa in being exactly globular, opaque, and glossy ; the sculpture consists of flexuous but slight and remote raised striz in the (direction of the) lines of growth; the last or body-whorl is not so disproportionately large ; the apex of the spire is depressed ; the mouth is angulated both above and below, and the umbilical chink is channelled. L. 125, B. 125.” P. Z. &, p. 94, pl. | xix, figs. 9-9a, Mar., 1883.
Ganesa Jeffreys, 1883 (sens. restr.)
Shell resembling a Zissospira in general appearance, but differing” in having the aperture modified by the body-whorl and angulated both above and below, and also in having the umbilical chink chan- neled and partly concealed by the columellar margin.
Operculum and animal unknown.
Fischer placed this genus as a subgenus of his Cyclostrema, but cited G. pruinosa for an example.
Until the odontophore is known, I consider it but a subgenus of Lissospira, for the reception of species which, like Lissospira Dalli, have the umbilical chink bordered along its outer margin by a raised rounded thread so that it appears channeled, and is nearly concealed by the more or less flattening and spreading out of the columellar margin of the peritreme. See also Cirsonella Angas, p. 120.
Cyclostrema, Adeorbis, Vitrinella, and related genera. 115
Granigyra Dall, 1889. Type, G. limata Dall. Off Cuba, 310 fathoms.
“Shell covered with small pustules or granules like those on Poromya or Plectodon.”
“This singular little shell is a typical Cyclostrema (i. e. Lissospira') in its conchological features, except for its granular surface. The latter recalls that of Poromya, but is finer and less regular.” Bull. Mus. Comp. Zodl., xviii, p. 395, 1889.
Specimens of G. limata Dall and G. spinulosa Bush in Professor Verrill’s collection have a small umbilicus or perforation; the aper- ture nearly circular, with a slight sutural angle, and the peritreme simple, entire, and but slightly attached to the body-whorl, so that the section will come under the new genus Lissospira, until some knowledge of the operculum and animal can be obtained, which may
give it generic distinction.
There is a group of small, white, solid, nearly smooth, porcella- nous shells which have been referred to Hthalia (since 1855-7), Teinostoma, Pseudorotella, Calceolina, Cirsonella, and Dillwynella. They have several features in common, agreeing especially in hay- ing the umbilical region wholly, or in part, covered with a callous deposit. It therefore seems probable that they may, upon further examination, prove to be closely related to each other. Their family relations, however, will be difficult to determine, as the operculum and animal even of most of the type species are unknown. The solid porcellanous character of the shells together with the callosity covering the umbilicus would indicate closer affinity to the genus Umbonium Link, 1807 (type, I vestiarium (Linné), sub-family Umboniine, H. and A. Adams), rather than to the thinner, more delicate, mostly semi-transparent species referred to the several genera belonging to the Vitrinellide. Therefore, instead of. placing the above genera chronologically with those already considered, I have, for convenience, grouped them as follows:
1 Heliciella Costa, 1861, non Helicella Ferussac, 1819 (1821?) nec Helicelle Lam.,
1812.
b I have been unable to consult the original description of this genus, and therefore
we not know whether it bears any relation to Lissospira or not. Tryon (Manual, x, p. 96) made Helicella (Heliciella index) mutabilis Costa a synonym of Cyclostrema Cutler- Zz tana Clark; (ix, p. 296) he mentioned Heliciella as equal to Megalomphalus Brusina,
meet), in part, but did not quote either the description or type.
AZ
116 K. J. Bush—Marine Gastropods referred to
Ethalia A. Adams, 1853. Type, ZH. Guamense (Quoy and Gaimard). Is. of Guam. and Philippines.
“Shell orbicular, turbinately depressed ; whorls smooth or trans- versely striated, convex, rounded at the periphery; umbilicus partly closed by a callous deposit.” P. Z. S., p. 189, 1853.
This was constituted as a sub-genus of Umbonium, for two spe- cies of variously colored shells, £ Guamense (Q. and G.) and £. striolatum A. Ad. (described), closely resembling that genus but having the umbilicus only partially covered by a callous deposit. Neither was mentioned as a type species. H. and A. Adams, in 1858 (Gen. Rec. Moll., p. 409), added to the description “columellar lip ending anteriorly in an obtuse dilated callus,” but cited the same two species as the only known examples.
P. P. Carpenter, in 1855-7 (Mazatlan Mollusca, p. 250), described five new species of small shells which he referred to Ethalia with some doubt ; for, although they had “the general aspect of Viétré- nelle they agreed with Globulus in having callous bases,” but dif-
-fered in having the callus “ generally not covering the umbilicus.”
A. Adams, in 1861 (Ann. Mag. N. H., viii, p. 305), described three new species as belonging to the genus Hthalia; but the first one #. atomaria is described as a small, semi-transparent, white shell, with the umbilical region entirely covered by a callus; outer lip produced; which shows it to be quite different from either of the two original species and much nearer to Teinostoma politum A. Ad. or Pseudo- rotella semistriata (d’Orb.) Fischer.
These two instances will suftice to show the erroneous application of the genus. Unfortunately, this has been adopted, apparently without question, by more recent authors, so that. a reconsideration of the subject is necessary before the right relations of the numerous species can be satisfactorily determined. 7
Mr. Pilsbry (Manual, xi, p. 457) called attention to the confusion into which the genus had been brought and gave Ethalia Guwamense (Q. and G.) as the type.
List of species belonging to the marine fauna of eastern America — which have been referred to Ethalia:
Ethalia diaphana (d’Orb.) Dall = Vitrinella diaphana (@Orb.) — Bush, Z. anomala (d’Orb.) Dall = Vitrinella anomala (d’Orb.) Tryon, E. semistriata (V’Orb.) Poulsen= Pseudorotella semistriata (dOrb.) Fischer, Z. multistriata Verrill = Vitrinella? multistriata (Verrill) Bush, Z. reclusa Dall, E. suppressa Dall, E. solida Dall, £. (Dill- wynella) modesta Dall. ;
Cyclostrema, Adeorbis, Vitrinella, and related genera. 117
Teinostoma A. Adams, 1853. Type, 7. politum A. Ad. Santa Elena, 8 fathoms.
“Shell orbicular, depressed, subspiral, polished, last whorl rounded at the periphery ; umbilical region covered with a large flat callosity ; the aperture transverse, rounded, greatly produced and elongated, ending anteriorly in a slightly canaliculated point ; inner lip smooth, and callous, not emarginate or truncate anteriorly ; outer lip thin, simple, not marginal or reflected. Animal? oper- culum ?”
“ Although but a single species has been found, I have made a genus of this singular shell, because I was unable fairly to refer it to any known form. It has resemblances to Cyclops, Camitia, and Rotella but agrees with neither.” P. Z. S., p. 183, pl. x, figs. 1-3, 1853,
H. and A. Adams, in 1858 (Gen. Rec. Moll., p. 123) stated: ‘This curious little genus very much resembles at first sight Camitia of Gray, a genus of. Trochide, from which, however, it is readily dis- tinguished. Its true affinity is with Meritula, from which it is known by the absence of the notch at the fore part of the aperture, and by the very peculiar elongation of the mouth.” 7. anomalum C. B. Adams was added as a second known species; but p. 615 they stated that the “anomalum C. B. Adams should be pusillum C. B. Adams” (see page 119), and added three other species to the list. They also considered the relations of the genus to be nearer Umbo- nium as suggested by P. P. Carpenter, and transferred it to the sub- family Umboniine.
Megatyloma (type T. wateleti Desh.) was proposed by Cossmann as a section under Jeinostoma (Cat. Coqu. Foss. Eocene, Paris, 111, p. 50, 1892).
P. P. Carpenter (Mazatlan Mollusca, p. 253) called attention to the fact that as the genus was “described from a single species, some of the characters before given may hereafter prove to be only specific.”
Certainly if “greatly produced and elongated, ending anteriorly in a slightly canaliculated point,” could be taken as a specific charac- ter and be omitted from the above description of the genus, it would be possible to rightly refer many more species to it.
Fischer associated Zéinostoma with Cyclostrema in his family Cyclostrematidx, and Tryon with Vitrinedla.
List of species belonging to the marine fauna of eastern America which have been referred to Tetnostoma :
Teinostoma diaphanum (d’Orb.) Tryon = Vitrinella diaphana
118 K. J. Bush—Marine Gastropods referred to
The Rotella cryptospira Verrill (these Trans., vi, p. 241, 1884) from off Cape Hatteras, N. C., in 142° . fathoms, would seem to be a true Zeinostoma, as “@ suggested by Mr. Dall. Figs. 1, 2.
It is a small, solid, porcellanous, white shell with — the umbilicus entirely covered with a rather thick, — clearly defined, flat pad of callous deposit, distinctly — x10 separated from the columellar lip; edge of the peri-— =n ) treme simple, rounded, not sharp, owing to the thick- 7 ness of the shell; but the aperture is not at all “ pro-
duced and elongated.”
Pseudorotella Fischer, 1857. Type, P. semistriata (d’Orbigny). West Indies.
“ Animal? Operculum ?
“Shell small, transparent, globular or flattened, whorls few, finely striated; umbilicus covered over with a brilliant transparent callos- ity; peritreme not continuous, right border curved, simple, acute.” Jour. de Conch., vi, p. 52, 1857.
Type, P. AIP a (VOrbigny).
“Shell orbicular, depressed, thin, transparent, white, above trans- versely (i. e. concentrically) striate, beneath polished; umbilical cal- lus, shining ; spire very short, obtuse, whorls four, slightly convex; aperture oval. Diameter 1°5™™; height 3™™.” Histoire L’ile de Cuba, Moll., ii, p. 61 ; atlas, pl. xviii, figs. 20-22, 1853.
Fischer (Manuel, p. 234) considered this a subgenus of Teino- stoma.
List of species belonging to the marine fauna of eastern America which have been referred to Pseudorotella :
Pseudorotella semistriata (d’Orb.) Fischer, P. pusilla (Pfeiffer) Fischer = diaphana (VOrb.) = Vitrinella diaphana (dOrb.) Bush, P. carinata (VOrb.) Fischer = Vitrinella carinata (VOrb.) Bush, P. anomala (VOrb.) Fischer = Vitrinella anomala (dOrb.) Tryon, P. striata (VOrb.) Fischer.
Pseudorotella minuscula sp. nov. Figure 3, a, }, c. A single, minute, dead shell from station 2283, off Cape Hatteras, N. C., in 14 fathoms, 1884, has the umbilical region entirely covered
Cyclostrema, Adeorbis, Vitrinella, and related genera. 119
by.a thin, very lustrous glaze or layer of enamel, not in any sense a thickened pad, as in Teinostoma.
Shell thick, solid, porcellanous, slightly tinted with yellow along the suture and on the base ; flattened above and below, with the in- dented umbilical region covered with a thin lustrous glaze or layer of enamel. Surface smooth and very lustrous, marked only by irregu- lar, microscopic, growth lines. Suture incon- spicuous. Whorls about 23, coiled in the same plane, lapping well on to each other, rapidly enlarging, with a very small nuclear whorl and large body-whorl. Aperture very oblique, some-
what ovate; peritreme not continuous, modified into a thin, inconspicuous glaze on the body- whorl, elsewhere with rounded edge, with a slight callous deposit beneath the suture where the outer-lip extends obliquely well for- ward from the body-whorl, with little, if any, curvature and forms a slight sutural notch.
Greatest diameter, about 1°5™"; height, about °5™”,
In form, this species approaches Teinostoma cryptospira (V errill) Dall, but it is a much smaller shell, with the whorls quite differently coiled and with the umbilical callus represented by a thin glaze.
nucleus:
Calceolina A. Adams, 1863. Type, C. pusilla C. B. Adams. Jamaica.
“Shell like a Neritina, oblong, depressed, with small spire. Whorls rapidly increasing; umbilical region callus. Aperture semi-circular, interior not pearly with the inner lip concealed by a large, broad callus covering the umbilicus behind, with a simple, straight, front edge.”
Type, C. pusilla C. B. Adams. Ann. Mag. N. H., xi, p. 267, 1863.
The type was described by C. B. Adams as a Neritina, which it very strongly resembles, so that I question its rightful relation to this group of shells, but H. and A. Adams, Fischer, and Tryon, all used the genus as a section under Zeinostoma.'
1The genus Discopsis de Folin, 1869, (type, D. omalos de Folin, West Indies) (Fonde de la Mer, i, pp. 190, 205, pl. 23, f. 6, 1869) is also placed by Fischer, Tryon, and others as a sub-genus under Zeinostomu. Judging from the description and figures as given by Tryon, it seems to me very distinct, The genus Cochiliolepis Stimpson, 1859 (type, C. parasiticus St., Charleston, S.C.) (Proc. Boston Soc. Nat. Hist., vi, p. 308, with three cuts, 1857-1859) was piaced by Mr. Dall next Disvopsis (Bull. U. S. Nat. Mus., No. 37, p. 162, 1889), but he also suggested its possible rela- tion to the genus Vitrinella (Bull. Mus. Comp. Zodl., xviii, p. 360, 1889).
120 K. J. Bush—Marine Gastropods referred to
Cirsonella Angas, 1877. Type, C. australis Angas. New South Wales.
“Shell minute, globosely turbinate, smooth, narrowly umbilicated; aperture circular ; peritreme continuous, slightly thickened.” Type, C. australis Angas.
“Shell globosely turbinate, narrowly umbilicate, semi-opaque,
smooth, shining, white; whorls 4, convex ; the last large, rounded at the periphery ; aperture circular, peritreme continuous, slightly thick- ened on the columellar margin. Alt. 1 line, breadth, 1 line.” P. Z. S., Pp. S85cpl: wy tr 1G, Vea 7.
Mr. Angas placed this genus provisionally among the Trochidw.
The “slightly thickened” as given for the peritreme in the generic
description is rather misleading, as it is definitely stated in the description of the type to be “on the columellar margin.”
At my request Mr. Smith has very courteously examined the type which is in the British Museum. The following is an extract from his letter under date of May 7th:
“We have the type of Angas’ Cirsonella australis, and it is placed at present in the neighborhood of Ethalia.
“It is a very minute, but solid, porcellanous shell, narrowly um- bilicate, smooth, with a circular mouth, continuous doudle peristome, the inner edge looking as if it might support a shelly operculum as
in Bithynia. The upper figure in P. Z. S., 1877, pl. v, f. 16, is a | good representation of the shell generally ; the figure of the under- —
side is out of drawing and does not give the idea of a round aper- ture and continuous lip. From its texture and general aspect I think it may be left in its present location. Some of >the small Ethalias approximate rather closely to it, but their peristomes are not quite so markedly continuous,” ete.
The open umbilicus would, however, exclude it from Zthalia in its
restricted sense, so that the proper relations of the genus are still ° doubtful. In outline, form of aperture, continuous peritreme, and —
small umbilicus, it strongly resembles Lissospira Bush, (p. 129,) but with the exception of ZL. (Ganesa) rarinota Bush, the species of that genus have shells of delicate texture and not at all solid and porcellanous. Its affinities may possibly prove to be such that the subgenus Ganesa Jeffreys 1883 (p. 114) would become synonymous.
Dillwynella Dall, 1889. Type, D. modesta Dall. Off St. Lucia, 226 fathoms,
“Shell resembling Diloma in form, but minute, depressed, porcel-
lanous, with a thin, horny operculum of comparatively few whorls;
Belp
Cyclostrema, Adeorbis, Vitrinella, and related genera. 121
imperforate, but with a depression bounded by a riblet in the um- bilical rib (region?) outside of the columella; whorls few with a thin fugacious epidermis; outer lip thin ; pillow without teeth, pro- jection, or folds, passing smoothly into the anterior margin.”
Type, D. modesta Dall.
‘This little shell will not fit into any of the groups detined in the text-books, resembling more than any other group the Rotellide, from which it differs in wanting the sutural fasciole, the nacreous layer, and the basal callus, as well as in possessing an epidermis. It is remarkably solid for its size, and of a peculiar opaque white- ness, like Mamma among the Naticide.” Bull. Mus. Comp. Zodl., Xvili, p. 362, pl. xxi, figs. 3, 3a, 1889.
This was proposed as a subgenus of Ethalia, but it is question- able if the species referred to that genus by Mr. Dall could right-
fully belong to it, when taken in its most restricted sense. In
some features it comes near the subgenus Ganesa Jeffreys. See pp. 114, 116.
The following genera have been classed with the Cyclostrematide by Fischer and Tryon. In several features, their types seem much nearer the genus Liotia Gray and related genera, than any of those of the preceding group.
Microtheca A. Adams, 1863, Type, Jf crenellifera A. Ad. Japan.
Globosely turbinate ; peritreme continuous; umbilicus wide, crenu- lated. Operculum unknown. Ann. Mag. N. H., xi, p. 265, 1863. Figured in Thes. Conch., copied by Tryon.
Haplocochlias P. P. Carpenter, 1864. Type, H. cyclophoreus Carpenter. Lower California.
Similar to a Collonia but not margaritaceus, with a continuous thickened peritreme having an exterior varix; young umbilicate, adult lacunate. Animal and operculum unknown. Ann. Mag. N. H., xill, p. 476, 1864.
Leucorhynchia Crosse, 1867. Type, L. Caledonica Crosse. New Caledonia,
~ In the peculiar callous rostrum prolonged past the umbilical region, this genus approximates to Craspedostoma Lindstrom. Journ. de Conch., xv, p. 319, pl. 11, f. 4, 1867.
122 K. J. Bush—Marine Gastropods referred to
VITRINELLIDZ. Cyclostrematide: auth., in part and Adeorbidz Dall, in part. Vitrinella C. B. Adams, 1850. See page 105. Type, Vitrinella helicoidea C. B. Adams.
Vitrinella helicoidea C. B, Adams, Monograph of Vitrinella, p. 9, 1850. non Tryon, Manual, x, pl. 34, figs. 40, 41, 1888.
PLATE XXIII. figs. 9, 9a.
“‘Discoidal: white, opaque and translucent in transverse alternat- ing lines: with a single impressed spiral line near the summit of the whorls, and very fine irregular transverse striw ; apex very obtuse ; spire slightly and convexly elevated ; whorls four, moderately con- vex, rapidly increasing with a lightly impressed suture; last whorl regularly rounded, a little compressed beneath. Aperture not modi- fied by the last whorl ; labium with a rather thick deposit ; umbili-— cus large and deep, with a spiral carina, exhibiting all the whorls. Mean divergence about 150° ; length of spire ‘01 inch ; total length ‘03 inch ; greatest breadth ‘075 inch; least breadth 06 inch,”
A specimen from station 2280, off Cape Hatteras, N. C., in 16 fathoms, agrees so closely with Adams’ description that I have ven- tured to figure it as an example of this, the type species of Vitrinella, —
It is a small, semi-transparent, smooth, shining shell of about three and one-half convex whorls, forming a very low spire with obtuse apex and large body-whorl. Suture inconspicuous with an internal spiral line just below it showing through the shell.” Umbili- cus moderate, deep, showing all the whorls, with straight walls, dis- tinctly angulated and defined on its outer margin by a rather small, rounded thread. Aperture oblique, nearly circular. Peritreme simple with a thin sharp edge, becoming thickened and rounded along the columellar margin and modified on the body-whorl into an inconspicuous, irregular, very thin glaze.
In a basal view the aperture is decidedly angulated below, the lip curving forward from the body-whorl, then backward and abr uptly inward to join the sinuous pillar-lip.
Under the microscope the surface is marked by very delicate sinu- ous raised lines in the direction of the lines of growth.
Greatest width, about 2°3""; height about 1™™.
The higher spire and sh onlarae and carinated umbilicus distin- guish this species from V. helicoidea as figured by Tryon.
Cyclostrema, Adeorbis, Vitrinella, and related genera. 123
Vitrinella Tryoni sp. nov. ? Vitrinella helicoidea Tryon, Manual, x, pl. 34, figs. 40, 41, 1888. PLATE XXII. figs. 11, lla.
A specimen from station 2278, off Cape Hatteras, N. C., in 16 fathoms, very closely resembles the figures in Tryon’s Manual given as V. helicoidea.
It is a smaller, more transparent shell than the preceding, consist- ing of about three flatly convex whorls, which lap well on to each other, forming a very much flattened spire, and large body-whorl. Suture inconspicuous, with an internal spiral line considerably below it showing through the shell and indicating the width of the lapping on of the whorls. Umbilicus moderate, deep, showing all the whorls, with rounded walls, not defined by a carina as in V. helicoidea. Unfortunately the lip is broken, but the general form of the aperture is as in the preceding species, but with the columellar margin less thickened.
Greatest width, about 2"; height, about :8™™.
Vitrinella diaphana (d’Orbigny).
Rotella diaphana d’Orbigny, Histoire L'lle de Cuba, Moll. ii, p. 62; atlas, pl. xviii, figs. 23-25, 1853.
A specimen from station 2113, off Cape Hatteras, N. C., in 15 fathoms, very closely resembles the figures given by d’Orbigny as Rotella diaphana.
It is a minute, smooth, shining, opaque white shell consisting of about three well-rounded whorls, forming a well-raised spire with minute nuclear whorl and large body-whorl. Suture distinct. Umbilicus rather small, deep, with rounded walls, showing only a part of the whorls. Aperture as in V. helicoidea, but with the columellar lip not only thickened but flattened and spreading slightly over the umbilical region. Surface marked only by delicate micro- scopic lines of growth. Greatest width, about 1:2" ; height, about -gnm
“Umbilical callus very minute,” as given by d’Orbigny for this species as well as for carinata, has doubtless led to their being
referred to Psewdorotella, Teinostoma and Ethalia. The thickened &
1 Pseudorotella Fischer, Journ. de Conch., vi, p. 173, 1857 = P. pusilla (Pfieffer) ; Teinostoma Tryon, Manual, x, p. 104, 1888; Hthalia Dall, Bull. M. C. Z, xviii, p. 361, 1889. This species seems to have been referred also to Cyclostrema by Poulsen.
124 K, J. Bush—Marine Gastropods referred to
and flattened columellar margin corresponds to this, but there is not the slightest indication of a callus pad found in species of any of those genera. My specimen is somewhat worn, so that it is not at all transparent.
Vitrinella carinata (d’Orbigny). Rotella carinata @Orbigny, op. cit., p. 62 ; atlas, pl. xviii, figs. 26-28.
I have identified two specimens from station 2278, off Cape Hatteras, N. C., in 16 fathoms, as the carinata of d’Orbigny.
They are minute, semi-transparent, smooth, shining shells, similar in form to the preceding, but having on the body-whorl a distinct peripheral thread or carina which in some positions appears double. The surface under the microscope is marked by delicate lines in the direction of the lines of growth, and on the base by a few inconspic-
uous, raised, revolving lines, more distinct in one specimen than in
the other. One of the specimens also appears to have an indistinctly flattened sutural area. Umbilicus, aperture, and columellar margin as in V. diaphana.,
Greatest width, about 1:°2™™; height, about ‘8™™.
Vitrinella? multistriata (Verrill).
Ethalia multistriata Verrill, these Trans., vi, p. 242, 1884; Expl. Albatross, Report U.S. Com. Fish and Fisheries for 1883, p. 568, 1885. Dall, Bull. Mus. Comp. Zodl., Xvill, p. 361, 1889; Bull. U.S. Nat. Mus., No. 37, p. 160, 1889.
PLATE XXII. tig. 7. PLATE XXIII. figs. 4 and 14.
The specimens (No. 35733) from station 2109, off Cape Hatteras, N. C., in 142 fathoms, which were described by Professor Verrill as belonging to Zthalia, seem much nearer Vitrinella.
The young specimens are semi-transparent and shining, like V. helicoidea, V. Tryoni, etc., but the more mature ones are somewhat weather-worn and appear quite opaque but have considerable luster. The umbilicus is of moderate size, deep, showing part of the whorls, with the walls somewhat flattened. The aperture is similar in form to that of V. helicoidea, but in the adult the columellar margin is considerably thickened and flattened so that its outer margin is decidedly angulated below, where it joins the outer lip. There is, however, no callus pad covering the umbilical region.
Rotella anomala VOrbigny (Hist. L’Ile de Cuba, ii, p. 64 ; atlas,
*
a eM lah EE A he ee
Cyclostrema, Adeorbis, Vitrinella, and related genera. 12
or
pl. xviii, figs. 32-34, 1853), should undoubtedly be referred to Vitrinella rather than to Hthalia as given by Mr. Dall (Bull. M. C. Z., xviii, p. 361, 1889).
Section Circulus Jeffreys, 1865. See page 110. Adeorbis Wood, 1842, in part. Type, Circulus striatus (Philippi) Pilsbry.
Adeorbis striatus Wood, Ann. Mag. Nat. Hist., ix, p. 530, pl. v, figs. 5-6, 1842; Mon. Crag. Moll., p. 137, pl. xv, fig. 7, 1848 = Valvata? striata Philippi (teste Wood).
Circulus Duminyi Jeffreys, B. C., iii, p. 315, 1865; y, p. 203, pl. lxii, f 5, 1869 = Solarium Philippii Cantraine (teste Jeffreys).
Circulus striatus Pilsbry, Tryon’s Manual, xi, p. 274, pl. 66, figs. 12, 13, 1889. (A very complete synonymy is given.)
PLATE XXIII. fig. 11.
Mr. Dall has very kindly loaned me specimens of Duminyi from both Naples and the coast of France, in the U. S. National Museum.
They resemble the smaller species, the /ivatus of our coast, in hay- ing the upper part of the whorls covered with raised spiral threads separated by grooves, but the grooves are less deeply cut, making the alternating threads less elevated. The whorls are less swollen or convex, the suture shallower, the nuclear whorl, although of about the same size, is less elevated, causing the shell to appear lower and flatter, with a flatter base, a feature which increases with age.
The sculpture on the series of specimens from the coast of France is even less conspicuous than that on the single larger specimen from
Naples. Circulus liratus (Verrill).
Omalaxis (?) lirata Verrill, these Trans., v, p. 529, 1882.
Skenea lirata Verrill, op. cit., vi, p. 452, 1885. Bush, op. cit., vi, p. 464, 1885.
Adeorbis supranitidus, var. Orbignyi Dall, Bull. Mus. Comp. Zodl., xviii, p. 278, 1889; Bull. U. S. Nat. Mus., No. 37, p. 150, 1889, in part.
Skenea lirata Bush, Bull. Mus. Comp. Zodl., xxiii, p. 240, pl. i, figs. 11, 12, 1893.
Piate XXIII. figs. 7, 12-120.
Found in considerable numbers off Cape Hatteras, N. C., in 8-43 fathoms, 1883-1884.
This species is considered by Mr. Dall to be the same as Fischer’s A. Orbignyi from the West Indies. That, however, is a very minute species (diam. 14™™; height 1™™), ornamented, “ with 12-15 elevate, acute, equidistant, concentric cost,” while the larger species, liratus (diam. 24™™; height, about 1™"), has but 9 or 10 cinguli or lire.
126 K. J. Bush—Marine Gastropods referred to
Circulus Smithi Bush. Cyclostrema tricarinatus Smith, P. Z. S., London, p. 737, pl. 75, fig. 26, 1871.
In studying the various figures published in Tryon’s Manual I was impressed with the strong resemblance of the African species, Cy- clostrema tricarinatus Smith, to the Crag species of Adeorbis Wood, esp. tricarinatus. I referred the matter to Mr. Smith, who writes me that upon comparing his species with those of Wood, he finds that there is unquestionably no generic distinction. He also states that his figures are accurate representations of the only specimen which he had of his species. As the specific name tricarinatus is preoccupied by Wood, I propose the name Smithi for this African — shell. It differs from any of those on our coast which belong to Circulus, in having the entire surface covered by conspicuous revoly- ing threads and grooves, in addition to three prominent carine.
Circulus Dalli sp. noy. PLaTE XXIII. figs. 3, 3a and 6.
A single dead specimen, found among foraminifera, at station 2655, N. lat. 27° 22’, W. long. 78° 07’ 30’, in 338 fathoms, 1886.
This deep-water species is of more delicate texture and more trans- parent than the more northern shallow-water species of similar form. It is ornamented on the body-whorl with two rather inconspicuous carine, one defining the base and the other on the periphery; above this the surface is cut by about seven delicate, unequal, microscopic shallow grooves or striz, the two uppermost being the most distinct; above these the surface is smooth and appears somewhat flattened ; there are also a few less distinct strize below the periphery and in the umbilical region.
Greatest width, about 3™™; height, about 1:4™™.
A smaller dead specimen (No. 44983) from station 2307, off Cape Hatteras, N. C., in 43 fathoms, agrees well with this species. It has however, in addition to the typical sculpture, a few smaller striz just below the suture, and the grooves and alternating raised lines appear more distinct, the specimen being very much worn and twice injured and repaired by the animal.
This larger species, in its inconspicuous sculpture, seems to be a connecting link between the distinctly grooved ones and the cari- nated ones, so that we have a series of gradations in sculpture from the smooth variety of supranitidus through supranitidus (typical), trilix, Dalli, Smithi, striatus, up to the strongly grooved liratus.
Cyclostrema, Adeorbis, Vitrinella, and related genera. 127
Circulus trilix Bush.
Skenea triliz Bush, Expl. Albatross, Report U. 8. Com. Fish and Fisheries for 1883, __ —p. 584, 1885.
Non Homalogyra densicosta Tryon, Manual, ix, p. 399, pl. 61, figs. 10-11, 1887,
j Adeorbis supranitidus Dall, Bull. Mus. Comp. Zo6l, xviii, p. 278, 1889; Bull. U.S. _ Nat. Mus., No. 37, p. 150, pl. xli, figs. 7, 7a, 1889, in part.
Puate XXII. figs. 6, 10, 10a and 12, a-g. Puare XXIII. figs. 10 and 15.
The most common species of this genus off Cape Hatteras, N. C., in 7-17 fathoms, 1883-1884.
The operculum of ¢fri/ix is thin, horny, circular, of about five whorls with central nucleus. The animal matter, the shell having - been removed. with nitric acid, is too much dried to dissect but shows stout tentacles, prominent eyes situated at their bases, and a rather broad, bilobed snout (fig. 6). The radula, although ex- tremely minute, measuring about °3"™ in length and ‘1™™ in width, _ reveals interesting characters and shows that this species unques- _ tionably belongs to the Rhiphidoglossa. The radula (fig. 12) con- sists of between 50 and 60 rows of teeth ; in each row there is a - comparatively wide central tooth (fig. 7) having a base with convex sides and bearing a long, finely serrated, strongly pointed hook; on either side a narrow, long, lateral tooth (fig. e¢) with a serrated hook ; and beyond twenty or more delicately pointed, sickle-shaped, mar- ginal ones (figs. f, g). These were revealed only when exceed- ingly high power objective and ocular, of a magnifying power of about 1000 diameters, were used, and even then only the higher lines could be seen; when using a lower power of about 500 diame- ters, only ten of the marginal hooks could be counted. See p. 128.
Circulus supranitidus (Wood) Jeffreys.
__ Adeorbis supranitidus Wood, Ann. Mag. Nat. Hist., ix, p. 530, 1842; Crag Mol- lusea, p. 137, pl. xv, figs. 5, a—b, London, 1848. — Circulus Duminyi Requien, var. supranitidus Jeffreys, B. C., iii, p. 315, 1860. Adeorbis supranitidus Jeffreys, P. Z. S., London, p. 42, 1889. _ Non Omalaxis supranitida G. O. Sars, Moll. Reg. Arct. Norv., p. 214, pl. 22, figs. 20 a-b, 1878=(0. (?) Sarsi sp. nov.) Non Adeorbis supranitidus Dall, Bull. Mus. Comp. ZoOl., xviii, p. 278, 1889; Bull. U. ‘S. Nat. Mus., No. 37, p. 150, pl. xli, figs. 7, 7a, 1889.
_
Pirate XXIIL, figs. 1 and 2.
‘ Through the courtesy of Mr. Dall, I have been able to study two authentic specimens of swpranitidus from the Crag, in the U. 8. National Museum.
128 K. J. Bush—Marine Gastropods referred to
They are about half the size of a full-grown ¢rilix and differ from specimens of that species of the same size, in lacking the distinct tricarination of the whorls, in having the whorls more convex and more regularly coiled, so that the body-whorl is not so abruptly enlarged, in having the spire relatively larger, and in the striation of the umbilicus. Both specimens of supranitidus have three conspic- uous revolving threads separated by deep grooves in the umbilicus; one has a very prominent basal carina, the other has it but partially developed; in this specimen the whorls are perfectly smooth, while | in the former the two faint upper carinze commence just back of the aperture; neither show any trace of the microscopic striations found | in trilix. In the many specimens of the latter which I have, there seems to be no variation in the size of the small nucleus, the rela- tively much smaller spire and abruptly erlarged body-whorl (a pecu- liarity which becomes more marked as the shell increases) and in the strong tricarination of the whorls, even in very small specimens. | They differ only in the number and prominence of the threads in © the umbilical region, and in the distinctness of the microscopic stria- tions on the upper part of the whorls. The constancy of these char-— acters ought to prove that the two species are quite distinct.
The four species, besides subcarinatus (Montagu), included in Adeorbis by Wood present two quite distinct forms ; pulchralis, which bears a strong resemblance to a Margarita, and supranitidus, — tricarinatus and striatus, which are quite different. Of these, swpra- nitidus has caused the most confusion in synonymy, as no two writers seem to agree as to what it really is. As mentioned above, Mr. Jeffreys identified it as a fossil variety of Trochus Duminyi Requien, but in vol. vy, B. C., 1869, he mentioned that that species should fall into the synonymy of Solariwm Philippii Cantraine.
In 1878, Prof. G. O. Sars described and figured, as supranitidus Wood, a very minute Norwegian shell,’ referring it to the genus Omalaxis. He stated that the spire is plane, not elevated; suture profoundly impressed, etc., etc. At the same time he defined the genus Adeorbis Wood, and described and figured a species very different in form from suwbearinatus (Montagu), but closely resem- bling pulchralis Wood, with which Mr. Jeffreys compares it (Proc. Zool. Soc., London, p. 41, Jan., 1885). Prof. Sars places both genera in the family Solariide at the end of the Gymnoglossa.
‘For this very distinct species, which does not seem to be very closely related generically to Circulus, | propose the name Sarsi, but very much doubt its near rela- tion even to Omalazis. :
Cyclostrema, Adeorbis, Vitrinella, and related genera. 129
Mr. Jeffreys in 1875 changed his mind in regard to supranitidus' Wood, as the “ Lightning and Porcupine Expedition,” obtained speci- mens which he mentioned as “agreeing with the Crag specimens in every respect (especially in being tricarinated) except in being spirally and rather strongly striated,” and added A. tricarinatus Wood as another variety. He further mentioned that “the opercu- lum is not known, and it is therefore questionable whether the pre- sent species belongs to Adeorbis or Homalamis.” He however referred it to Adeorbis, but did not mention his genus Cireulus.
Lissospira gen. noy. Cyclostrema auth, in part.
This genus is proposed for a group of deep-water species which have been erroneously referred to Cyclostrema. They are small, thin, of rather delicate texture, opaque white, slightly lustrous, of few convex whorls forming an elevated spire, with relatively large, prominent nuclear whorl and large body-whorl. Suture deep, Umbilicus small, deep, not showing any whorls. Aperture somewhat oblique, circular with a slight sutural angle, not modified by the body-whorl to which the simple, continuous peritreme is but slightly attached, often having an indistinct thread just within the inner lip, fading out above and below, so that it extends but about half way round the aperture ; it is much nearer the edge along the columellar margin than at the ends and is evidently to prevent the thin oper- culum being drawn in too far. The operculum (fig. 4) is cireular, thin, of a delicate horn-color, with central nucleus, of about seven whorls, defined by a distinct spiral thread ; often showing delicate, microscopic transverse growth lines. The radula consists of numer- ous rows of delicate teeth ; each row having one broad central, or median tooth, with a broad, blunt, delicately serrate, curved tip and on either side four more slender lateral teeth also with blunt, curved, delicately serrate tips, beyond which is a seriesof numerous, between 30 and 50, long, very slender, somewhat sickle-shaped hooks some- times with delicately serrate tips. (G. O. Sars, Moll. Reg. Arct. Norv., tab. iii. f. 6 6. and L. diaphana.)
_. 1! The locality (New England, Verrill) given by Mr. Jeffreys, I have been unable to 2 Wr erity, as I find no such species on record or mention of it in any of Professor Verrill’s _ papers. TRANS. Conn. ACAD., VoL. X. JULY, 1897. 9
130 K. J. Bush—Marine Gastropods referred to
Type, Lissospira proxima (Tryon).
Cyclostrema rugulosum Verrill, these Trans., v, p. 533, 1882, non G. O. Sars, 1878. Cyclostrema affine Verrill, op. cit. vi, p. 199, pl. xxxii, fig. 15, 1884, non Jeffreys, 1883. ; Cyclostrema sp. Verrill, Expl. Albatross, Report U. S. Com. Fish and Fisheries for 1883, p. 569, 1885. " Cyclostrema proxima Tryon, Manual of Conchology, x, p. 98, pl. 33, fig 4, 1888. ; Cyclostrema trochoides Dall, Bull. Mus. Comp. Zodl., xviii, p. 393, 1889; Bull. U. S. Nat. Mus., No. 37, p. 166, 1889.
Figure 4. PuLate XXII. fig. 3.
A comparatively few specimens were obtained by the U.S. F. C. — at about twelve stations between N. lat. 41° 53’, W. long. 65° 35’ and N. lat. 35° 49’ 30”, W. long. 74° 34' 45”, in 365 to 858 fathoms, A young specimen (No. 78460) from near stations 2692-6, N. lat. 46°+, W. long. 44°+, in 73-105 fathoms, was also identified as this. species. |
The operculum (fig. 4) is thin, of a delicate horn- color, slightly concave in front. The radula appar-— ently had the same number of teeth in each row as in that of Z. diaphana and L. basistriata,’ but was unfortunately lost before the form of the median tooth could be satisfactorily determined.
Mr. Jeffreys seems to have had a very confused idea as to the exact form he intended to stand as the type of his — species trochoides. A specimen which he identified for Dr, Friele as trochoide (published by the latter in 1875, not described) proved — to be the Zevigatum G. O. Sars, 1878 (teste Friele and Sars). Speci- mens identified as trochoide for Professor Sars, described and figured by him in 1878, must stand for the species, especially as Mr. Jeffreys himself never published a description or figure of it; Sars’ species, -
) There, seems also to be some question in regard to the true Cyclostrema basistriatum Jeffreys, 1877, non Brugnone. In Ann. Mag. N. H., p. 234, 1877, under this species Mr. Jeffreys described three differently sculptured forms: a, ‘spiral strize on base encircling the umbilicus”; }, ‘‘a single ridge-like stria at the top so as to give an angulated appearance to the summit”; c, “umbilicus surrounded by a strong keel- like stria”’; while in P. Z.S., p. 90, 1883, he added the fourth, d, “striated through- out,” and “ C. profundum Friele is this form” (p. 141.) In 1886 Dr. Friele very decidedly stated that his species was not the basistriatum of Jeffreys. G. O. Sars described and figured the form having striae on the base only and this ought to stand for the true or typical form and is undoubtedly a Lissospira. The basistriatum of Brugnone was said by Jeffreys to have been changed to curvistriatum Brugnone.*
Cyclostrema, Adeorbis, Vitrinella, and related genera. 131
however, has been found to be a variety of Friele’s Petterseni (1877) (teste Friele and Sars). In P. Z.S., p. 91, 1883, Mr. Jeffreys men- tioned “ The umbilicus is sometimes encircled by one or more strong spiral strie,” which leads me to think that he had still other forms erroneously referred to the species, more especially as Mr. Dall (op. cit., p. 393) states that “A careful comparison of types leaves no doubt as to the identity of these two forms” (affine V. and trochoides J.); but the figures given by both Friele and Sars are very unlike affine V. Petterseni is described as solid, perfectly smooth, white, with no umbilicus proper, but a more or less distinct umbilical fissure, and the figures show the form of the aperture and peritreme to be very simi- lar to that figured by Jeffreys for the type of Tharsis= Tharsiella, as do also those given by Sars. As the specific name affine was used in 1883 by Jeffreys for another species of Cyclostrema which now would be referred to Lis- sospira, the name proxima, given by Tryon in 1888, is adopted for Professor Verrill’s species.
Lissospira diaphana (Verrill).
Cyclostrema diaphanum Verrill, these Trans., vi, p. 199, pl. xxxii, fig. 16, 1884; Expl. Albatross, Report U. S. Com. Fish and Fisheries for 1883, p. 569, 1885. Tryon, Manual of Conchology, x, p. 91, pl. 31, fig. 47, 1888. Dall, Bull. Mus. Comp. ZoiL., XVili, p. 393, 1889; Bull. U.S. Nat. Mus., No. 37, p. 166, 1889; Proc. U. S. Nat. Mus., xii, p. 355, 1889.
Puate XXII. fig. 2.
Two specimens (No. 38409), station 2084, off Martha’s Vineyard, in 1290 fathoms, 1883, south to Brazil, in 281-1019 fathoms (Dall). The locality, station 2004, as given by Professor Verrill was a typo- graphical error.
The operculum is similar to that of the preceding species and basistriutam as figured by Sars. The radula, however, shows inter- esting, though slight, differences from that of the latter. The median tooth being not broadly elliptical, but narrower above than below, the sides at first straight but curving strongly outward below, with a broad, blunt, delicately serrate, curved tip; on either side are four narrow, curved, lateral teeth, also with blunt delicately
“serrate hooked tips ; beyond these are numerous, between 30 and 40, very fine, hair-like, delicately hooked marginal teeth ; no serrations were found on these.
132 KE. J. Bush—Marine Gastropods referred to
Lissospira striata sp. nov. Figures 5, a.
Fig. 5. One live specimen, station 2213, off Mar-
= tha’s Vineyard, in 384 fathoms, 1884.
Shell consisting of about three convex. whorls forming a comparatively low, some- — _what depressed spire. The entire surface be- low the relatively large, little raised, smooth nuclear whorl, is covered with raised, round- — ed, well-separated, revolving microscopic threads, most distinct on — the base. Umbilicus small, scarcely more than a chink. Aperture and operculum typical.
Greatest width, about 2™™ ; height, about 1°5™™.
This species, in general form and sculpture, resembles Cyclo-_ strema (= Lissospira) Willet Friele (Den. Nor. Nord.-Expd. 1876-78, xvi, Moll. ii, p. 34, pl. xi, fig 19, 1886), but that is a much smaller species with larger umbilicus.
Lissospira cingulata (Verrill).
Cyclostrema cingulatum Verrill, these Trans., vi, p. 198, pl. xxxii, fig. 14, 1884 (non Philippi nec Dunker); Expl. Albatross, Report U. S. Com. Fish and Fisheries for 1883, p. 569, 1885.
Cyclostrema Verrilli Tryon, Manual of Conchology, x, p. 90, pl. 31, fig 46, 1888.
Cyclostrema cingulatum Dall, Bull. U.S. Nat. Mus., No. 37, p. 166, 1889.
One specimen (No. 38100), station 2048, off Martha’s Vineyard, in 547 fathoms, 1883.
As this species was erroneously referred to Cyclostrema, I do not adopt Tryon’s specific name Verrilli for it.
Lissospira (?) convexa sp. noy.
A much mutilated, live specimen, station 2233 off Delaware Bay, in 630 fathoms, 1884,
Shell very fragile, consisting of four regularly increasing, very convex whorls, forming a high spire with abnormally relatively large, prominent, somewhat twisted, nuclear whorl. Suture very deep, umbilicus round, of good size, deep, with rounded walls. Aperture somewhat circular but modified by the body-whorl, to which the simple, continuous peritreme is attached for a consider- — able distance; columellar margin very straight, angulated at its
Cyclostrema, Adeorbis, Vitrinella, and related genera. 133
junction with the strongly curved outer-lip and reflected along the umbilical region, but not thickened. Surface appears smooth but
under the microscope it is covered, except the nucleus, with delicate, much separated, spiral threads which extend up into the umbilicus. The animal is drawn so far into the shell that the operculum, if present, cannot be seen.
Greatest width, about 1.5" ; height, the same.
Although at first sight this species strongly resembles a Lissospira, in the large size of its nucleus, regularly increasing whorls, and form of its aperture it approximates to Rissoa (Setia) triangularis Watson (Report Voy. Challenger, Zod]. Scaphopoda and Gasteropoda, xv, p. 611, pl. xlvi, fig. 2, 1885), but is much smaller with more swollen whorls. 2
Subgenus Ganesa Jeffreys, 1865. See page 114. Lissospira (Ganesa) Dalli (Verrill).
Cyclostrema trochoides Verrill, Notice of Recent Add. to Mar. Invert., Part 11, Proce. U.S. Nat. Mus., iii, p. 378, 1880, non G. O. Sars, 1878.
Cyclostrema Dalli Verrill, these Trans., v, p. 532, pl. lvii, fig. 39, 1882; Expl. Albatross, Report U.S. Com. Fish and Fisheries for 1883, p. 569, pl. xxvii, fig. 99, 1885. Tryon, Manual of Conchology, x, p. 97, pl. 33, fig 100, 1888.
Cyclostrema fulgidus Dall, Bull Mus. Comp. Zodl., xviii, p. 393, 1889, non (Trochus fulgidus) Jeffreys, 1883.
Cyclostrema fulgidwm Dall, Bull. U. 8. Nat. Mus., No. 37, p. 166, pl. Ixiii, fig. 99, 1889.
About a dozen specimens have been recorded as found at five stations between N. lat. 41° 53’, W. long. 65° 35' and N. lat. 38° 01’ 15’, W. long. 73° 44’, in 390-1188 fathoms, 1880-1886,
Specimens from very deep-water are not of so firm a texture as the typical examples, the entire surface appears chalky without much lustre, and the lines on the base are less distinct. There is little resemblance between G. Dalli and the figure of Trochus fulgidus as given by Mr. Jeffreys, who described that species as having the peritreme “not continuous or complete ;” while Dal/i has the peri- treme continuous, attached to the body-whorl for a considerable dis- tance so that the form of the aperture is somewhat modified, and the columellar margin is flattened so that it spreads slightly over the
~ umbilical region. The umbilical chink or fissure appears channeled and is defined by a raised thread, as in other species of Ganesa.
134 K. J. Bush—Marine Gastropods referred to
Lissospira (Ganesa) ornata (Verrill).
Cyclostrema Dalli, var. ornatum Verrill, these Trans., vi, p. 255, pl. xxxii, fig. 17, 1884; Expl. Albatross, Report U. 8. Com. Fish and Fisheries for 1883, p. 569, 1885.
Cyclostrema ornata Dall, Bull. Mus. Comp. Zodl., xviii, p. 393, 1889; Bull. U.S. Nat. Mus., No. 37, p. 166, 1889.
One live specimen (No. 35610), station 2115, off Cape Hatteras, N. C., in 843 fathoms, 1883. ‘ As the principal distinguishing feature of the species of this genus lies in the arrangement of the microscopic sculpture, I follow Mr. —
Dall in giving specific distinction to the varietal name. The umbili- —
cal fissure, margined by a raised thread, would place the species — with Ganesa rather than Tharsis (see p. 1138), as suggested by Professor Verrill.
Lissospira (Ganesa) abyssicola sp. nov.
Ganeza sp. Verrill, these Trans., vi, p. 202, 1884.
Ganesa sp. Bush, Bull. Mus. Comp. Zo6l., xxiii, p. 219, 1893.
As described by Professor Verrill, this species in general appear- ance resembles a true Lssospira, but the umbilical chink is channeled and defined by a raised thread and the aperture is modified by the body-whorl to which the peritreme is more attached than in typical species.
It is similar in form to G, Dailli but larger, without sculpture and with the columellar lip less flattened. ss One dead specimen, station 307 Ag., east of George’s Bank, in 980 fathoms, 1880, ‘‘ Blake Expedition.”
.
Lissospira (Ganesa ?) rarinota sp. nov.
Shell smooth, white, semi-opaque, of firm texture, with brilliant luster so that it appears somewhat porcellanous. Whorls three, - convex, with a single, delicate, raised, microscopic, spiral line just below the distinct suture, defining a narrow, inconspicuous, slightly concave or channeled, sutural area. Nucleus relatively small, little raised ; body-whorl large ; spire low. The form of the aperture and peritreme and the channeled umbilical chink, bordered by an incon- spicuous, rounded thread, place this species with Ganesa, although the texture of the shell is firmer and more porcellanous than any of — the other species referred to it. Just within the aperture and extending completely around it is a delicate, opaque white line, — which in some places appears raised.
This may prove to be a Cirsonella, p. 120.
Cyclostrema, Adeorbis, Vitrinella, and related genera. 135
Greatest diameter about 2:2"; height, about 1°6™™. One dead specimen, among foraminifera, at station 2150, N. lat. 13° 34’ 45”, W. long. 81° 21’ 10’, in 382 fathoms, 1884.
Section Granigyra Dail. See p. 115.
Type, Granigyra limata (Dall),
Cyclostrema (Granigyra) limatum Dall, Bull. Mus. Comp. Zodl., xviii, p. 395, 1889; Bull. U. 8. Nat. Mus., No. 37, p. 166, 1889.
A single dead specimen, found among foraminifera, from station 2150, N. lat. 13° 34' 45”, W. long. 81° 21’ 10’, in 382 fathoms, 1884, agrees, except in size, with Mr. Dall’s description of his type from off Cuba, in 310 fathoms. My specimen measures about 2™™ in greatest diameter and about 2°5™™ in height, and in this respect comes nearer
the G. pruinosa of Jeffreys, the measurements of which are given as
“TL, 0.175, B. 0.15.”
Granigyra spinulosa sp. nov.
A single dead specimen, found among foraminifera, from station 2655, N. lat. 27° 22’, W. long 78° 07’ 30”, in 338 fathoms, 1886.
This is a larger shell than the preceding but is of the same size as the type-specimen of that species, of a delicate yellow color with the granulations of the surface much coarser and more prominent.
It has about three well-rounded, rather loosely coiled whorls, forming an elevated spire with relatively large, prominent nuclear whorl and very large body-whorl. Suture well-marked, rather deep. Umbilicus of good size, deep, with rounded walls. Aperture as in typical Lissospira, with the peritreme but slightly attached at the suture. Granulation of the surface very conspicuous, the granules, when seen in profile, appearing like little spines.
Greatest width, 2°5™™, height the same.
Leptogyra gen. nov.
This genus is constituted for a group of three species of minute, semi-transparent, dull, dirty white or faintly brown shells covered with a thin, rather tough, delicate straw-colored epidermis, consist- ing of a few convex whorls forming an elevated spire with relatively
5 large, smooth, slightly twisted nuclear whorl and large body-whorl.
a a »
a
tt a
Suture deep, somewhat channeled. Umbilicus relatively large, round, deep, showing some of the whorls, with rounded walls.
136 K. J. Bush— Marine Gastropods referred to
Aperture very oblique, somewhat ovate. Peritreme simple, contin- uous, modified on the body-whorl into a thin glaze, sometimes in the adult having a free edge; strongly sinuate along the umbilical region and anteriorly, slightly angulated below, at the junction of the two lips; above, arching well upward, forward, then backward — from the body-whorl forming a distinct sutural notch.
Interior of the aperture smooth and very lustrous, with the con- spicuous, exterior, transverse lines showing through by transparency. There is no internal opaque line ; in all the specimens the operculum _ is drawn well into the shell. :
The operculum is very thin, circular, of a delicate horn-color, with — central nucleus, of about seven whorls defined by a fine spiral line.
The animal of the type species is too much dried to dissect, the shell being removed by hydrochloric acid, but the radula shows exceedingly interesting features. The entire length is but about °35™™ and the width about ‘08™™. It consists of numerous rows of —
very delicately colored teeth. In each row there is a long, rather
broad, strongly hooked median tooth; on either side, three or four lateral ones of about equal size; all of these, when seen in profile,
appear distinctly triserrate and the outer ones somewhat larger than
the others ; beyond, there is a series of between 30 and 40, long, very
slender, marginal hooks.
Type, Leptogyra Verrilli sp. nov.
Pratge XXIII. figs. 13, 13a.
.
Shell consisting of three whorls which form a moderately elevated
spire with obtuse apex due to the scarcely raised nuclear whorl. —
The surface, below the nucleus, is covered with microscopic, — impressed, spiral lines, and more conspicuous, sinuous, raised, irregu- lar, transverse lines or wrinkles which are most distinct near the suture and upon the upper part of the whorls, in addition to the very fine lines of growth; on the middle of the body-whorl and on the base, the spiral striae become broader so that the alternating
spaces appear like raised threads. Operculum and radula described _
under the genus.
Greatest diameter of one of the largest specimens, about 1°5"™; height, about -9™™,
Eight live specimens, station 2174, off Delaware Bay, in 1594 fathoms, 1884.
Cyclostrema, Adeorbis, Vitrinella, and related genera. 137
Leptogyra inconspicua sp. nov.
Shell consisting of about 25 whorls, forming a little elevated spire with very obtuse apex (the nuclear whorl being coiled on the same plane as the succeeding one). Surface nearly smooth, slightly lus- trous, having, besides the delicate microscopic growth lines, irregu- larly dispersed, ill-defined, sinuous wrinkles extending from the suture a little way below the summit of the whorls.
Greatest diameter, about 1:3"; height, about :8™™,
Two live specimens, station 2174.
Leptogyra eritmeta sp. nov.
Shell consisting of three whorls which are more loosely coiled than in either of the preceding species, causing the spire to appear more elevated and the suture not so deep. Nuclear whorl very prominent. Surface below the nucleus, crossed by prominent, regular, evenly separated lines or threads which extend from suture to suture and over the base well up into the umbilicus, but are not so conspicuous here as on the upper part of the whorls. Under a high power, the summit of these appears uneven or very inconspicuously serrate.
Greatest diameter, about 1°5™"; height, about 1™™.
One live specimen, station 2174. It is an interesting fact that all three species were found at the same station.
Molleriopsis gen. nov.
Shell small, relatively thin, white under a golden ‘brown or olive brown epidermis. Whorls few, convex, forming an elevated spire and large body-whorl. Suture distinct. Umbilicus of moderate size, round, deep, showing some of the whorls. Aperture circular, slightly oblique; peritreme continuous with a thin, sharp edge, appearing thickened within, attached to the body-whorl only for a short distance. Animal and operculum not known.
This genus is proposed for a very deep-water shell which closely resembles WM. costulata (Miller) Jeffreys, the type of the genus Molleria, in several features, as the form of the aperture and peri- treme and especially in the conspicuous carination of the base bordering the umbilicus ; but differs from that rather solid, strongly
sculptured species in being much larger, of more delicate texture,
with a nearly smooth surface. The Cyclostrema sulcatum Watson
_ (non A. Adams, 1850) = Watsoni Tryon, 1888 (Report Voy. Chal-
_ lenger, Zod]. Scaphopoda and Gasteropoda, xv, p. 121, pl. viii, fig. 11,
138 K. J. Bush—Marine Gastropods referred to
1885), a very small species from off Brazil, in 675 fathoms, and Adeorbis sincera Dall (Proc. U. 8. Nat. Mus. xii, p. 338, pl. xii, f. 2, 1889), from off Florida and Brazil, in 294 and 391 fathoms, have several features in common with the above deeper water form and can doubtless be referred to the same genus.
Type, Molleriopsis abyssicola sp. nov. Figures 6, 7.
One dead, imperfect specimen (No. 52496), station 2572, N. lat. 40° 29’, W. long. 66° 04’, in 1769 fathoms, 1885. 4
Shell small, opaque white, under a golden brown — epidermis, of few convex whorls (the tip is broken away) forming a well-elevated spire and large body- whorl. Surface lustrous where rubbed, ornamented — with a single conspicuous carina which defines a _ broad, flattened sutural area and makes the whorls _ slightly angulated. This is roughened by the cross-— ing of the growth lines, which are elsewhere incon- spicuous. On the base surrounding the umbilicus there are also four more prominent carine, about — equal in size and evenly separated by wide, slightly — concave interspaces ; the first one, situated about the — middle of the base, and the last one well up in the umbilicus; on all of these the lines of growth are so conspicuous as — to give them a distinctly beaded appearance ; at their termination they form distinct points on the somewhat expanded thin edge of — the peritreme. The aperture within is dull, opaque white, with a narrow, much thinner, semi-transparent, somewhat expanded, sharp- edged border. This gives the appearance of thickening, but there is- no raised opaque white line, said by some author to be the dis- tinguishing character of MWélleria,
Greatest diameter, about 3:2"; height of body-whorl, about gO as
Choristella gen. nov. Figure 8.
This genus is proposed for two species of small shells of few con- vex whorls forming a flattened, little elevated spire with minute, scarcely raised, nuclear whorl and large body-whorl. Suture very deep, somewhat channeled. Umbilicus small, round, deep, showing
Cyclosirema, Adeorbis, Vitrinella, and related genera. 139
some of the whorls with rounded walls. Aperture oblique, nearly circular. Peritreme simple, continuous, slightly attached to the body-whorl, reflected over the umbilicus.
Operculum (fig. 8) of the type species is thin, round- ovate, delicate horn-color, of few abruptly enlarging whorls indistinctly defined by a spiral thread and showing sinuous transverse lines of growth, nucleus slightly excentric. il ee
The animal has a broad emarginate head with one \ =< pair of long slender tentacles ; with a rather broad, short, tapered, ciliated verge just beneath the base of the right one. Eyesnone. Gill attached to the left side lying across the top of the body just within the mantle edge. Jaw plates thin, delicate horn-color with a broad band of very dark brown along the strongly serrate, cutting edges. Inner surface strongly reticulated, as in species of Velutina. The form of these plates is quite irregular: the cutting edge is oblique, forming an angle of about 135° with the inner or middle, straight edge; the distal outline is very strongly sinuously curved, forming a wide, shallow upper portion and a much narrower basal portion.
The radula (pl. xxiii, fig. 16) consists of numerous rows of delicate colored, rather stout, non-serrate teeth, each row having a series of thirteen :—a very small central or median tooth with rather long, strongly curved tip, placed a little above and alternating somewhat with the rest of theseries; on either side, one broad strongly hooked
lateral, and a much broader second lateral one with correspondingly broad, more pointed hook; beyond, three, about equal, much nar- ‘rower, somewhat sickle-shaped, marginal ones with a small triangular, scarcely perceptible, platelike one on the outer edge.
The form of the shell and operculum strongly resemble a Choristes elegans var. tenera Verrill of medium size, but the radula shows marked and interesting differences. In that species, or rather variety, there are but eleven teeth in each series, the second or outer Jateral tooth having a double or bilobed tip (these Trans., v, p. 541, pl. lvi, figs. 27, 27a, 1882; vi, p. 256, pl. xxix, figs. 9-96, 1884.
Type, Choristella leptalea sp. nov. Figure 9. Puate XXIII. figs. 16, a. P Shell without sculpture, dull opaque white, of very delicate tex- ture, consisting of about three and a half whorls. Operculum, animal, and odontophore described under the genus. Greatest width, about 3°5™™; height, about 2°5™™.
140 K. J. Bush—Marine Gastropods referred to
One imperfect, live specimen (No. 52504), station 2547, off Martha’s Vineyard, in 390 fathoms, 1885.
Owing to the very interesting character of the odontophore I have ventured to make this shell, although imperfect, the type of a new genus which is closely related to Choristes, for which Professor Verrill constituted the family Choristide and. placed it among the Tectibranchiata. Mr. Dall (1889) placed it between family — ? genus Separa- tista Gray and Calyptraside, genus Mitrularia Schumacher, ete.
Choristella brychia sp. nov. Figure 10.
This is a larger species of firmer texture than the preceding, although of the same number of whorls. Sculpture none. Color dirty white tinted with brown. Where not worn, the surface is slightly lustrous. Interior of aper- ture very smooth and lustrous, showing a sutural band of delicate rose color. Greatest width, about 4mm: height, about 3™™.
ub dead, imperfect specimen, station 2234, off Martha’s Vineyard, in 810 fathoms, 1884.
The Cyclostrema pompholyx Dall (Bull. Mus. Comp. Zodl., xviii, — p- 394, pl. xxviii, fig. 9, 1889), may prove to be another species of - this genus.
Cyclostremella gen. nov.
Shell minute, thin, semi-transparent, when fresh, planorbiform, of — few convex whorls, nearly symmetrically coiled, forming a con- cavely depressed spire and large umbilical cavity. Epidermis thin, nearly colorless. Nuclear whorl relatively large, smooth, turned downward, seen only in a basal view, leaving a small pit above. Suture deep and channeled. Aperture triangular-ovate, expanded — below, angulated above, with a relatively wide deep sinus just below the suture. Peritreme thin, simple, continuous, not modified, slightly attached.
The dark animal was drawn well into the shell. The operculum is very thin, almost colorless, broad-ovate, with the nucleus below the center and represented by a small smooth space indefinitely
Cyclostrema, Adeorbis, Vitrinella, and related genera. 141
defined by an indistinct line. From this arise numerous raised lines in the direction of the lines of growth, which are at first near together but diverge toward the outer margin, where they terminate just within the edge. Others arise between these about two-thirds their length.
Two large black eyes, close together on the top of the head, were the only feature that could be discerned in the much dried animal, the shell being removed by acid. The radula could not be found.
Type, Cyclostremella humilis sp. nov. PLATE XXII. figs. 8-8b.
Shell minute, white, nearly smooth, consisting of two whorls, besides the nucleus. Surface somewhat shining, but under the microscope it is covered with rather conspicuous sinuous lines of growth which are crossed on the peripheral region by several raised, unequally separated, spiral threads, varying in number in different specimens, so that in some they extend tarther above and below than in others.
Diameter of one of the largest specimens, about 1°5™™ ; height, about ‘7™™. One live and several dead specimens at stations 2112, 2274, 2277
> <4 “id,
2278, off Cape Hatteras, N. C.. in 153-16 fathoms, 1883-1884.
Lissospira profunda (Friele). See page 130.
A specimen of Cyclostrema profundum sent to Professor Verrill by Dr. Friele is much more transparent than any species from our coast which has been referred to Lissospira. The three very con- vex whorls form a small, elevated spire and abruptly enlarged body- whorl. The nuclear whorl is smooth, below which the entire surface is covered with numerous, raised, rather regular, evenly separated spiral lines which, under the microscope, are crossed and inconspicu- ously roughened by regular, raised, transverse lines which are more
evident on the interspaces ; the spiral sculpture becomes coarser on the base. The umbilicus is of moderate size, deep. The aperture is circular ; the peritreme is slightly injured but shows that it is con- tinuous and slightly attached to the body-whorl. The operculum is also typical. _ Greatest diameter, about 2°5™™ ; height the same. Yale University Museum, July, 1897.
142 K. J. Bush— Marine Gastropods referred to ;
EXPLANATION OF PLATES.
Puate XXII.
Figure 1.—Delphinoidea serpuloides (Montagu) Brown, p. 100. Front view of a speci. men (No. 9450) from Guernsey, England, in the Yale University Museum; x about 22. e Figure la.—The same, Basal view of another specimen from the same locality; x about 22. Figure 1b.—The same. Top view of another specimen from the same locality x about 22. Figure 2.—Lissospira diaphana (Verri}l) Bush, p. 131. Basal view of the type speci- — men (No. 38409) from station 2084; x 12. . Figure 3.— Lissospira proxima (Tryon) Bush, p. 130. Front view of the type specimen (No. 38443) from station 2115; x 12. } Figure 4.— Cyclostrema cancellata Marryatt, p. 97. Top view of the type specimen. Figure 4a.—The same. Basal view. Both are enlarged copies of the original figures. Figure 6.—Adeorbis subcarinatus (Montagu) Wood, p. 102. Top view of the nuclea whorl of the same specimen as figure 9; x about 35. Figure 6,— Circulus trilix Bush, p. 127. View of an animal with operculum, from station 2280; the shell having been removed with nitric acid; much enlarged. Figure 7.— Vitrinella multistriata (Verrill) Bush, p.124. Top view of a young speci men (No. 35733) from station 2109; x about 16. Figure 8.— Cyclostremella humilis Bush, p. 141. Basal view of one of the type speci- mens; x about 22. ; Figure Sa.—The same. Front view of another specimen; x about 22. . Figure 8b.—Top view of another specimen; x about 22. All are from station 2278. Figure 9.—Adeorbis subcarinatus (Montagu) Wood, p. 102. Basal view of a specimen (No. 9428) from Guernsey, England, in the Yale University Museum; x about 14. 2 Figure 10.—Circulus trilic Bush. Basal view of an adult (type) specimen (No. 35365) from station 2113; x 12. Figure 10a.—The same. Front view of the same specimen; x 12. Figure 11.— Vitrinella Tryoni Bush, p. 123. Front view of the type specimen from station 2278; x about 16. Figure 1la.—The same, Top view of the same specimen; x about 16. Figure 12.—Circulus triliz Bush. a, 6, c, Camera-lucida drawings. a, Portion of the radula; greatly enlarged, 0, A series of marginal teeth which turned outward in mounting; much enlarged. c, A single marginal hook; more enlarged. d, é, g, Free hand drawings. 4, Broad median tooth; e, slender lateral tooth; f, series of twenty or more sickle-shaped, marginal teeth as seen when magnified about 1,000 diameters. g, Series of marginal teeth with the invisible lower lines introduced. c Figures 2, 3, 10, and 10a, drawn by J. H. Emerton; figures 6 and 12, by the author ; the others, by A. Hyatt Verrill, are camera-lucida drawings (except figures 4 4a),
q =
,
eye: =
Cyelostrema, Adeorbis, Vitrinella, and related genera. 143
PLATE XXIII.
Figure 1.—Circulus supranitidus (Wood) Jeffreys, p. 127. Front view of an authen-
; i tic specimen from the “Crag”; x about 18.
‘Figure 2.—The same. Top view of another specimen (smooth variety) from the
same locality; x about 20. Both are from the U. S. National Museum.
‘Figure 3.—Circulus Dalli Bush, p. 126. Front view of the type specimen; x about
mn 19.
Figure 3a.—The same. Basal view; x about 12.
Figure 4.—Vitrinella multistriata (Verrill) Bush, p. 124. Basal view of an adult
specimen (No. 35733) from station 2109; x about 10.
Figure 5.—Skenea planorbis (Fabricius) Fleming, p. 100. Basal view of a specimen
from Provincetown, Mass.; x 15,
Figure 6.—Circulus Dalli Bush. Top view of the early whorls of the type specimen; -X about 17.
Figure 1.—Circulus liratus (Verrill) Bush, p. 125. Top view of the nuclear whorls
of a young specimen from station 2277; x about 20,
Figure 8.—Skenea planorbis (Fabricius). Top view of another specimen from Proy-
i incetown, Mass.; x about 15.
Figure 8a.—The same. Front view of another specimen from the same locality; x
about 13.
Figure 9.— Vitrinella helicoidea C. B. Adams, p. 122. Top view of a specimen from
station 2280; x about 13.
igure 9a— The same Front view of the same specimen; x about 13.
Figure 10.— Circulus trilix Bush, p. 127. Front view of a young specimen from
station 2276; x about 20.
Figure 11.—Circulus Duminyi (Requien) Jeffreys, p. 125. Front view of a specimen
_ from Naples; x 11. From the U.S. National Museum.
Figure 12.— Circulus liratus (Verrill) Bush, p. 125.. Front view of a young specimen
from station 2277; x about 15.
Figure 12a.—The same. Basal view of the same specimen; x about 16.
Figure 12b.—The same. Top view; x about 13.
Figure 13.—Leptogyra Verrilli Bush, p.136. Front view of one of the type speci-
mens from station 2174; x about 19.
Figure 13a—The same. Basal view of another specimen from the same station; x
about 19.
Figure 14.—Vitrinella muJtistriata (Verrill) Bush, p. 124. Front view of a young specimen from station 2109; x about 10.
Figure 15.— Circulus trilix Bush, p. 127.. Top view of the early whorls of a speci-
men from station 2276; x about 22.
e 16.— Choristella leptalea Bush. p. 139. Camera-lucida drawing of a portion of
_ the radula; greatly enlarged. a, One of the marginal teeth which was turned outward in mounting.
16 is by the author; the rest are camera-lucida drawings by A. Hyatt Verrill.
+r J
INDEX OF GENERA AND SPECIES.
Architectonica.............+06- 112 |
Archytea...... e 118 catenulata . -113 MGMIGRUUI on aes creme neice 118
Adeorbis. .....97. 102, 105, 125, 129 DSCC) ERR ABA ars stlos soAcac 104 SCAU Sicasidccs-cciccwepcwncue 104 GBUWIAUS ccs ccc sclesisleeiinss's 104 cyclostomoides ...........-- 104 | CLE RAD Bo iirc ico weitsic nc sin’aln 104 PPRVIIG occ vce we euss's teres 113 5 TMOVNGLUMLe.: p oseneeceee 104 | NEMALG: acl ce tciieae pivete = dar 104 | TINtICOIGES see aeene ila senor 104 | Nauti formisi. ede. pases 104 OHVACCUG sce. cecseemericane 104 | OPDIS NWI ince cecaaleete 104, 125 PULCHTALIS Cec eseeeane 102, 128 SINCELAy se. aoe eee
38 striatus ...102, 103, 104, 125, 128 | subearinatus. .102, 103, 105, 1a subimbricatus.............. supranitidus 103, 104, 125, ier | var. ee tricarinatus
CynisCaysasciececeen ae seb eeoe ue QTANUIALA cnc nc sees eens 107 WADOWICH waiecisisecabaios aie’ 108
Maronias, cach epcise cma es 102, 108, 109 spirula.. 108, 1
| Delphinoidea . 100, 101, 107, 109 PEORIA pond cist a cinns Ae depressa . ; resupinata , serpuloides . ,
| EO ee qrnSentor sac and 100
Delphinula.. 98, 102
STBCTA}IS eee toes oa 107 CANCEIATA Se sects seb ec ciess 98 MTN VAS. sam ra es o/elvieivio'a se 103 FETC HENIE. aoe aisle pees a oleate 98 WV IB want eiccisisiemews shana 10
Dillwynella........ wiesee 115, 120 TMLOGEBUR IS vais cle slsivis siclas + niriale 120
EDISCOPSIAge re cteer dee ates tees 119 OUALOB perseeisie ois peeled ieee 119
4) SPISC VOLS) jostle weloeisloisleee siete 111 AMOTWALA © apcm mets nets see 111 Multicarinata: 2c 2c. ess seen 107
Ethalia .... 106, 113, 115, 116, 120, “A
GULL Soe ioae anaaeoninéoarycc 104 |
Calceolinag ©... -....seeeceldo. 119 | pusilla 119 Choristella DPYChiIan =. asc leptalea pompholyx Choristes elegans var. tenera 139 _
a Rants 103, 107, 110, 111, es a BRO AISCEC OOOO AOOOSING MUM Glseoeses eee 110, 125, be var. sunanivaan niaeighaitaciek HIP ACUS eile seins eterna c 125, 126 [Tall thes aonSdegonnosaas aneiG 126 RUriatisyssce- acces 110, 125, 136 supranitidus ..... 126, 127 128 [Bebb been espinado as 126, 127, 128 | Cirsonella......... ~. e114, 115, 120 RUStrALIB eek tenement cas 120 | Cithna cingulata........ y Clathrella naticoides.. | Cochlholepisy Wee cecesss see | PATASIDIC ARS ans eeetate | Craspedostoma .... ...... elegantulum) ........00.-.. 108 Cyclostrema 97,100,101, 107, ie 129 | By ith) Se gaearsiodoc ood 99, as SPM Babar sacnasoocsocs areolatum ....... Aasoacsons 103 basistriatum .......... 180, 181 | Beaniiy cas accebectcrceraee rs 99 Dicarinatam Sess cesses cee 99 canecellata ........... 97, 98, 99 Cingulatum: 25 .g25. 22: 99, 132 cistronlamy - oo ace senses ss 99 curyistriaftum .......5.... 130 Cutlerlanaye cin spss aslcinte 115 Das owns -eetaeete 99, 1338 Var. ornatum...... .... 99, 134 GiaphanumM. cone. cee 99, 131 eDUrMen) io.c jc sbeaneeaes 99 CRCAV ALA. c.- claisiomia. oe le aisette lion 99 fnlPiduss sy. essen eee 99, 183 granulata......... isisenease 107 PUAN UUM sates scenes 99 Ase Vig atu &.cacecce. Madedant 130 AUUTT RULE ere crateteteieenieatreett 99, 185 THLVGAy neste sitet ne omiaeiste 110 ornate’. 255: .icnencet meee 1384 Petteraent o.yaecc. caseee ee 181 POM PHOly 0. westerns 99, 140 | DrotunduM ay .e-2se eee 130; 141
PIOXMA.~cen nee seam eenn pseudocancellata rugulosum......
Schrammii . serpuloides sp. spirala .
subexcatata sulcatum . tr. caripatas trochoides . tubercnlosa turbinum . valvatoides . Verrilli Watsont WO lice occa niainte Cyclostremella humilis
ANG Ala) Hesse Meee
atomaria . Sisie(naislohlelat ata ie diaphana. .. wach were weiss 116, 123 GUSMENRE! (jensen sess enone 116 PRE PBBLOMIA cine on scarne cane 106 INLOGEStaie cect mislo ca iateistene 116 multistriata ............... 116 FOCLUSH getngni ce itecne pete 116 pemistriate csc. acer nsnoee 116 syelh Gb tae Ancor Gonseacon soe eklG BETIOUR CUM & 5: ose <.ocieelsisies nies 116 SWPP CSAM fics tees 'ssiclee clea see 116
Ganesa....... 107, 114, 120, 121, 133 ADYSSICOLA 7. . cet acces eee 134 Dallinsc., Gesecessen cece 138, 184 nitidiuscula .......-.-.... 114 OPnBtars cts eaczcnacasesbes 134 PIULD OBB er cer cisesispieics 114, 135 PATIND tas tee tdeceaeeels 120, 1 SD. wee eeceeeceeesrsecececeese 134
Granigyra ........ 107,114, ay 1385 VISE Rete rs raceme toes 5, 185 pruinosa....... Hapntnd Side 135 BpiInDlOsa <7. cecceee ese , 135
Haloceras...... . 105
Maplocochlias . 121
| cyclophoreus 121
Helicella ........ 115
Helicelle . mah iS
Heliciella . 115 MULAD LIS cere eree cectnoee caer 15
LET Ga sGe soautlec Acoseod 100, 106, 109 GEpressa...., .<.0ssececce 98, 100 SENPUIOIGER we aacaenenass 98, 102
LOMA ex 8 errtamiee sicietwicistots are winrsis 129
Homalogyra densicosta ...... 127
Leptoeyra... ...22....- 107, 118, 185 @TAUNVC PE fh esis cw caine asians 137 inconspicua 137 Vernilli<. ... 186
Leucorbynchia 108, 121 Caledonica
Tilt eee. s - 98, 121
ilula we
Liotina.
Lissospira. 107, 118, 115, 120, 13 ADVARICOIA Cos. ot isc ecs ces SHAN Gere eeenaronaeatninn sone i DABSIStIIGUBI cae sewers 2 <s 129 CINTA ER Sriae cet clerte!<clsinicis'e. 132 BOIVOE@! wes te cet +e vcciesnie 132 DST eres ecpcetresesc’s 133 Gisphank ean tevhcee vawasen 129 Thee) NGA oc (ann oo Sena 185 OTTSUR LS. 148 Galen's oan cne naps 134 POLINA gen cine vee esinens 141 PFOMIMAM ewklse civic sie s dsiroie ele 129 DITUIMOBA VEN Ce Linkins sbieaeiets 185 TATINGUD Peeee: ctiyarmclne 120, 134 BDLNUIONB oe paiiceais nis 5s vine 135 METIGGDE Joadadenca Jaden Ssiacers 182 IWVAINGL seats ceteiisiare:s ctor lersiciewiers 132
Margarita......... 102, 106, 111, ie UOMO cnc csistenside ns Sone is helicina....... 101, 108
MECRREVIOMB. cs ccanwelcenc'se Saas M7 WALCLOUL, i soncniscn encase: «ALT
| MOlleriopsis
| Rossiteria ; 2. ..cueveaaea
| Tubiola
Microtheca......... aia crenellifera MGllOrda oc vccarwcne cle senna costulata
abyssicola ... sincera ....... - 137 as oeley sap aeee 108, 110 | Obyoluta...........0+. «oo. 108
Pseudorotella anomala Garinata.--., veces diaphana ...... cana minuscula ...... seat chidgipe DUBIN. 2. Coc eas eesees semistriata .... striata
Rissoa (Setia) re
Rotella . Seen anomala..... carinata.... eryptospira . BA : diaphana a onde ;
Separatista Chemnitzii <.. asses eee .- 105 eingnlata) 3... cuwekes = Grayil .... 3 sees : separatista ............65 a!
Serpulari@..:.. seas centrifuga
Skenea.. lirata ..: .o2: .oeaeee planorbis) << -o--0sem sich Trilieseeee unto eceeee ee
Solanderia...
Solariorbis
Solarium ...
apo
Teinostoma ..106, 108, 115, 117, 119 anomalum....... eercesc i CarinatuM............6006 1 pi tenee age Sexea cane iis; 119 diaphanum.. 17 minuta
Thawielis ask wo manemers romettensis ....... Tharsigh2...ceeeee F Trachysma ... delicatum . VAT. OX PANGS... .. 05-5052 anie 1 Trochus
striatus’ <2). eee
cornuella .......
Turb helicinus sc ncnes ese oen helicoides ......... brats HIVEA « <u cee eens eee eee separatista ......05 sees Umbonium ........ .. 115, 116, 1 vestiarium ....... <stocsaeee
Valvata striata........102. 108, 125 Vitrinella 97, 105, 107. 108, 8 1 117
earinata diaphana 107, 116, 117, 118, 3 helicoidea 105, 106, 107, 1 23 hyalina... .2 sees qu2 106 interrupta.............. 106, 1 Megastoma ...........- multicarinata.......... 0%, 11 multistriata.... ins striata
x sss Bene Sn Be ®oR Ge tor “rr SS SSS a a a ee a a er a eer ee ee eee ee <r i ee ee Oe is Se ee ee Se eet 66 ee ee Oe, Se ee a ae
_ IV.—Revision oF cERTAIN GENERA AND SpEcIES OF STARFISHES WITH DESCRIPTIONS OF NEW FoRMS. By A. E. Verritt.
Family GONIASTERIDZ.
Goniasteride Verrill, Trans. Conn. Acad. Sci., i. p. 348, 1867; Perrier, Revision, Arch. Zool. Exper. et Gen., iv. pp. 281, 283, 289, 291, 1875; Speci, V., p. 1, 1876.
_ Goniasteride (pars) Forbes, 1840.
Pentacerotide (pars) Gray, p. 275, 1866.
Pentagonasteride Perrier, 1884; Sladen, p. 260, 1889.
Tuer generic nomenclature in this family has become very much confused for several reasons.
- The genera themselves are difficult to limit and define, and scarcely any two investigators, in the past, have agreed as to their number or limits. Nor have they agreed as to what characters should be con- sidered as of generic value. This was the case, in a very marked degree, and very unfortunately, when the ancient and comprehensive genus Asterias was first divided into numerous genera by J. E. Gray, in 1840, and by Miiller and Troschel, in 1842. In these two works, issued within a brief period, there was very great diversity, both as to the number of genera and their names. In the genus Astrogonium of M. and Tr. four of Gray’s genera were reunited into one. In - Goniodiscus M. and Tr., Jive of his genera were also reunited. For about half a century most subsequent authors have tried to take an intermediate course, but gradually more and more of Gray’s genera have been adopted, though often with their limits more or less modi- fied. Fortunately Gray assigned definite types to his genera, and in his later works he described and figured many of the species, so that in most cases his groups can be readily understood. Moreover he followed, pretty closely, the generally accepted rules of zoological nomenclature, which has not always been done by later writers.
_ The failure of several writers to follow the ordinary and accepted tules of priority has led to much needless confusion of names. The failure to recognize the priority of Gray’s generic names has been he cause of more confusion than any other one thing, throughout e group of starfishes.
‘ he efforts that certain writers have made, from time to time, to restrict or apply certain generic names to species or groups to which they were not originally given, has repeatedly led to confusion and ‘Trans. Conn. Acap., Vou. X. Aveust, 1899. 10
146 A. EF. Verrilli—Revision Genera and Species of Starfishes.
uncertainty. This applies strikingly to Goniaster, Astrogonium, Dorigona, ete.
A few recent and prominent writers, especially Perrier aud Sladen, have restored the ancient names given by Linck (1733) to certain genera and species of starfishes, thus displacing names well estab lished under the binomial system. Linck was a very able naturalist for his period, but he was not a binomial writer, and his names cannot. properly be allowed priority over those established under the binomial system.
The name Pentagonaster is the only generic name in this family to which this remark applies. 4
Perrier himself, although he restores several of Linck’s names of species, does not go so far as to try to restore others that have equal _ claims to priority, for to do this would overthrow the well known names of several common European species.* Nor has he proposed” to restore the names of Seba, which have equal claims to recognition,
In the following pages I propose to briefly review the history of some of the earlier names and of the more important groups to which they have been applied by various writers, in order to show, if possi- ble, to what particular groups certain of these names ought rightly to be applied, in accordance with the generally accepted rules of biological nomenclature.t
* Among the names adopted by Perrier, and also by Sladen, from Linck, are oculata, under Cribrella; planus, under Hippasterias; corniculatus, under Ctenodiscus. Neither of these can be justified.
+ Among the recognized rules that I follow, and which need to be applied to this group, are the following :
A.—Strict priority to be applied to all names properly published in actual binomial works, in general dating only from Ed. X. of the Syst. Nat. of Linné,
B.—Exclusion from the rule of priority of names taken from earlier polynomial - writers, unless adopted by later binomial writers. In that case they should date only from their introduction into binomial literature.
C.—When an old composite genus has been divided by a later writer, the original name must be kept for one of the component groups, and for one or more of the species originally included by name. If a definite generic type was given by the original author, the name must remain with that type. If no type was mentioned, the mere position on the page cannot fix the type. Nor does it follow that the jirst species named was the type, unless so stated originally, for many early writers arranged their species alphabetically, or in some other arbi- trary way.
D.—A composite genus having been subdivided and the original name definitely applied to one of its parts (in accordance with rule C), it must ever after be kept for that group (or some part of it) just the same as if it had been originally sO
A, E. Verrill—Revision Genera and Species of Starfishes. 147
This brief review is, however, by no means intended as a complete history of the subject. Generally only the works that seem essential to the object in view will be referred to here. <A fuller discussion must be left to a much more extensive work on American starfishes, which is now well advanced towards completion and in which most of the genera and species will be well illustrated.
1, The first generic name applied by binomial writers to any sub- division of the old genus Asterias (Linné), and pertaining to the present family, was Goniaster. This name was proposed by L. Agassiz, in 1836, for the pentagonal starfishes, collectively, includ-
ing representatives of more than one family.
This name was adopted by Forbes in 1841 (Brit. Starfishes), in the same sense, for he included in it such diverse forms as Hippas- teria and Asteropsis, without assigning to it any definite type.
~Miiller and Troschel used it in the same way, in 1840. Dr. J. E.
Gray, in 1840, adopted the name for a very restricted group, with a definite diagnosis, and named as a type, G. cuspidatus, a well- known species and one of those given by Agassiz as examples of his genus. This species should, therefore, remain as the type of the restricted genus.
In 1842, Miiller and Troschel reunited Goniaster, Pentagonaster, Tosia, and Hippasteria of Gray into a single genus, to which they applied the new name, Astrogoniuwm. If these four groups really
constitute only a single genus, it is evident that Goniaster (emended)
should have been adopted as its name. Il. The name Pentagonaster was first used, under the binomial system, by Gray, in 1840. He applied it to a particular type (P.
applied. In other words, a generic name correctly applied to a restricted group has just as much claim to priority, in the new sense, as a new name would have.
E.—When a generic name is a real synonym of another earlier one it should be dropped from the system, unless it had a different type-species when first pro- posed. In case the two types belong to different subdivisions of a composite genus both names may be retained in a modified sense. In cases where two names are only partially synonymous, both may be used if they can be properly
restricted to distinct subdivisions of the groups to which they may have been
originally applied (in accordance with rule C). F.—The application of an old or discarded name to a species or group not included in the group to which it was originally applied is to be avoided as lead-
ing to confusion and instability. A name once dropped from the system, for
good cause, should fall into disuse in every other sense. To use a discarded generic name for a new genus in the same class or order (as if it were a new name), should never be thought of, for it is sure to cause confusion. (Goniaster and Astrogoniuwm among starfishes afford examples of incorrect transposition of
names.)
148 A. #. Verrilli—Revision Genera and Species of Starfishes.
pulchellus) and gave it a definite diagnosis. His use of the name should, therefore, have priority, and the name should not be applied to any other group, unless P. pulchellus be included in a larger generic group, as was done by Perrier, in 1876, and by Sladen. But in the latter case, Goniaster should have had precedence over Penta- gonaster, for such a comprehensive group, on the ground of priority.
In a later work (1894) Perrier separated Gray’s Pentagonaster as a distinct genus, but he ignored the original application of the name by Gray, and adopted the later name, Stephanaster of Ayres, for Gray’s genus.
At the same time he retained Pentagonaster for a large group of species closely allied to P. australis, which was the type of Tosta Gray, 1840. This arrangement was based on the fact that Linck, in 1733, had figured an indeterminable species, apparently of the latter group, under the name of Pentagonaster. But Linck, however great his merits may have been, was certainly not a binomial writer. Most of his names were trinomial or polynomial, and there is noth- ing to be gained, except increased confusion, by trying to give priority to the names used by such polynomial writers, in place o later binomial names that have been definitely defined ey fixed in the binomial system.
Perrier, in 1876, restricted Goniaster to a genus containing only a single species, Pentaceros obtusangula (Lam.) Gray. This species was not mentioned by Agassiz in connection with the genus Gonias- ter. Its use by Perrier is, therefore, in a new sense and like that of
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an entirely new name, and was not justifiable.*
Sladen (1889) has also restricted Goniaster to the same type.
Ill. Yosia was also proposed by Gray, in 1840, for a definite group of this family, with 7. australis as the type. Several other species were added to it by him in 1847. This name has been ignored by most later writers on starfishes, or else it has been placed as a partial synonym of Pentagonaster (Sladen, Perrier) or Astrogo- nium (M. and Trosch.), If Gray’s restricted genus Pentagonaster be deemed a valid one, as by Perrier (1894), then Zosia should be used for the large group of species, called Pentagonaster by Perrier and Sladen, agreeing well with Gray’s diagnosis and type-species (7. australis).
IV. /Hippasteria was proposed by Gray, in 1840, for the single type- species, JZ. phrygiana (as H. Europea, ete.). This name has been so genel ig) adopiae wis later writers that it needs no discussion here.
i A I a, a Sle ee Cee pee ee ae Ee
* The restriction of Goniaster to its correct eer leaves this genus without a name. Therefore I would propose for it Pseudoreaster, with P. obtusangulus (Lam.) as its type and only known species. :
A. E. Verrill— Revision Genera and Species of Starfishes. 149
V. Calliaster proposed by Gray, in 1840, for the single type- species, C. Childreni, is very distinct from the genera already named, not only on account of the spinose plates of both surfaces, but also by reason of its very different adambulacral spines, ‘ VI. Hosia. When Gray established this genus, in 1840, he referred to it only H. flavescens. Perrier (1876) has redescribed the types of this species and refers them to two distinct species of true Anthenea (Gray, 1840). Therefore Hosia becomes a synonym of Kho latter. In 1847 and 1866, Gray added another species (HZ. spi- nulosa) to Hosia, but according to Perrier (1876), who reéxamined Gray’s type, this species belongs to a different genus. He referred it to his section C of Pentagonaster. It has spinulose marginal plates, and also vaivular pedicellarie. It is probably an immature species of Zosia, or of some closely related genus.
_ VIL. The names proposed by Miiller and Troschel, in 1842 (Syst. Ast.), are next in order.
_ Astrogonium, as stated above (p. 145), was formed by uniting four ‘of Gray’s genera. It thus became a composite group without any definite type, and not very different from the original group called Goniaster (1st section) by Agassiz. In 1847 and 1866, Gray applied the name to a more limited group, including A. granularis (Retz.), which is nearly allied to Zosia, together with species now referred to. Odontaster.
_ If it were to be used at ali in the modern system, it should be restricted to the group containing A. granularis. But as it was an artificial group and should have had no real status originally, it should properly drop out of use except as a synonym of Gray’s
_ By Sladen (1889) Astrogonium was restricted to Gray’s genus Pentagonaster = Stephanaster Ayres.
Perrier (1894) has used it improperly for a very different group, (= Pseudarchaster + Aphroditaster Sladen), including several deep-sea
VIL. Goniodiscus. M. and Troschel (1842) constituted this nus by reuniting five of Gray’s genera, together with forms un- own to Gray. Perrier (1894) has very judiciously restricted the name to those species that have stellate or 6-lobed abactinal plates, ncluded in it by M. and Trosch., such as G. euspidatus (Lam.), and 0 1 a sense it should be hl
cu a here.
150 A. FE. Verrill—Revision Genera and Species of Starfishes.
Goniaster (Agassiz), Gray (restr.). Type G. cuspidatus Agassiz.
Goniaster (pars) Agassiz, Prod. Mem. Soc. Neufch., 1836.
Goniaster Gray, Ann. and Mag. Nat. Hist., vol. vi, p. 280, 1840. Type G. cus- pidatus. Synopsis, p. 10, 1866 (non Perrier, 1876, nec Sladen, 1889).
Pentagonaster (pars) Perrier, Revis., Arch. de Zool., v, p. 24, 1876. Sladen, Voy. Chall., xxx, p. 264, 1889.
Astrogonium (pars) Mill. and Trosch., Syst., pp. 52, 56, 1842.
Phaneraster Perrier, Exp. Sci. Trav. and Talisman, pp. 334, 337, 387, 1894, (Type G. semilunatus = cuspidatus.)
As already explained, the genus Goniaster was restricted by Gray, in 1840, to a definite and well known type (G@. euspidatus). Perrier, in 1894, has, quite unnecessarily, applied a new name (Phaneraster) to exactly the same group, with the same type. Whether Gonéaster, as here restricted, is worthy of generic separation from the great group called Pentagonaster by Perrier and by Sladen, must remain, for the present, a matter of personal opinion, but if they should be reunited under a single generic name, Goniaster would be the name that ought to be chosen for the whole group, if we are to follow the generally accepted rules of binomial nomenclature. (See above, p. — 146.)
The principal character by which the present group has been distinguished is the presence of one or more large, stout conical tubercles or spines on more or less of the dorsal marginal and- abactinal plates, in adult specimens; or of verruciform swellings in the same situations, in the young. In most adults these conical spines form a central group on the disk and five large radial groups, but the number of plates that may bear spines is variable; sometimes they occur on nearly all the dorsal plates.
The marginal plates are large, thick, convex, not numerous, and usually naked, except for one or two marginal series of granules, but they are more or less granulous over the surface in the very young. They are more numerous in the ventral series. Those in the dorsal series may not decrease regularly distally; the last one is sometimes as large as, or even larger than, the one that precedes it. The apical plate is small, conical. The actinal plates are large, polygonal, and crowded, mostly in series parallel with the adambulacral plates, and covered with coarse granules; the granules on the center of the plates are often larger and may be like small tubercles. Sometimes part or all of these plates bear high, slender, spatulate pedicellaria. The adambulacral plates and spinules and the dentary plates are essentially the same as in Pentagonaster or Tosia. ‘The adambulacral spinules are numerous and closely crowded in three or more rows 3 the row next to the furrow-series is largest.
A, £. Verrili— Revision Genera and Species of Starfishes. 151
The abactinal plates are rather large, polygonal or roundish, coy- ered with crowded, short, angular granules, with a larger marginal series; sometimes they also bear pedicellariw. Between these there are often, in adults, many small ossicles, usually bearing groups of few granules. Papular pores are present between most of the abac- tinal plates, except in the small interradial areas.
In adult specimens some of the distal, lower, marginal plates bear small conical spines or tubercles, in some species.
Pedicellariz have been observed only in G. Americanus, where they are sometimes numerous, both on the actinal and abactinal plates, and they occur also on the sides of the dorsal spines and marginal plates. They are small, high, slender, pincer-shaped, with spatulate blades and corresponding sockets on the plates. (See below.)
Perrier erroneously states (1894) that pedicellariz are not found
in this genus. They were described by me in 1871.
When very young (up to 12 or 14™™ in diameter) there is no appearance of dorsal spines or tubercles and the marginal plates are few in number and granulated. In this stage there appears to be no obvious distinction between this genus and Pentagonaster or Tosia.
Such specimens were mistaken by Perrier for a distinct species
_ (Pent. parvus). They will be more fully described on a later page.
Goniaster Americanus Verrill.
Goniaster Americanus Verrill, Amer. Journ. Sci, vol. ii, p. 280, 1871.
Pentagonaster semilunatus (pars) Perrier, Arch. de Zool. exper., V, p- 24, 1876.
Phaneraster semilunatus (pars) Perrier, Sci. Exp. Trav. and Talis., p. 388, 1894.
Pentagonaster parvus Perrier, Mem. Etoiles de mer, Nouv. Archives du Mus. d@’Hist. Nat., vi, p. 231, pl. vii, figs. 7, 8, 1884. (Young.)
PLaTE XXIVa. Ficures 1, 2. Puate XXVI. Ficures 1-6.
This species was originally described by me so minutely that it is not necessary to repeat the general description of the adult. The type was from rather shallow water, off the coast of South Carolina,
This type specimen, which is in the museum of Yale University,
has a large number of high, pincer-like pedicellarie, with two
slender spatulate or spoon-shaped blades, and a slightly enlarged articulating base; the blades are sometimes straight, but often more or less strongly curved to the right or left. The blades, when fully _ expanded, rest in socket-like depressions of the plates, which corre- i spond in shape and curvature with the blades, so that the two belonging to a pedicellaria with curved blades, form, when taken
152 A. £. Verrill—Revision Genera and Species of Starfishes.
together, a crescent-shaped or semicircular pit, with a round central pore and a wider rounded depression at each end. Sometimes one or two granules exist close to the pedicellariz, and when rubbed of the pits that they leave make the markings on the plates still more complex.
Pedicellariz of this peculiar form are present on a large propor- tion of the actinal plates; on some of the marginal plates; on the borders of the spinose abactinal plates, around the bases of the spines, 1 to 6 on a plate; on the basal part of the spine itself; and on those abactinal plates that do not bear spines, 1 to 4 or more.
On the actinal plates they are variously placed, and irregularly oriented ; most of the plates have but one, which is most commonly near the center, but many have two; those plates in the row next to the adambulacral plates usually have two or three. The pedi- cellariz on the abactinal plates and on the spines are smaller than those of the lower surface, but have the same form and similar sockets. Each pedicellaria of the actinal and abactinal plates occu- pies a small, slightly elevated, smooth, rounded or ovate area, sur- rounded by granules. A pedicellaria and a stout blunt tubercle co-— exist on some of the actinal plates, near the jaws.
Between all the abactinal plates, except those of the small inter- radial areas, where there are no papular pores, there are small inter- mediate ossicles, the larger of which bear small circular or angular - rosettes of about 5 to 9 prismatic, flat-topped granules, like those | around the margins of the large plates. One to three of the granules occupy the center of these groups. Between these sma}l rosettes ‘there are many small irregular groups of two or three similar angu- lar granules, intervening between the numerous and rather large papular pores, of which there may be ten or twelve around the larger plates.
The madreporic plate is very large, somewhat swollen, with fine gyri. The apical plate is small and conical, similar in size and form | to the tubercles of the distal marginal plates.
The two distal pairs of dorsal marginal plates are in contact medially. On each of the distal adambulacral plates there is.a_ single large, obtuse conical spine, outside the furrow-series of slen- der spinules. These spines are longer and larger than the more ~ numerous corresponding spines of the more proximal plates. There are usually, in large specimens like the type, four stout, prismatie, blunt, crowded spinules on each plate, in the furrow-series, as in G: Dinara, instead of three, present in G. cuspidatus.
A, FE. Verrill— Revision Genera and Species of Starfishes. 153
A fine series of specimens, of various ages, was dredged by the _“ Albatross,” off Florida and in the West Indies. ,, Some of the variations noted among these are as follows:
A.—Station 2363. One large example.
a Lesser radius, 35™™ ; greater, 65™™,
_ Most of the dorsal marginal plates, except distally, bear a high, acute, conical spine. On each ray the last dorsal marginal plate is elongated, subconical, with a small terminal spine. It looks as if it might have been about to divide into two or more plates ; or as if two or more had abnormally consolidated. The distal lower marginal plates bear rudimentary conical spines. There is also a group of 2 to 4 small obtuse tubercles on each jaw, around the mouth. The abactinal plates bear a central group of 9 or 10 large, high, acute, conical spines and four or five rows of about 6 or 7 on each radial area, with shorter rows of 2 to 4 on each side of these.
B.—Station 2373. One large example. Nearly normal, but 4 to 6 of the interradial plates have very large, stout, transversely compressed spines, bilobed at the tip.
C.—Station 2318. One very large.
Similar to A, with 18 or 19 pairs of marginal plates, the distal ones regularly decreasing. ‘Three or four of the distal, dorsal mar- ginal spines bear acute conical spines.
No tubercles around the mouth on the jaws.
D.—Station 2316. Six examples.
_ These vary in size from 40 to 54™™, in larger radius. ‘They have m 13 to 16 pairs of marginal plates on each side. All have tapered, acute rays, with a small conical apical plate. They agree pretty closely as to the lower surface. The more spinose examples ve a central abactinal group of 6 or 7 conical spines and there are 4 to 6 (usually 5) spines in each radial row ; nearly all the upper marginal plates have a single stout, conical spine. In others there are but 3 to 5 spines in each radial row; in some 8 to 10 spines are irregularly scattered over the abactinal surface. In some more or less of the spines have been broken off, leaving a oth scar in their places. In some examples the dorsal marginal es bear high, acute, conical spines; in others low, blunt, cones or tubercles with broad bases.
The lower marginal plates bear a variable number of spines, toward the end of the rays; most frequently there is a small group
154 A. #. Verrill—Revision Genera and Species of Starfishes.
of spines on the 4 or 5 distal plates ; in others only one; in some none at all. These variations may occur on different rays of the same example.
E.—Station 2406. Three examples.
The two larger are much like those described above. The smallest has the smaller radius 15™™; the larger radius, 23™™. It has 12 pairs of marginal plates on each side. The dorsal ones are thick, convex or rounded, and some are beginning to swell up in the middle to form tubercles or spines. The distal lower marginal plates show sub-conical elevations, where the spines are beginning to grow. Some of the abactinal plates show an elevation in the center, where the spines are beginning to develop.
F.—Station 2315. One example.
Similar in size to the one last described. Ten pairs of marginal plates on each side. They are of the usual shape and many of the dorsal ones show a central, low, conical elevation or rudiment of a developing spine. The abactinal plates are without evident tubercles,
G.—Station 2374. Four examples. ; Similar to the last in size. Three have 10 pairs of marginal plates — on each sides; the other has 12. Most of the dorsal marginal plates’ have conical or subspiniform elevations. The abactinal radial plates have 3 or 4 conical spines in a row, nearly like those of adult ex- amples. There is also a central group of spines on the disk, some-— times as many as five.
H.—Station 2370. One, very young. (Specimen figured, Pl. xxv, fig. 6.)
The larger radius is 5™™, Six pairs of marginal plates on each side.
Apical plate broadly triangular. No marginal nor abactinal tubercles.-
Abactinal and actinal plates surrounded by a single series of granules.
I.—Station 2370. No. 18,457, specimen figured. One young example.
Radii 12 and 17™", Dorsal marginal plates 10 pairs on each side, all short, broad, strongly convex. Ventral marginal plates 14 on a — side. Small, slender pedicellarie with long spatulate blades oceur on many of the actinal and abactinal plates. No trace of spines on the abactinal plates, which are finely granulated over the central area, with larger marginal granules; a few very small intermediate ossicles occur between them, each with 1 to 3 small granules. Papular pores are few and small, but seattered over most of the radial areas, The adambulacral spinules are in four longitudinal series; those next
A. E, Verrill—Revision Genera and Species of Starfishes, 155
to the furrow-series are distinctly larger than the others, two to a plate on the proximal half of the series ; those of the furrow-series are slender, equal, obtuse, regularly placed in a row, four to each plate. One or two of the distal lower marginal plates of each series bears a small conical tubercle, but the dorsal plates are smooth, naked, without tubercles, though considerably elevated centrally.
I have also examined four young specimens of this species from the “ Blake” Exp., preserved in the Museum of Comparative Zodlogy.
Three of these (a, c, d) were types of Pentagonaster parvus Perrier. They agree perfectly with those of similar size collected by the steamer “Albatross” in the West Indies. (See H and I above.) With the latter they form a complete series, connecting the smallest with the full grown examples from the same region.
The smallest of the specimens (4) from the “ Blake” Exp. is from station 253. It is enumerated under P. parvus by Perrier, but is not marked as a type, but it agrees with the others. Its lesser radius is 7™™; greater radius, 10™™. In this there are, for the most part, four upper marginal plates, above and below, on each side; but in one case there is a small triangular plate interpolated between the first interradial and the next normal one, while there is a normal plate next the apical one, so that there are four marginal plates on one side of this ray and the adjacent semi-margin. On another interradial margin there is a small, triangular, odd interradial marginal plate of the upper series, similar to that in Odontaster.
The lower marginal plates are usually six to a side, but on one margin there are seven. The distal plate of some of the series is small and only recently developed. The marginal plates of both series are covered with granules. The papular pores are few in number, in small radial groups.
The specimens next in size are 25 to 35™™ in diameter (types of P. parvus) and usually have six marginal plates on each side, above and below. In the smaller of these the upper and lower marginal plates and the actinal interradial plates are nearly or quite covered with small granules, but in the somewhat larger specimens more or less of the central area of these plates is naked. Most of them show a distinct central swelling where the conical tubercles would have appeared later. In some the abactinal plates are entirely covered
with granules, but in others the central area is naked, the amount of naked surface increasing with age, but not regularly so, The papular pores increase in number with age and cover more and more of the Median radial areas and the central area of the disk, but these areas have no sharp boundaries.
156 A. E&. Verrill—Revision Genera and Species of Starfishes.
List of Specimens of Goniaster Americanus taken by the Albatross, in the West Indies.
Station 2315, 37 fathoms, No. 10071, 1, young.
“ 9316,50 « 10076, 6, young.
tc DS Beas re 10821, 1, very large.
Cy PBIB AS ib 10876, 2, half-grown. 5 “« 9363,21 « 10618, 1, large.
es 2370, 25 CT) pean Pace oe 7, young.
eS Sie. ea BD eee 3, young.
SSE POM os 20 =e 10387, 1, large; 11, young.
Mo 2814, OG ee 10820, 6, young and half-grown. * “2374, 26 Y 103840, 5, young.
. 2406, 26 4 10459, 3, half-grown.
6 AO TS Oe ee te ee 6, young and old.
Specimens examined from the Blake Expedition.
a -Station 32, 95 fathoms. b 953, 92 c “ 276, 94 « d. “ 996, 84-129 «
Goniaster Africanus Verrill. Goniaster Africanus Verrill, Amer. Journ. Sci., vol. ee p. 131, 1871. PLATE XXV. Ficures 1,
Perrier (Revis. 1876, p. 24) united this species and G. Ameri- canus with G. cuspidatus of the East Indies. He showed very well, by his comparison of a large series of specimens, that the number and precise form of the dorsal spines and marginal plates are vari- able in specimens from each region and cannot be depended upon separate the species.
It has long been known that the number of marginal plates in all starfishes increases with age, and that their shape also varies with age, also that the spines increase with age.
Perrier, however, did not make careful comparisons of the much more se tant ai acters to be derived from the size and character of the granules, tubercles, and spines of the plates; presence absence and shape of pedicellarix ; form and character of the adam bulacral spines; size and form of the small ossicles and gran between the abactinal plates.
In all these characters G. Africanus differs decidedly from @. Americanus, if the numerous specimens of the latter that I have
f
A. E. Verrill—Revision Genera and Species of Starfishes, 157
been able to examine are to be depended upon. I have not had a
sufficient series of the Kast Indian G. cuspidatus for examination to
warrant me in making so positive a statement as to its distinctness from G. Africanus, but the published descriptions indicate impor- tant differences.
In addition to the marked differences between G. Africanus and G. Americanus in the number and character of the dorsal spines and marginal plates, I wish to call attention to the following points. G. Africanus (type) has no pedicellarix, above or below. The actinal plates mostly have a central cluster of three to six or more unequal rounded tubercles, much larger than the granules. No specimens of G@. Americanus that I have seen have this character, though some of the largest ones may have one or two central tuber- cles, on a few of the plates near the mouth. The granules of the abactinal plates are much smaller than in the latter and are more rounded, with less differentiation of the marginal series. The small, intermediate abactinal plates seldom bear distinct rosettes of gran- ules; but usually appear as small, round or oval, smooth-topped ossi- cles, on a level with the other plates.
In having the distal pair of dorsal marginal plates larger and more swollen than those that precede it, and largely in contact medially, this species agrees with Pentagonaster pulchellus and allied species. I have seen no specimens of G. Americanus having this character, nor is it said to occur in G. cuspidatus.
It is my intention to fully illustrate these species in another arti- ele, now in course of preparation.
G. Africanus is native of the West Coast of Africa.
G. Lamarckii (Astrogonium Lamarckit Mil. and Trosch., p. 56, 1842; Pentagonaster Lamarckii Per. (Revis., p. 29, 1876) is from
an unknown locality. It has rounded dowal marginal plates and
seems quite distinct from the other species.
> _ Pentagonaster Gray, 1840, 1866. Type P. pulchellus Gray.
x
Stephanaster Ayres, Proc. Boston Soc. Nat. Hist., iv, p. 118, 1851. Type S. elegans Ayres=pulchellus Gray.
Pentagonaster (Sect. A, a, pars) Perrier, Revision, Arch. Zool., Vis Ps den Ley Gs
Astrogonium Sladen, Voy. Chall., xxx, pp. 263, 280, 1889.
Stephanaster Perrier, Exp. Trav. and Talisman, p. 402, 1894.
The name Pentagonaster was first introduced into binomial
nomenclature by Gray in 1840. By him it was definitely detined,
a
and a well-known species was given as the type (see p. 147, above).
‘Therefore, the name should be used for the group thus limited by
him,
158 <A. E. Verrilli—Revision Genera and Species of Starfishes.
The following species apparently belong to the genus as restricted by Gray (1840):
P. pulchellus Gray. New Zealaud to China.
P. abnormale Gray. Unknown locality.
P. Bourgeti Perrier. Cape Verde Islands.
P. Gunnii Perrier. Tasmania and Australia,
P. Dubeni Gray. South and West Australia.
Tosia Gray, 1840. Type T. australis Gray. Astrogonium (pars) Mill. and Troschel, Syst., Ast., 1842. Tosia Gray, Synopsis, p. 11, plates iii and xvi, 1866. Pentagonaster (Sec. A, b, pars) Perrier, Revision, p. 20, 1876. d Pentagonaster (pars) Sladen, Voy. Chall., xxx, p. 264, 1889. Perrier, Exp. Tray. et Talism., pp. 389, 390, 1894.
Under the ordinary rules of priority, in zoological nomenclature, there is no valid reason why Zosia should not be adopted for a large part of the species included by Perrier and by Sladen in the genus Pentagonaster, providing we are to consider this group generically distinct from Pentagonaster Gray. This question of the generic dis- tinction must still be regarded as doubtful by many students of the group, though Perrier, in his later works, has definitely separated them, as shown above.
The only obvious difference, hitherto pointed out, that may be considered as of generic value, is the gradual decrease in size of the marginal plates distally, so that the rays are sub-acute, instead of — the distal ones being larger and swollen, asin the true Pentagonaster,
But there are species of the latter in which the distal plates are only slightly larger than the others, while the amount of decrease in the plates of the species of this group is variable. Moreover in the restricted genus Goniaster, one species (G@. Africanus, type) _ has the distal dorsal plates more swollen and larger than those that precede it.
Nevertheless, since the marked enlargement of the distal plates indicates a different law of growth in species so characterized, it seems desirable to keep the two groups separate, at least until truly intermediate forms become known.
In the typical species of Zosia, the marginal, abactinal, and usually some of the actinal plates have a naked central area, with one or more rows of granules around the margin. But the extent of — the granulation varies more or less individually, and also according to the age of the specimen. Therefore we cannot regard this as a matter of much importance, generically.
A, EF. Verrill—Revision Genera and Species of Starfishes. 159
The typical species appears to be destitute of pedicellariz, but small, high, pincer-like pedicellariw, with spatulate blades or chisel- _ shaped blades, occur in many of the allied species more recently described (e. g. Perrieri, Vincenti, hesitans).
In many of the species of this group all the dorsal marginal plates, except the last or two last pairs, are separated by one or more rows of abactinal plates. But in several deep-sea species (Perrieri, etc.) three to five distal pairs of marginal plates are in contact medially. In some species different individuals have been found to vary, in this respect, from those having only one pair joined, to those with three or four pairs joined.
Considerable variations also occur among the species, in respect to the character and arrangement of the adambulacral spinules, in the number and arrangement of the papular pores, and in the form of the abactinal plates.
Most of these characters are not sufficiently constant nor important for generic divisions, but may well afford grounds for dividing the group into convenient sections. (See p. 160.)
Sladen, in his great work on starfishes, included in his genus Pen- tagonaster not only those that are here separated as typical Goniaster, but also others that apparently belong to Mediaster and Hoplaster, besides some that belong perhaps to undescribed genera.
To Mediaster I refer three of his new species: viz. P. Japonicus, P. Patagonicus, and P. arcuatus. But as the existence of internal connecting ossicles between the abactinal plates has not been ascer- tained for either of these species, this reference is based on the general appearance and on the character of the plates, spinules, and pedicellariz.
His P. lepidus appears to be a true Hoplaster. It has odd inter- radial marginal plates and all the plates are spinulose. LP. gibbosus Perrier also appears to be generically distinct, as well as P. inter- medius and P. dentatus. If these forms be eliminated, the genus becomes more homogeneous and better capable of definition, though it still remains an extensive group.
Perrier, in his later works, has generically separated numerous species that he formerly referred to Pentagonaster, such as Losaster, Odontaster, and the forms that he refers to Dorigona. But some of the other species that he has described as belonging to this genus, especially P. intermedius and P. dentatus, also appear to be worthy of generic distinction.
In this article, I have constituted several new genera to include some of these peculiar forms, hitherto referred to Pentagonaster, together with some new species.
160 A. EF. Verrili—Revision Genera and Species of Starfishes.
Tosia Gray, emended.
The genus Zosia, as here limited, will include not only the typical group of species named by Gray, in which more or less of the marginal and abactinal plates are naked in the middle, but also those that are granulated over the whole surface, as in granularis and its allies, the extent of the granulation having been found to be variable in many. species. In each section there are species with pedicellarie and others in which they appear to be lacking.
The marginal plates are regular and generally correspond pretty closely in the upper and lower series, except distally ; an odd inter- radial plate sometimes occurs abnormally. Apical plate small.
The abactinal plates of the radial areas are polygonal, most often hexagonal, or roundish, crowded pretty closely together, without distinct, intervening, connecting ossicles and without secondary plates of small size.
The papular pores are usually rather numerous, generally place singly in the angles between the plates of the basal radial areas, and sometimes on the central part of the disk, but not on the triangular interradial areas, where the plates are angular and closely in con- tact. The pedicellarize when present are small, elevated, usually _ with two spatulate blades, higher than broad, and often set in’ special pits of corresponding shape. They may occur on any or all the kinds of plates, either above or below, or on both sides. |
The adambulacral spinules are numerous and crowded, and grade into the actinal granulation ; the furrow-series form a simple row, usually not much longer than those of the next series‘and not separated from them by a wide space. Distally some of the spines — of the second series usually become much longer than the rest.
In the following table we give an arrangement of most of the figured species of Zoséa, in sections and sub-sections.
Section A—Typical. Distribution Indo-Pacific and Australian.
More or less of the marginal and abactinal plates are naked in the center, margined by one or two rows of granules. Adambulacral plates narrow, each with few spinules, usually only two or three in the furrow-series. ‘”
Pedicellariz absent or not recorded in most species. Only one or two of the distal, dorsal, marginal plates are usually in contact medially.
6.—The actinal as well as abactinal and marginal plates are usually naked centrally.
T. australis Gray, Synop., p. 11, pl. 16, fig. 1. West Australia,
T. rubra Gray, Synop., p. 11, pl. 16, fig. 3. Australia.
T. tubercularis Gray, Synop., p. 11, pl. 16, fig. 4. Australia.
A. E. Verrill—Revision Genera and Species of Starfishes. 161
fT. magnifica (M. & Tr.), Ast., p. 53, pl. iv, figs. 1 Tasmania.
T. astrologorum M. & Tr., Ast., p. 54, Australia. A variety has pedicellariz (¢. Perrier).
ame b, 1842.
6b.—The actinal plates are usually entirely covered with granules.
T. aurata Gray, Synop., p. il, pl. 16, fig. 2 (= Astrogonium australis M. & Tr., non Gray, t. Perrier). Australia.
Pedicellariz size of granules on abactinal plates.
’ T. grandis Gray, Synop., p. 11, pl. 3, fig. 1. West Australia.
T. tuberculata (Gray). Port Natal.
4 Section B. Plinthaster.—Pedicellarie with narrow blades are pres- _ ent, of small size, about equal to the granules, or but little larger. _ Adambulacral plates are wider, about as large as the actinal plates, and bear many crowded spinules ; usually four to six in the furrow ‘series. Marginal and abactinal plates usually naked in the middle and often areolated. Three to five of the dorsal marginal plates are usually in contact medially. Atlantic.
Z. Perrieri (Sladen) = P. Perrieri Perrier, 1894, p. 391, pl. 25, figs. la, 10.
Off Morocco, 930 to 1590 meters.
Pedicellariz occur on the abactinal and on both series of marginal plates. They are set in special bilobed pits. Upper marginal and abactinal plates are granulated only around the edges.
2 T. compta Ver. West Indies, 683 fathoms. : T. nitida Ver. West Indies, 335 fathoms. _ Section C. Ceramaster.—All the plates, above and below, are usually granulated nearly or quite all over, unless rubbed ; in some
species the marginal plates may often have a small, naked, central
area. Adambulacral plates with four to six furrow spines. Atlantic. c.—Pedicellarie absent or not recorded. Only one or two dorsal
marginal plates are usually in contact medially.
_ T. granularis (Retz.). Arctic Ocean and both coasts of the North
Atlantic, 20 to 750 fathoms.
| _ The variety Deplasi Per. (1894, p. 401) has some of the marginal
_ plates naked in the middle; the same occurs in some of our examples. T. simplex Ver., 1895, p. 135. Off Martha’s Vineyard, 640 fathoms.
The type of this species has a small naked spot in the middle of
the marginal plates, above and below.
~~ T, ewimia Ver., 1894, p. 264. Off Nova Scotia, 80-122 fathoms. T. Greenei (Bell, 1889); 1892, p. 74, fig. Off Ireland, 1000 fathoms. T. placenta (M. & Tr.). Mediterranean, 40-50 fathoms.
_ T. mirabilis (Per.). Mediterranean.
Trans. Conn. Acap., Vou. X. AvuGust, 1899.
ll
162 A. BE. Verrill—Revision Genera and Species of Starfishes.
T. mammillata (M. & Tr.). Locality unknown. Pedicellariz absent on type (t. Perrier).
cc.—Pedicellarie are present; their blades are higher than broad, — usually spatulate or spoon-shaped. Only two to three pairs of dorsal © marginal plates are in contact medially.
T. Vincenti (Per.), 1894, p. 396, pl. 26, fig. 2. East Atlantic, 946 to 1105 meters. ; In this species there is a regular row of spatulate pedicellariz on
the row of plates next the adambulacral series.
T. hesitans (Per.), 1894, p. 397, pl. 23, fig. 7, pl. 25, fig. 2. Hast Atlantic, 2210 meters.
Pedicellariz small, numerous ; they occur on the actinal, adambu- lacral, abactinal, and on both series of marginal plates. In the type some of the marginal plates are naked in the middle, perhaps acci- dentally rubbed. Three of the dorsal marginal plates are in contact medially.
T. Grenadensis (Per.), 1881, 1884, p. 232, pl. viii, fig. 2. West Indies, 176 fathoms. Pedicellariz few, small, abactinal.
T. Gosselini (Per.), 1894, p. 399, pl. 26, fig. 4.
East Atlantic, 946 to 1440 meters.
Small spatulate pedicellariz, with special pits, occur on the abacti- nal and both series of marginal plates. .
T. pulvinus (Alcock, 1893), India, 1200 fathoms.
Tosia granularis (Retzius).
Asterias granularis Retzius, K. Vet. Akad. Nya. Handl., vol. iv, p. 288, 1788. Abilg., in Zool. Dan., fas. 3, p. 19, pl. xcii, 1788. Bruzelius, Diss. Sys. Ast., p. 10, 1805.
Astrogonium granulare Miiller and Trosch., Syst. Asteriden, p. 57, 1842. Gray, Synop., p. 10, pl. 1, fig. 4, 1866. Verrill, Expl. by the Albatross in 1883, p. 542, pl. 18, figs. 48, 48a, 1885.
Goniaster granularis Liitken, Vidensk. Medd. nat. Foren., p. 146, 1865.
Pentagonaster granularis Perrier, Revis. Stell. du Mus., p. 224, 1876. Sladen, Voy. Challenger, vol. xxx, p. 268, 1889. Bell, Catal. British Echinod. in
3ritish Museum, p. 73, pl. x, figs. 4,5, 6, 1892. Verrill, Distrib. of Echinod., Amer. Journ. Sci., vol. xlix, p. 185, 1895. Danielssenand Koren, Asteroidea, Norske Nordhavs-Expd. Zo6l., xi, p. 58, 1884. ‘
Pentagonaster balteatus Sladen, Proc. Royal Irish Acad., i, p. 688, pl. xxv, 1891 (t. Bell).
Pentagonaster concinnus Sladen, op. cit., i, p. 690, pl. xxvi, 1891 (t. Bell).
A large specimen, from off Halifax, N. S., has the following char- acters :
The inner adambulacral spinules form a simple marginal row, with three or four spines on each plate, of which the proximal is smaller
A, E. Verrili—Revision Genera and Species of Starfishes. 163
and sets farther back, so as to be partly overlapped by the distal one
_ of the preceding plate; the others are rather short, stout, blunt, scarcely tapered, about as long as the breadth of the adambulacral plates. Outside the furrow-series, each plate bears an actinal group of about seven to eleven short, stout, polygonal spinules or granules, one of which occupies the center, and the others surround it ; those on the side next the furrow-series are much larger and somewhat longer than the rest. Oral spinules numerous, short, stout, poly- gonal, seven or eight on the border of the dentary plate, and a median or sutural group consisting of a row of six or eight on each plate, with two shorter intermediate or central rows of three or four smaller ones. —
The actinal interradial plates are crowded, polygonal and closely covered with small polygonal granule-like spinules with rounded
tips, about thirty on the larger plates, their size decreasing toward the marginal plates, where they are very small.
The marginal plates, above and below, are closely covered with similar but smaller granules. The plates of the upper surface are hexagonal on the radial areas of the bases of the rays, and are mostly transversely elongated, and surrounded by six papular pores, corresponding to the angles. In the interradial areas they are trans- versely rhombic, often with the acute angles truncated, where pores intervene. All are closely covered with small angular granules.
Madreporic plate small, with conspicuously convoluted, deep grooves and high ridges. It is nearer to the center than to the margin.
Taken on the American Coast, by the “ Albatross,” at several stations between N. lat. 44° 28’ 30” and 41° 47’. Also taken by the Gloucester fishermen on the Banks off Nova Scotia. Occurs off the coasts of Norway and Great Britain.
Bathymetrical range, 50 to 471 fath. on the American coast, Rarely taken below 150 fathoms.
Z Tosia (Plinthaster) compta Ver., sp. nov.
& PLATE XXVII. Ficwre 2.
& Pentagonal with regularly incurved margins and short, tapering é subacute rays. Radii as 13:8.
* Marginal spines large, mostly nearly square, slightly convex, the _ upper and lower ones nearly corresponding along the margins of the q : disk, but alternating on the distal part of the rays. There are : usually, in the type, 16 upper and 18 lower plates on each side of “$y
164. A. EB. Verrill—Revision Genera and Species of Starfishes.
the body, but on one margin there are two ventral plates corre- sponding to one of the upper dorsals nearest the median line, so that — there appears to be an odd, lower, interradial plate on this side.
The dorsal marginal plates are smooth, microscopically strigillate, and naked except for a single row of small, round marginal granules and a central irregular cluster of large, well spaced, round granules, each implanted in a pit and easily detached. They are lacking on the small distal plates. Three or four of the distal plates are in contact medially. The apical plate is of moderate size, wedge-shaped proximally and prominent at the tip.
The ventral marginal plates have most of the surface covered with — implanted round granules, like those of the upper ones, and distinctly larger than those of the marginal row.
The abactinal plates are flat, even, closely crowded, regularly arranged, and mostly of about the same size, though the median radial rows are easily distinguished. They are mostly rounded or _ hexagonal with rounded angles. They are covered with small hemispherical bosses, but are not granulated, having only a single row of minute grains around the edges. A group of these grains, of somewhat larger size, surrounds each papular pore. The latter are few and small, but easily visible; they are confined to the basal radial areas. The madreporite is small, convex, prominent, with fine gyri. }
The proximal adambulacral plates bear each a strait, regular row of five or six short, blunt, prismatic spinules in the furrow-series. The actinal side bears a second row of about four stouter conical spinules, of about the same diameter, but larger than the actinal granules ; the outer margin bears four to six granules, like those of the actinal plates. Distally the plates have an angular inner edge, with fewer and more slender spinules in an oblique row, while one or two of those in the actinal row become much longer and larger.
The actinal plates are large, mostly rhombic, well defined, and covered with rather coarse, somewhat conical granules, which are not closely crowded.
Pedicellariz, about like the granules in size, with narrow oblong blades, oceur very sparingly on the adambulacral and some of the actinal plates. The dentary plates are large, covered with spaced, conical granules, similar to those of the actinal plates, but larger ; those near the apex become stouter and prismatic, like the apical teeth ; there are 7 or 8 in the furrow-series, similar to the adambu- lacral spinules. “7
A. E, Verrili—Revision Genera and Species of Starfishes. 165
Greater radius, 44™™; lesser, 27™™,
Taken by the U. S. Fish Com. steamer Albatross in the West Indies, at station 2117, in 683 fathoms, and by the Blake, station xi, in 555 fathoms, 1880,
This species is very similar to 7: Perrieri of the East Atlantic. — Without a direct comparison of specimens it is impossible to say _ whether our form may not be merely a variety of the latter. How- ever, the American form differs from the photographic figures of 7. Perriert in having larger marginal plates and in the details of _ the actinal surface. Moreover its pedicellariz are much fewer and apparently are different in form.
- Tosia (Plinthaster) nitida Ver., sp. nov. PLATE XXVII. Ficures 1, la, 1b.
Pentagonal with regular incurved sides; 18 or 19 upper marginal _ plates on each side; 20 lower ones; 4 or 5 upper marginal plates are in contact medially.
Closely allied to the preceding species in form and most of the details of structure. It differs chiefly in the finer granulation and in having the abactinal plates more closely crowded and even, with less evident sutures between them and with the areolation of their surface much finer; the granules around their margins are also much smaller and lacking in many places, but a group of rather larger ones sur- rounds each of the very small, unequal papular pores, so that these _ appear quite distinctly over a limited basal radial area. They are lacking on the central part of the disk and on the large interradial areas,
The dorsal marginal plates are partially naked and smooth, but have a central group of well spaced, rounded, implanted granules, as in 7. compta, but the granules are much smaller; the lower third of these plates is closely covered with small round granules, like those of the ventral plates. The adambulacral plates, on the proximal part of the groove, usually have five or six slender, compressed furrow
the actinal plates. - Pedicellariz of small size, similar to the granules in appearance,
_ Greater radius, 27™™; lesser, 15™™. _ Taken by the Albatross, in the West Indies, at station 2396, in 335
166 A. & Verrill—Revision Genera and Species of Starfishes.
This species is so similar to the last, in most of its characters, that it might prove to be only a variety, if we had a large series for study. But although the type is not much more than half as large — as that of 7. compta, it has rather more marginal plates and more — numerous adambulacral spinules and actinal granules. The reverse — would usually occur in the young of this genus. Hence I am dis-. posed to consider it a very closely related, but distinct, species.
Pyrenaster Ver., gen. nov. Type, P. dentatus Perrier.
Form flat, more or less pentagonal, or stellate with a broad disk. Rays tapered. Marginal plates rather large, those of the two series — similar and generally paired ; sometimes there is on one or more of — the margins (rarely on all) an odd interradial plate,—but this seems to be more or less abnormal. .
In the type the upper marginal plates are partially naked, and the — abactinal plates usually have a small naked central area, surrounded ~ by marginal granules, but this is not constant. The upper marginal - plates in the type species are sometimes all separated by a row of — abactinal plates; in other specimens of the same species two to five pairs are in contact medially. Actinal and inferior marginal plates granulated.
Pedicellariz occur sparingly on the adambulacral plates; they are similar to the granules in size and height and have short chisel-shaped blades,
The dentary plates are large, triangular, with numerous prominent granules on the actinal surface, and with somewhat enlarged pris- matic spinules on the oral margin.
Adambulacral plates large, squarish, with 4 to 7 furrow spinules in a regular marginal series; these are decidedly more elongated than the granules of the actinal surface and are separated from them by anaked space, as in Mediaster. Distally these plates become small, with the furrow end prominent and bearing a convex group of spin- ules, while one or two of the spinules of the second row, on the actinal side, become much longer and larger than the rest, as in Tosia and most of the allied genera.
The actinal plates are flat, rather large, polygonal, crowded and arranged in series parallel with the furrows.
The abactinal plates of the radial areas are rounded, convex and — of two kinds, smaller, secondary, rounded plates being interpolated — between and around the larger or primary plates. The smaller plates are, however, of the same form as the others, and are granu-
—— ie a
A. E. Verrilli— Revision Genera and Species of Starfishes. 167
lated in the same way, but their presence gives an appearance of irregularity to the arrangement of the plates. «
Papular pores are of moderate size and not very numerous; they are confined to the median radial areas. In young specimens these areas are small and well defined and the pores few. Each pore seems to be surrounded by a special group of granules.
This genus is distinguished from TYosia and Pentagonaster espe- cially by the existence of smaller’ secondary, rounded plates between the primary abactinal plates, and also by the greater specialization of the furrow-series of adambulacral spines, for these do not grade into the actinal granulation, as they do in the genera referred to. In this respect this genus is more nearly like Mediaster, but the latter does not have the secondary abactinal plates, but has concealed, radiating connecting ossicles between the distinctly separated abac-
tinal plates. Peltaster also has secondary abactinal ossicles, but
they are different in character, and it also differs in having broad valvular pedicellariz and graded adambulacral spinules.
Pyrenaster dentatus (Perrier) Ver.
Pentagonaster dentatus Perrier, Nouv. Arch. du Mus., vi, p. 242, pl. viii, fig. 3,
1884. Sladen, op. cit., pp. 265, 744, 1887.
Puate XXVII. Ficures 3, 3a, 3b.
I have had an opportunity to examine Perrier’s types of this species, in the Mus. of Comp. Zodlogy, and to compare them with those dredged by the “ Albatross.”
Among the latter there are both large and smallspecimens. They show remarkable variations in several respects.
Two large examples of the same size, Nos. 10,370 and 18,433, are of special interest. In the former, four or five distal pairs of dorsal marginal plates are in contact medially. In No. 18,433, which is closely similar in other respects, all the plates, or all but the last pair, are separated by abactinal plates. In this specimen, on one margin, two upper plates correspond to one lower, so that there is an odd median plate above. In some of the young specimens one or more odd, interradial marginal plates may occur both above and below.
~*~ This species was taken by the Blake Exp., in 41 to 1500 fathoms.
By the Albatross it was dredged in several localities, in the West _ Indies and off the Carolina coasts, in 478 to 1639 fathoms.
168 A. B. Verrill—Revision Genera and Species of Starfishes.
Pyrenaster affinis (Perrier) Ver.
Pentagonaster affinis Perrier, Nouy. Arch. du Mus., p. 248, pl. viii, fig. 4. 1881. Sladen, op. cit., pp. 265, 744, 1889.
This is, perhaps, only a variety of the last. The coarser granula- — tion and the differentiation of the granules around the margins of the abactinal plates, in the papular areas, are the special characters cited. by Perrier. J have not seen the type.
Some of the younger examples, which I refer to this species on account of the last peculiarity, are not in other respects distinguish-— able from dentatus.
It was dredged by the “Blake” in 1131 and 1323 fathoms, in the West Indies, and by the Albatross.
Peltaster Ver., gen. nov. Type, P. hebes Verrill.
Form nearly pentagonal, with very short, obtuse rays. Marginal . plates rather large, regular, decreasing in size distally, covered like all the other plates, above and below, with fine nearly uniform gran- ules. Apical plate small.
Abactinal plates numerous, not large, closely crowded, of two kinds. ‘The primary plates are mostly hexagonal. Between, and often surrounding them, are smaller roundish or irregular plates granulated like the larger ones, but with fewer granules. Papular pores small, numerous, arranged singly around the pee y plates and occupying large radial areas.
Pedicellariz, in the type, large, bivalve, sessile, with broad, lamel- liform jaws, as wide as half the diameter of a plate. They occur mostly on the actinal plates next the adambulacral series. In P. planus wone have been observed, but only one specimen is known. Adambulacral plates large, with several series of spinules, which are short, crowded, prismatic and grade into the granulation of adjacent plates. The furrow-series form regular rows of four to six on each plate ; they are smaller and not longer than those of the next series, — and there is no naked space between the series.
Distally one or two of those in the second series gradually change to much longer and larger blunt or conical spines. Dentary plates not prominent, covered with numerous blunt prismatic spinules, like those of the adambulacral plates, but rather coarser.
Actinal plates numerous, squarish or rhombic, closely crowded, the outlines obscured by their close, uniform granulation, They run in series parallel with the adambulacral furrows.
A. FE, Verrili—f—Revision Genera and Species of Starfishes. 169
This genus is separated from TZosia on account of the small, irregular, secondary plates or ossicles between the primary abactinal plates, and the large, broad bivalve pedicellariw of the type.
The characters of the marginal plates, actinal plates, and of the adambulacral spinules are like those of Zosia, of the granularis group (section C).
The second species (planus), although, so far as known, without pedicellariz, is placed in this genus because it agrees with the type in the characters of the skeleton.
Peltaster hebes Ver., sp. nov. Puate XXVIU. Ficure 4.
Form broadly pentagonal, with very short rays and a rather thick, flat disk and large, slightly convex marginal plates, decidedly higher than long. Radii as 7:8. All the plates are closely and uniformly granulated, above and below, and many actinal plate shave, in the type, central, large, bivalve pedicellarie with broad blades. Upper marginal plates about 20 on each side of the body ; lower ones about 24 in the type. Along the sides of the disk the upper and lower ones are pretty closely paired and nearly of the same size and shape, though the vertical sutures are not strictly coincident, except between the middle plates, owing to the slightly wider lower plates. In each series the plates are nearly twice as high as long, and this form holds good except for the last two upper and last four or five lower plates, which decrease in size and change form very rapidly, the last ones being very small. The apical plate is very small, obconic, not prominent. The abactinal plates are closely crowded, and so closely granulated that the outlines are concealed, unless denuded. The primary plates are rounded or polygonal, with many rounded angles, and are sur- rounded, in the radial areas, by many smaller secondary plates, having the same form and granulation, but variable in size, and mostly less than half the diameter of the larger plates. All are closely covered with small round granules, the marginal series scarcely different from the rest. The larger plates may have 40 to 50 granules, of which 18 to 24 may form the marginal row.
The papular pores are very small and numerous, placed singly, and occupy wide radial areas.
The adambulacral plates bear a closely crowded group of graded spines on the actinal side ; the furrow-series consists of five or six short, thick, blunt, prismatic or compressed spinules, in a nearly
170 A. E. Verrill— Revision Genera and Species of Starfishes.
straight row ; next, and close to these, there is a row of three or four larger, angular blunt spinules of the same height; these are followed — by another row of three or four similar but small spines, in a slightly curved row; then there is a group of five or six, sometimes forming rows, on the outer part of this plate, of the same form and size as the actinal granules. .
The actinal plates are numerous and even, closely crowded, mostly rhombic or squarish, covered with granules that become angular where most crowded.
Large valvular pedicellariz occupy the center of many of the plates in the series next the adambulacral ; they are about as broad as half the diameter of the plate, or more.
The dentary plates are not prominent, but are covered with numerous prismatic granules and spinules, larger than those of the adambulacral plates.
The madreporic plate is large, round, with numerous fine gyri. The dorsal nephridial pore is surrounded with granules larger then those of the surrounding plates.
Greater radius, 56™™; lesser, 50™™.
Taken by the Albatross, in the West Indies, at station 2668, in 294 fathoms, gray sand.
Peltaster planus Verrill.
Pentagonaster planus Verrill, Distr. Echinod., Amer. Journ. Sci., xlix, p. 135, 1885. , Puate XXVIII. Ficures 3, 3a. 5
Form pentagonal, with the sides only slightly incurved ; rays very short, triangular, and obtuse, with the tip turned up and terminated by a small, conical plate.
Marginal plates large, median ones nearly square, higher than long, — the upper and lower nearly corresponding, fourteen in the dorsal series and sixteen in the ventral series, all uniformly covered with rather coarse, rounded granules, standing a little apart; the margins of the plates with a regular row of granules of about the same size. The three distal dorsal plates are in contact medially. Apical plate small, obovate. :
Abactinal plates nearly flat, the primary ones rather large, rounded or hexagonal with rounded angles, with many small, rounded, unequal secondary ones interspersed ; all are uniformly covered with rather coarse, spaced granules, like those of the marginal plates, so that the whole of the upper surface has a remarkably uniform granular coat-
A. FE. Verrill—Revision Genera and Species of Starfishes. \71
ing. The larger plates often bear fifty to seventy granules; the small intermediate plates frequently carry but nine to twelve, one or two being central. Actinal plates large, rhombic, uniformly covered with coarse, angular granules, distinctly larger than those of the marginal plates.
Adambulacral plates numerous and crowded, similar to the actinal plates, but slightly larger and longer; toward the ends of the rays the plates are smaller and one or two of the first actinal row of spinules become much larger and longer, round and blunt. Each plate usually bears three or four marginal spines in a simple row ; outside of these there are usually nine to twelve thicker, obtuse, angular spines, forming four irregular longitudinal rows, the outer ones smallest and like the actinal granules.
Dentary plates not prominent, covered with numerous blunt, angu- lar spinules, similar to the actinal spinules, but larger.
The papular pores are numerous, placed singly, and occupy large radial areas, extending nearly to the center of the disk.
No pedicellariz could be found.
Greater radius of the type, 50™™; lesser radius, 35"™; thickness at margin, 8™™,
N. lat. 39° 53’, off Martha’s Vineyard, in 156 fathoms, one speci- men (No. 13,362).
Litonotaster Ver., gen. nov. Type, P. intermedius Per.
Stellate, with a rather broad, flat, flexible disk and tapered rays, becoming slender distally. The dorsal integument is so thin that it is wrinkled in the dried specimens. Marginal plates unusually small for this family. The dorsal ones encroach but little on the upper surface of the disk ; distally they become irregular near the tip of the rays, in the type; two to four pairs are in contact medially (a single oblong plate, equal to two or three of the usual distal plates, may replace the latter on some of the rays).
Abactinal plates polygonal, flat, thin, closely united, finely granu- lated, with two or more rows of granules around the edges, but with a small, central, round, naked area, in the type.
Papular pores rudimentary, few, small, obscure, not visible except when the plates are denuded ; they occur only between the three central rows of plates, in a very circumscribed basal radial area.
Actinal plates granulated, rather large, angular, of various forms, not forming regular rows.
172. A, E. Verrill—Revision Genera and Species of Starfishes.
Adambulacral plates are large, as wide as the adjacent actinal plates or wider. Each one bears seven or eight small, compressed furrow spines, in a regular row; the spinules of the actinal side are very small, on the proximal plates, and form an irregular group on the outer half or else stand more or less in three or four rows ; most of them are scarcely larger than granules; distally one or two of the second row become much larger conical spines.
A small elongated pedicellaria, with two, three, or four spatulaté blades, occurs on the center of many of the adambulacral plates and on some of the actinal plates.
The dentary plates are large, separated by am open suture ; acl one bears an actinal triangular group of numerous small granules and a furrow-series of about ten or twelve small, prismatic, blunt spinules, those toward the apex becoming larger.
This genus is separated from its allies mainly on account of the few and minute papular pores and the very limited area on which they occur; the thin and small marginal plates; flexible dorsal sur- face of the disk; and large number of’ adambulacral spines.
The type is the only species determined. Mr. Alcock has recorded this species from the East Indies. Possibly this may indicate a second species of the genus.
Litonotaster intermedius (Perrier). Pentagonaster intermedius Perrier, Etoiles de Mer., p. 243, pl. v, figs. 5, 6, 1884. Sladen, Voy. Chall., xxx, p. 746, 1889.
» .
Puate XXVIII. Ficures 5, 5a, 5b.
This species was taken by the Blake Expedition in the West Indies, in 1930 fathoms. It was also taken by the Albatross at station 2379, in 1467 - fathoms, yellow ooze (two examples, No. 18,424). I have compared the type described. by Perrier, from the Blake Expedition, now in the Museum of Comp. Zodl., with those taken by the Albatross. They agree closely. The larger Albatross specimen has the radii 33™™ and 14™™,
Eugoniaster, gen. noy. Type, £. investigatoris (Alcock).
Form broadly pentagonal, with short rays. Abactinal plates uni- formly small and rounded, naked, except for a marginal series of granules ; some of them haa een: bivalve pedicellariz. Papular pores numerous, placed singly, radial. Marginal plates mostly naked with a border of granules and also some in a central group.
a
A. E. Verrill— Revision Genera and Species of Starfishes. 173
Bivalve pedicellariz, with wide blades, occur on some of them, as well as on the adambulacral and actinal plates.
Adambulacral plates are covered with actinal granules in longi- tudinal rows; there are six or more prismatic spinules in a regular furrow-series. Actinal plates are granulated and extend to near end of rays.
This genus is related to Peltaster, but differs in having the abac- tinal plates all small and similar, and also naked centrally, and in having the marginal plates mostly naked, except around the margin. The large bivalve pedicellariz are similar in the two genera. The character of the pedicellarie differentiates the genus from 7osia and its closer allies.
Eugoniaster investigatoris (Alcock). Pentagonaster investigatoris Alcock, Ann, and Mag. Nat. Hist., xi, p. 88, 1893.
This large species has, on the abactinal surface, “ uniformly small round tabular plates, which are distinctly isolated from one another and are fringed with a single row of flat squamous, membrane-clad granules flush with the general surface, but are otherwise naked, except that some of the plates (perhaps one-fourth) bear a very excentric or quite marginal, broadly bilobed pedicellaria.”
The marginal plates are also bordered with squamous granules and bear bilobed pedicellariz ; some also have a central group of granules.
Bivalve pedicellariz also occur on the adambulacral plates and broad ones on the actinal plates near the jaws.
Adambulacral plates bear crowded, graded actinal granules in two or three longitudinal rows, and a furrow-series of six or seven prism- shaped spinules.
Antheniaster, gen. nov. Type, A. sarissa (Alcock, sp.).
This genus resembles Anthengides Per. in having a thin, finely granulous membrane over the abactinal surface of the plates, but it differs so much in other respects that it cannot properly be referred to the same genus. The pedicellariz are papilliform or spoon-
‘shaped; not large and bivalvular as in the latter.
s
<4
It has two kinds of abactinal plates, which is not the case in Anthenoides. The larger plates are “stellate or somewhat poly- onal,” arranged in radial rows; the small secondary plates are
_ “inlaid everywhere between the large plates.” Papular pores exist
on large radial areas. Marginal plates are large and partly granu- lated; the dorsal distal plates are in contact medially and mostly naked,
174. A. EL Verrill—Revision Genera and Species of Starfishes.
Pedicellarix of a simple papilliform structure occur on some of the
upper plates. The lower marginal plates have two or three spines
in a horizontal row; one distally.
The adambulacral plates have a divergent or palmate-series of —
furrow spines and a larger spine on the outer actinal end; many have also a central pedicellaria with spoon-shaped blades.
The actinal plates are numerous, in cheyvrons, and extend to about 7
the 13th or 14th adambulacral plates; they are covered with a granulose membrane, and some bear papilliform pedicellaria. The dentary plates are very prominent and bear large granules actinally,
but the oral spines are large. The ambulacral feet have a terminal —
sucker. Antheniaster sarissa (Alcock). Anthenoides sarissa Alcock, Ann. Mag. Nat. Hist., xi, p. 99, 1893.
Andaman Sea, 139 to 250 fathoms.
Subfamily HIPPASTERIIN&, nov.
This group is established for those Goniasteridx that have large elongated, divergent, and differentiated adambulacral spines, one or
two larger ones situated on the central part of the plate. ‘The dorsal —
and marginal plates are bordered or covered with large granules and often have one or more central tubercles or stout spines.
Bivalve pedicellariz, often of large size, are usually present. The
abactinal plates are thick, closely joined, and polygonal or roundish.
Hippasteria Caribzea Ver., sp. nov.
Puate XXVIII. Ficures 1, la.
Form stellate with a rather broad disk and tapered acute rays; disk convex. Radii about as 2:1.
Marginal plates regularly paired, those of the interradial margin nearly square ; all are bordered with coarse rounded granules, and some granules are scattered on the central parts; in some cases these form a central cluster on the lower plates. Most of the upper plates are naked centrally, and rise into a low conical tubercle, often sur- mounted by a small, round, ovate, blunt spine or large granule.
Many of the lower plates have a central large bivalve pedicellaria”
with low, broad blades; their breadth is about half the width of ye plates,
A. E. Verrill—Revision Genera and Species of Starfishes. 175
The abactinal plates are rounded, with a marginal row of coarse round granules; the center is occupied, in most cases, by a broad, low, bivalve pedicellaria, nearly as wide as the plate. Each plate of the radial areas and of the center of the disk has five or six papu- lar pores around it. The apical plate is irregularly ovate, with a pair of small apical spines.
The actinal plates are not numerous, much crowded, and closely united, so that their outlines are obscure. They have marginal granules and a large central pedicellaria, like those of the actinal plates, but rather larger.
The adambulacral plates are narrower; each has two, or some- times three, flattened, blunt or spatulate, often crooked furrow spines; a larger clavate spine in the next row, standing on the cen- ter of the plate; and three to five much smaller, unequal, conical or
clavate spinules in a group on the actinal end.
The oral spinules are numerous, crowded, and much compressed.
Greater radius, 17.5™™; lesser, 8.5 to 9.5™™,
Taken by the Albatross at station 2668, N. lat. 30° 58’ 30”, W. long. 79° 38' 30”, in 268 fathoms, gray sand (No. 18,425, one young).
The discovery of this tropical species is of special interest, for the genus was previously represented by only two species; one (Z. phrygiana) found in the boreal parts of the North Atlantic, on both coasts, extending on the American coast to Cape Cod in moderately deep water ; the other (HZ. magellanica) from the region of Pata- gonia. The occurrence of the genus in the intermediate tropical region is, therefore, significant.
Cladaster Ver., gen. nov.
Stellate, with a broad, flat disk; interradial margins regularly in- curved; rays tapered.
Marginal plates of both series rather large, not numerous, en- croaching upon both sides of the disk, regularly paired, except dis- tally, decreasing regularly in size ; about four pairs of the dorsal ones are in contact medially. No odd interradial plates. Apical plate and those adjacent, small.
The marginal plates and all the actinal and abactinal plates are normally granulated, but in the type many of the marginal and abactinal plates have irregular, partially naked central patches, covered with small pits where granules have fallen off.
Abactinal plates all polygonal with rounded angles, rather large, not numerous; the median row of the rays is distinct, and bordered
176 A. EB, Verrill— Revision Genera and Species of Starfishes.
on each side by a regular row of about the same size and form, arranged alternatingly.
Papular pores are arranged singly around and between the three central radial rows of plates, except distally, usually six to a plate, but are absent from the small interradial areas.
Actinal plates are few, rather large, angular, rather irregular and do not extend beyond the second pair of marginal plates, in the type; they are covered with well spaced, coarse granules. Pedicel- larize with two elevated spatulate blades occur on the middle of some of the actinal plates,
Adambulaeral plates bear relatively large and long, prominent, interlocking, spatulate or club-shaped spinules. Two of these, on each plate, belong to the furrow-series and are much flattened. Out- side of these, there is a stouter median spine, of the same length, usually not much flattened, clavate or blunt at the tip; outside of this there are usually two smaller, conical spinules, on the actinal - margin. The larger spines of the first actinal row do not increase in size distally, as in Zosia, etc., but gradually decrease.
The dentary plates are rather large, flat, and bear marginal and sutural rows of elevated, flattened or spatulate spines, like those of the furrow-series.
This genus seems to be more nearly allied to Hippasteri ta than to any other.
Cladaster rudis Ver., sp. nov.
Puate XXVIII. Ficures 2, 2a, 2b, 2c.
Rays narrow distally with four dorsal marginal pairs of plates in contact medially ; these aca plates are small and not regularly paired. Radii about as 2:1; greater radius, 25"; lesser, 12™™.
Dorsal marginal plates vary from 13. to 15 on different sides of the body. Ventral plates 15. Those of both series are similar in form and size, thick, somewhat convex, rectangular, higher than long on the interradial margins, and encroach considerably upon both sides of the disk; they are separated by deep sutures, Four larger pairs form the margins of the disk; those on the rays become rapidly smaller and more square. Those of both series are sparsely covered with coarse, rounded, well-spaced granules, many of which have fallen off, leaving small, shallow, rounded pits on the central portions of some of the plates. The marginal granules are of the
A, E. Verrilli— Revision Genera and Species of Starfishes. 177
same form, but rather smaller and more closely arranged. The granules are higher than broad with a rounded top.
Abactinal plates are covered with granules like those of the mar- ginal plates. They are roundish and slightly convex, in contact by their angles, between which some of them are slightly notched or incurved, to make room for the papular pores.
The madreporic plate is rather small, with many fine gyri.
The granules of the actinal plates are well-spaced and rather larger and more conical than those of the upper side; they form a marginal series, mostly of six to ten, which are somewhat divergent, and surround one or two larger central ones, which are sometimes replaced by a central pedicellaria, having broadly spatulate blades, rather higher than the granules.
The two outer spinules or granules of the adambulacral plates are like those of the actinal plates; there is often a minute conical spinule on the proximal side of the larger central spine.
Taken by the Albatross, off Florida, at station 1415, N. lat. 30° 44', in 440 fathoms, coarse sand, shells and foraminifera (No. 18,426, one example).
Subfamily MEDIASTERINA,, nov.
This subfamily is proposed for those Goniasteridx that agree very closely with the typical Goniasterine in the structure of the actinal side and marginal plates, but have paxilliform abactinal radial plates in the papular areas. These plates may be in the form of protopaxille or parapaxille, but are usually covered by close gran- ules and not by spinules, but sometimes they are spinulose. They stand a little apart, when denuded, and may appear stellate at base, Mediaster and Nymphaster are the leading genera. In these there are bivalve pediceilariz.
In Mediaster, and probably other genera, these plates are united by small dermal radiating ossicles that do not show distinctly at the surface (see pl. xxv, figure 8). In Mymphaster there are no connect- ing ossicles, but the columnar plates have enlarged and six-lobed bases. Some of the forms placed here show strong affinities to some
Pseudarchasterine. e
Trans. Conn. Acap., Vou. X. Aveust, 1899. 12
178 A. BE Verrili—Revision Genera and Species of Starfishes.
Mediaster Stimpson. Mediaster Stimpson, Journ. Boston Soc. Nat. Hist., vol. vi, p. 490, pl. 23, figs. 7-11, 1857. Mediaster Sladen, Voy. Challenger, Zool., vol. xxx, pp. 263, 752, 1889. Tsaster Verrill, Proc. U. S. Nat. Mus., vol. xvii, p. 257, 1894.
The original description and figure of this genus and of the type species by Stimpson were incomplete and rather imperfect, so that the genus has not been well understood by most subsequent writers who have referred to it. Ihave, therefore, thought it desirable to rede- scribe and figure the type species at this time.
Form stellate, with a broad flat disk and moderately long tapered rays. Marginal plates well developed, not swollen, granulated, rather numerous, higher than broad, paired, upper and lower series nearly equal in size and number and with their sutures more or less closely corresponding vertically; oblique in the type. No odd interradial plate. Abactinal plates or parapaxille are regularly . longitudinally arranged, of moderate size, somewhat elevated, mostly roundish, covered with a rosette of short, obtuse spinules or granules. When these are removed the plates on the central part of the disk and along the median region of the arms appear as roundish or oval convex bosses. They are connected together by five or six internal radiating ossicles, between which are the pores for the papule.* The papule may be single, or (in the type) clustered. Thus the plates appear to be stellate at the base, though they are not actually of that shape. The median row of abactinal plates extends to the apical plate of the rays in the type, but not in some of the other species. Some of the abactinal plates bear a central, broad, sessile, valvular pedicellaria, which, in the type species, is nearly as wide as the plate. They are sometimes lacking.
The adambulacral plates bear a regular marginal row of three to -
seven slender spinules, and usually two exterior longitudinal groups or rows of shorter spinules, which may be angular and obtuse, and toward the tips of the rays, some of them, in the type, become larger and longer, as in Pentagonaster of authors. Some of these spinules may be replaced by spinuliform or clavate, two or three-bladed pedi- cellariex, The actinal disk-plates are angular, often rhombic, closely arranged in rows parallel with the ambulacral grooves, covered with a rosette of granules, the central granules often replaced by a wide valvular pedicellaria. The dentary plates are not very prominent ;
*T have found these ossicles in M. equalis and M. Bairdii. Other species have not been examined as to this feature.
A. EF. Verrill—Revision Genera and Species of Starfishes. 179
each has an actinal row of larger spinules, similar to those of the oral margin.
This genus is closely allied to Pentagonaster, as limited by Sladen and some other recent writers. The principal differences consist in the somewhat more elevated and convex abactinal plates, especially in the papular areas, where they are more widely separated by the large papular pores and united by intervening small internal ossicles, which give them a stellate appearance. On other parts of the disk, as near the interradial margins, the plates are angular and closely joined in a mosaic, as in the former genus. The large valvular pedicellariz are also, to some extent, characteristic, but the marginal, actinal, and dentary plates and their spinules are essentially the same in the two genera. The spinules on the adambulacral plates are, however, more definitely triseriate, and the furrow series is more differentiated in all the known species, though this is perhaps of no more than specific value.
The type inhabits the Pacific coast of North America, in rather shallow water. No other species seems to have been described until a second one, from deep water off our north-eastern coast, was described by me in 1882, under the name of Jsaster Bairdii. Its close affinity to Mediaster was not recognized at that time, though I have referred it to that genus, for several years, in my MSS. lists and in the museum catalogues.
Mediaster sequalis Stimp. Journ. Boston Soc. Nat. Hist., vol. vi, p. 490, pl. 23, figs. 7-11, 1857.
Prats XXIV. Ficurss 10, 11, 12.
Rays five, in length about equal to the diameter of disk, regularly tapered, slender at the tip. Radii nearly as 1:3. Marginal plates on each side of a ray 22, above and below, in a specimen having the greater radius 36™°. The plates on the margin of the disk are higher than wide with the intervening sutures somewhat oblique. The lower marginal plates are similar in size and shape. All are closely covered with small rounded granules. Abactinal areas of the rays are wide at the base, where they may consist of seven or nine rows of plates, but they rapidly decréase to three rows, and only the
@ median row reaches the apical plate. The papular areas are large, | covering nearly the whole width of the proximal half of the rays, as well as most of the central disk. In these areas the plates are rounded or elliptical, convex, somewhat elevated, and separated by
180 A. E. Verrill—Revision Genera and Species of Starfishes.
intervening spaces, in which there are usually five or six groups of papular pores, the individual pores being small and unequal, two or three or more forming each group.
Each of the larger abactinal radial plates is covered with a rosette consisting of about five to seven central, and twelve to fourteen marginal, short, blunt or clavate, granule-like spinules, rather longer than broad. Some of the disk-plates are larger with more spinules. A large valvular pedicellaria often replaces the central group of spinules on some of the plates. These occupy nearly the whole breadth of the central area of the plate, and are narrowly oblong, not much elevated, with a nearly even and straight margin. Similar pedicellariz, as well as some much narrower ones, occupy the central area of some of the actinal disk-plates.
The madreporic plate is small, sunken, with narrow, acute gyri. The central nephridial pore is small but distinct.
The actinal disk-plates are crowded and closely united; those_ next the adambulacral plates are squarish or rhombic and form regular rows, but those in the angles are smaller, irregular, and more rounded. All are covered with rosettes of granules, or short, obtuse, often prismatic spinules, rather larger and less regular than those of the upper side. A central valvular pedicellaria occurs on some of the plates, as stated above.
The adambulacral plates are squarish, not very large. Each bears _ a marginal row of three or four, small, oblong, more or less prismatic or compressed, blunt spinules, the middle one usually a little larger than the others. External to these are two sets of shorter spinules, about three in each series ; these sometimes form two rows, but in other cases are in a rosette-like group; those next the inner or groove-series are longer than the others; one or more of these, especially distally, may be replaced by a spinuliform pedicellaria with two or three blades. On the distal part of the ray one or two of the spinules on the central part of these plates becomes considera- bly longer and larger than the rest. The oral spinules are similar to the adambulacral, but those at the tip of the oral plates are rather larger and more angular. The apical plates are rather small, promi- nent, somewhat obovate.
Radius of disk, 13™™; of rays, 36™™.
Off Wilmington, Cal., 27 fathoms, U. S. Nat. Mus.
A, E. Verrili— Revision Genera and Species of Starfishes. 181
Mediaster Bairdii Verrill.
Archaster Bairdii Verrill, Amer. Journal Sci., vol. xxiii, p. 139, 1882.
Isaster Bairdii Verrill, Proc. U. S. Nat. Mus., vol. xvii, p. 258, 1894. Amer. Journal Sci., vol. xlix, p. 156, 1895.
Mediaster steilatus Perrier, Mem. Soc. Zool. de France, iv, p. 268, 1891. Re- sults des‘Campag. Scient., fas. xi, p. 46, 1896, pl. iv, figs. 1-1%.
Puate XXIV. Ficures 1-9. Puate XXVI. Ficures 8, 8a.
A comparison of this species with the type-species of Stimpson has convinced me that they are very closely allied and should be referred to the same genus, though the Atlantic species is often nearly or quite destitute of pedicellariz. But when pedicellariz do occur they have nearly the same valvular forms seen in those of JZ equalis, though they are narrower and more elevated.
Mediaster Agassizii, sp. nov.
Five-rayed; regularly stellate, with a large disk and rather long tapered rays. Radii nearly as 1:3. Interradial angles are broadly curved.
Marginal plates large, nearly square, slightly convex, but not swollen; 36 dorsal and 38 ventral ones in the type, on each of the five sides; the transverse sutures between those of the upper and lower series are narrow and shallow and usually do not coincide.
The upper plates are sparsely granulated centrally, having only a few rather distant, rounded granules ; their margins are surrounded with a close row of angular granules, but these do not form distinct fascioles. Some of the upper marginal plates have also small valvu- lar pedicellariz.
The lower marginal plates are coarsely granulated over the whole surface, the granules being larger than those of the upper ones ; most of them also have one to three or more, oblong, valvular pedi- cellariz, larger than those of the upper plates.
The abactinal plates are regularly arranged in radial series, very unequal in size, mostly roundish in outline, naked in the middle, but with a marginal row of coarse angular granules.
Many of them have a central, large, oblong, valvular pedicellaria, sunken in a pit; on the larger plates the pedicellaria is about one-half the diameter of the plate, but on the smaller plates it often occupies nearly the entire breadth of the top. Some of the plates lack the pedicellaria and have a central granule in its place. The valves of these pedicellarie are usually higher than broad, with the blade broadly spatulate distally.
182 A. FE, Verrili— Revision Genera and Species of Starfishes.
The abactinal plates become suddenly reduced to three radial rows, about opposite the fourth or fifth pairs of marginal plates, and a little farther out only the median row remains ; this disappears about opposite the ninth pair of plates, beyond which the nine distal pairs of marginal plates are in contact medially. The apical plate is rather small, subconical, prominent. The papular pores are small — and scattered singly over large baso-radial areas.
The actinal interradial plates are angular and polygonal, rather large, closely crowded together, forming rows parallel with the ambulacra; they are covered with large, crowded, rounded granules; most of them have, also, a single, large, oblong or elliptical, valvular pedicellaria, usually occupying about one-half the width of the plate.
The adambulacral plates have each four or five, short, stout, blunt, angular or prismatic spines in the furrow-series, placed in a regular row; next to these, on the actinal surface, there is also a row of three similar, but shorter, compressed spinules; on the outer end there are - one or two rows of smaller and shorter, thick spinules; a large val- vular pedicellaria usually occupies the center of the plate, but when it is absent there is a central row of spinules, making four rows on the actinal surface.
The dentary plates are large ; each one, of a pair, bears about five stout, blunt, prismatic or compressed spinules in the furrow series, and two rows of short, thick, obtuse spinules on the actinal surface, those next the mouth being largest ; at the apex of the jaw there are two larger, thick, blunt, prismatic and compressed spines.
Lesser radius, 25™™; greater radius, 75™. Taken by the Blake Expedition, in the West Indies. ’
This fine species appears to be closely allied to MM. pedicellaris. It is referred to the genus Mediaster with some doubt, for the char- acter of the abactinal skeletal plates could not be satisfactorily ascertained by an external study of the single alcoholic specimen.
Mediaster (?) pedicellaris Verrill.
Goniodiscus pedicellaris Perrier, Nouv. Arch. du Mus., vi, p. 245, pl. iv, fig. 3, 1884. Sladen, op. cit., p. 756, 1889. ’
The following notes were made upon one of the original types of Perrier, in the Museum of Comp. Zodlogy.
Radii as 7:19. Dorsal or abactinal plates, large, roundish, the summit convex when naked, but flat when covered with the spinules; the largest have about sixteen marginal, tapered, acute spinules, and one to five or more somewhat larger acute central ones. Intervening
eee.
bs”
A, FE. Verrill—Revision Genera and Species of Starfishes. 183
papular pores large, single, about six around each plate, except that there are none between those plates in the middle radial rows ; arow on each outer border of the abactinal space extends nearly to the end of the rays, or to within about ten marginal plates of the end, and as far as the rows of lateral plates extend.
The median series of plates extends about four or five plates farther than the lateral, but ceases within four or five plates of the tip; from thence the marginal plates are in contact.
Upper marginal plates bevelled and covered with small, sharp, spaced spinules; the upper spinules are shorter than the lower ones, larger, stouter, acute, divergent; those around the margins are similar and do not form regular fascioles.
Lower marginal plates large, roundish, with one or two marginal series of sharp, divergent, stout spinules, and a central larger one.
Sometimes there are three to five central spinules on the dorsal plates
and on the row of plates next to the adoral plates. Pedicellariz small, narrow, ‘elevated, spatulate in form and rather numerous on the dorsal side. Lesser radius, 18™™"; greater, 59™™.
Station 295, Blake Exped., 180 fathoms. This species was also taken by the A/batross in the West Indies.
Mediaster arcuatus (Sladen).
Pentagonaster arcuatus Sladen, Voy. Chall., Zool., vol. xxx, p. 277, pl. xviii, figs. 5,6; pl. lii, figs. 1, 2, 1889.
This species has a few small pedicellariz on the abactinal plates, similar in size to the granules. South of Yeddo, Japan, 345 fathoms.
Mediaster Japonicus (Sladen).
Pentagonaster Japonicus Sladen, op. cit., p. 272, pl. xlvi, figs. 1, 2; pl. xlix, figs. 1, 2, 1889.
This species has rather large, sessile, bivalve pedicellarie with broad valves; on the adambulacral plates; others of smaller size occur on many of the actinal plates. Some of the pedicellarie have three valves. ~ South of Yeddo, Japan, with the last.
184. A, E&. Verrill—Revision Genera and Species of Starfishes.
Mediaster Patagonicus (Sladen).
Pentagonaster Patagonicus Sladen, op. cit., p. 269, pl. xlvi, figs. 1, 2; pl. xlix, figs. 3, 4, 1889.
This species has rather small, sessile, somewhat elevated pedicella- riz, sparingly scattered on the abactinal and superior marginal plates ; their blades are usually chisel-shaped or spatulate, and variable. Simi- lar ones occur sparingly on the adambulacral plates. Larger ones, with broader valves, occur on the actinal plates ; some of them have three or four blades. The dorsal marginal plates and some of the ventral ones have a small central naked area.
Near Atlantic entrance to Straits of Magellan, 55 fathoms ; of | entrance to Smyth Channel, 245 fathoms.
The Mediaster roseus (Alcock, 1893, p. 98), from India, 740 fathoms, is not a true Wediaster. It appears to belong to Pseudar- chaster and resembles P. granuliferus V.
Nymphaster Sladen.
y
Nymphaster Sladen, Narrative Chall. Exp., i, p. 612, 1885. Voy. Chall., vol. xxx, p. 294, 1889. :
Pentagonaster (pars) Perrier, Etoiles de Mer, p. 283, 1884.
Dorigona Perrier, Exp. Trav. et Talism., p. 365, 1894 (not of Gray, 1866, p. 7, nor of Perrier, 1876, p. 44.)
PEATE XGOVi.. Rigor) 7,
This genus is closely allied to Wediaster. It differs chiefly in hav- ing, in the typical species, the dorsal marginal plates in contact medially for the greater part of the length of the rays. The charac- ter of the pedicellariw, adambulacral plates and spines, jaws, and marginal plates is essentially the same in both, though the pedicel- lariz are usually higher and spatulate in this genus. f
The abactinal radial paxille, in the papular areas, differ in strue- ture from those of Mediaster. In WN. ternalis these plates, when separated, have no basal connecting ossicles, so characteristic of the latter. They are short, thick, columnar, with the basal portion somewhat swollen and slightly six-lobed ; they articulate by means of the lobes, while the papular pores are situated in the spaces corresponding to the emarginations. The lobes are so slight that they can hardly be called stellate. The stellate appearance, as seen — from the exterior, is due to the radial connecting bands of soft tissues between the pores. These plates are less stellate at base than those of Pseudarchaster, and rather more so than those of Plutonaster.
.
A. E. Verriil— Revision Genera and Species of Starfishes. 185
The name Dorigona, used for this genus by Perrier, is untenable. The type of Gray (1866) was D. Reevesii= Ogmaster capella, and the only other species mentioned by him was longimana (Mobius). The genus, as understood by Gray, was synonymous with Ogmaster (Von Mart.) of earlier date, and therefore should be dropped. If it were desirable to retain it at all, it should have been restricted to D. longimana. For the latter, Sladen, 1889, established the genus Iconaster, thus excluding Dorigona from the system.
Perrier, 1876, p. 44, used Dorigona for a section or a subgenus of Pentagonaster, and included in it P. longimanus and P. capella (as Mulleri), thus closely following Gray. But in his later work (1894), he has restricted it to Nymphaster Sladen, a group totally unknown to Gray and to Perrier, himself, in 1876.
_ This total transposition of the name is not justifiable. Perrier, himself, has disapproved of such a course in other cases of the same kind.
Seven species of Mymphaster have been described from the Atlantic and others from the Indo-Pacific. Probably the number pf Atlantic species may be hereafter reduced when direct compari- sons of the types shall have been made. I have studied only the three American species, from the types of Perrier and a good series dredged by the Albatross.
Atlantic species of Nymphaster. “Nymphaster ternalis (Per.). Pentagonaster ternalis Per., 1881, p. 20; 1884, p. 288, pl. x, fig. 1. Nymphaster (?) ternalis Sla., 1889, p. 752. Dorigona ternalis Per., 1894, p. 371.
Puate XXVI. FIGuRE 7.
West Indies, in 416 and 734 fathoms (Blake Exped.). Also dredged by the Albatross, at nine stations in the Gulf of Mexico and West Indies in 196 to 1181 fathoms, muddy bottoms, and at two stations off Brazil.
Nymphaster subspinosus (Per.). 1; 1884, p. 2384, pl. vi, fig. 1.
Pentagonaster subspinosus Per., 1881, p.
Nymphaster (2) subspinosus Sla., 1889, p. _ Dorigona subspinosa Per., 1874, p. 375. _ West Indies, 163 to 209 fathoms (Blake Exped.). Also dredged by the Albatross at two stations in the West Indies, in 338 to 388 fathoms, coral sand and gray sand.
2 i
186 A. EB. Verrilli—Revision Genera and Species of Starfishes.
Nymphaster arenatus (Per.). ’ Pentagonaster arenatas Per., 1881, p. 21; 1884, p. 236, pl. vii, figs. 3, 4.
Nymphaster (?) arenatas Sla., 1889, p. 752. Dorigona arenata Per., 1894, p. 379, pl. xxii, fig. 6, pl. xxiv, figs. 5, 6.
Found on both sides of the Atlantic. It was taken by the Blake in the West Indies, in 164 to 874 fathoms, and by the Travailleur” and Talisman, at many localities, in 157 to 1635 meters. It was also dredged in the West Indies by the Albatross.
Nymphaster Jacqueti (Per.).
Dorigona Jacqueti Per., 1894, p. 383, pl. xxi, fig. 4, pl. xxii, fig. 5. Dorigona prehensilis Per., 1885; 1894, pp. 32, 33. Nymphaster (2) prehensilis Sladen, 1889, p. 752.
East Atlantic, from N. lat. 44° 5’ to 28° 35’, in 540 to 1238 meters. —
Perrier does not explain why he has changed the name of this ~ species from prehensilis to Jacqueti, except that he states (1894, p. 426) that the former is a ‘ variety” of the latter. If no other rea- son exists, the earlier name should be retained. 6
Nymphaster protentus Sladen.
Voy. Chall., xxx, p. 303, pl. 1, figs. 3, 4, pl. liii, figs. 9, 10; 1889. Off Canary Islands, in 1525 fathoms.
Nymphaster albidus Sladen.
' Voy. Chall., xxx, p. 306, pl. li, figs. 1, 2, pl. liii, figs. 5, 6, 1889. Off Cape Verde Islands.
Nymphaster basilicus Sladen.
Voy. Chall., xxx, p. 308, pl. lvii, figs. 8, 9, 1889. Off Brazil, 1200 fathoms.
Two Indo-Pacitic species, described by Sladen, differ from all the others in having a single median row of abactinal radial plates between the dorsal marginal ones, nearly or quite to the tip of the rays. In this group the pedicellarie are high and spatulate, as in Goniaster, and the adambulacral spines are in very regular parallel rows. This group may, therefore, well deserve a distinet generic or subgeneric name, and I would suggest Nereidaster, with WV. sym- bolicus as the type. The two species are as follows :—
A. E. Verrilli— Revision Genera and Species of Starfishes. 187
Nereidaster symbolicus (Sla.), op. cit., p. 297, pl. 1, figs. 1, 2, pl. liii, figs. 7, 8, 1889.
East Indies and Philippines, 28 to 140 fathoms.
Nereidaster bipunctus (Sla.), op. cit., p. 301, pl. lii, figs. 3, 4, pl. liii, figs. 11, 12, 1889.
Off Admiralty I., 150 fathoms.
Subfamily. PSEUDARCHASTERINZ Sla. (emended).
Pseudarchusterince Sladen, Voy. Chall., xxx, p. 109, 1889. Astrogoniine (pars) Per., Exp. Trav. et Talism., pp. 337, 338, 1894.
This subfamily is remarkable for combining, in various ways, the structures that are generally characteristic and distinctive of Gonias- teride and Plutonasteride. The intermediate character of the group is so marked that Perrier and Sladen have differed as to its place: Perrier placed it in his Pentagonasteridx, while Sladen placed it in his Archasteride. In fact, its affinities appear to be nearly evenly balanced between the two groups. True pedicellariz, which might throw light on the subject, are generally absent from all the known species of the typical genera.
The abactinal radial plates are arranged in radial rows, the medium ones larger. They are paxilliform, more or less columnar, with round or elliptical convex tops, and with an enlarged six-lobed or stellate basal portion, the projecting lobes articulating and leaving spaces between them for the papular pores, which occupy large radial areas. About six are arranged singly around each of the plates.
Marginal plates are thick, moderately large and paired ; they have deep fasciolated sutural grooves between them. The dorsal plates are rarely in contact medially, unless close to the tip of the rays. They are covered with close granules or small, crowded, appressed spinules, and the lower ones often have several larger central spines of the same character, but in some species the plates are all evenly granulated.
The adambulacral plates are broad, and usually angular or convex on the furrow margin, so that the furrow is constricted opposite each pair, especially distally. The furrow spines are usually in a ‘curved or divergent series; those of the actinal side may be in longitudinal rows or clustered.
The actinal plates are often numerous, angular, arranged in chey- rons, with the rows parallel to the ambulacral furrows. More or less
188 A. BE. Verrill—Revision Genera and Species of Starfishes.
of those in the rows next the adambulacral plates have their trans- verse edges bordered by specialized spinules, forming with those of the plates opposed to them, special fascioles (pedicellaires fascio- laires of Perrier). But they are not real pedicellariz.
True papilliform pedicellariz occur very rarely on the actinal and marginal plates in Paragonaster.
The jaws are rather large and prominent and bear numerous elongated spinules, both on the actinal surface and the margin. At the oval apex of the jaw there is usually a larger odd median spine, but this is not constantly present in all species, and in certain species it is generally, if not always, lacking. Some specimens may have the odd spine on some of the jaws and lack it on others, so that it cannot be considered of much morphological importance, though its presence is a useful indication of the affinities of certain doubtful species,
The ambulacral feet have well formed suckers, as in Mymphaster
and Mediuster.
In the general appearance of the abactinal and marginal plates and the granulation of the dorsal surface this group agrees essentially with Mediaster and allied genera. It differs from that group mainly in the more prominent margins of the adambulacral plates, the lack
of bivalve pedicellariz, and the more divergent groups of furrow-
spinules, together with the usually spinulose covering of the actinal and inferior marginal plates; but this last character is not constant.
The singular actinal fascioles are characteristic of many species, but are not constant. The same is true of the odd apical spine of the jaws.
If true bivalve pedicellarize were present we should not hesitate to combine the group with Mediaster and Nymphaster in one sub- family.
On the other hand, the adambulacral plates and spines, the jaws, and the form and structure of the dorsal paxille are very much like
those of Plutonaster and allied genera, though the latter does not —
have the regular serial arrangement of the radial abactinal plates. Nor is there in this group any distinct arrangement of the actinal plates in rows running from the adambulacral plates to the marginals, as in Plutonaster and allied genera.
The actinal plates have nearly the same form and are imbricated in the same way as in Mediaster. The adambulacral plates next the dentary plates are somewhat oblique and slightly modified, but much less so than in /7utonaster, and more so than in Tosia and Mediaster.
A. E. Verrili— Revision Genera and Species of Starfishes. 189
The jaws are also intermediate, in respect to size, form, and amount of elevation of the actinal ridges, between Mediaster and Plutonaster. The enlarged stellate bases of the abactinal radial paxille may be considered as a farther development of the slightly enlarged and lobate bases at the columnar paxille of Plutonaster. There are no separate, internal, radiating connecting ossicles between the plates, such as exist in Mediaster. At least, I have not found them in anatomical preparations of Psewdarchaster intermedius and Parago- naster formosus, both of which have enlarged six-lobed bases on the paxilliform plates.
But in Nymphaster (ternalis) the connecting ossicles are also lacking, and the plates are short-columnar, with the bases only slightly enlarged, thick, and but slightly six-lobed, the lobes being rounded and often indistinct. The same is true of the correspond-
ing ossicles of Rosaster.
Considering all these points, the affinities of the group seem to me rather more with NMymphaster and Mediaster than with any other
genera. ‘This is also the view taken by Perrier (1894).
Pseudarchaster Sladen. Type, P. discus Sladen.
Pseudarchaster Sladen, 1885, p. 617. Voy. Chall., xxx, p. 109, 1889. Astrogonium (pars) Perrier, Exp. Trav. et Talism., p. 338, 1894 (not of M. and Troschel, nor of Gray).
The principal characters of this genus have been mentioned in the above description of the subfamily.
The adambulacral plates have a divergent or palmate series of furrow spines and a group or radiant cluster of actinal spines. The actinal plates are generally closely covered with small appressed spinules, often somewhat squamiform, rarely elongated, but fre- quently a few larger spinules exist among the smaller ones. Lower marginal plates are usually spinulated like the actinals, rarely granu-
_ lous, often with one or more central rows of larger appressed
spinules,
Specialized fascioles usually (but not always) exist between more or less of the larger actinal plates, especially toward the jaws. The ambulacral feet have well formed suckers.
The abactinal plates and upper marginals are usually closely
‘granulated. More than one series of abactinal plates usually extend
nearly to the ends of the rays. Perrier united ApAroditaster with this genus, but the type seems to me sufficiently distinct. However, the presence or absence of the
190 A. EB. Verrill—Revision Genera and Species of Starfishes.
specialized actinal fascioles cannot be made a character by which to distinguish them, for they may be present or absent in the same species (e. g. intermedius). Their presence seems to be the normal — condition.
Six species are known to me from off the American coast. Several other species have been described from the East Atlantic.
Pseudarchaster intermedius Sladen.
Pseudarchaster intermedius Sladen, Voyage of the Challenger, vol. xxx, p. 115, pl. 19, figs. 8, 4; pl. 42, figs. 5, 6, 1889. Verrill, Proc. Nat. Mus., vol. xvii, p. 249, 1894. Amer. Jour. Sci., xlix, p. 131, 1895.
Archaster Parelii (pars) Verrill, Amer. Journ. Sci., vol. vii, p. 500, 1874 (not of Diiben and Koren); vol. xxiii, p. 140, 1882; Rep. U. S. Com’r Fish and Fisheries, vol. xi, p. 548, 1884.
Puate XXX. Fuicures 1, 1a, 1b.
This is the most common species off the eastern coast of the United States and Canada.
It was taken at about 33 stations by the Albatross and Fishhawk, 1880 to 1887, in 85 to 1608 fathoms, from N. lat. 44° 26’ to 37° 59’ 30”. It has also been brought from the fishing banks, off Nova Scotia, by the Gloucester fishermen.
The variety (énsignis) named and described by me in 1895 (p. 132), is probably only the fully adult form of this species. The largest example has the larger radius, 75™™; the lesser, 23". It lacks distinct actinal fascioles. These exist, however, in variable numbers, on other similar specimens, of somewhat smaller size, as well as on quite young examples. Their presence does not depend upon age, for they may be absent or present in specimens of equal size. Most specimens have the odd apical oral spine somewhat larger and longer than those adjacent. The genital pores are oppo- site and close to the first pair of dorsal marginal plates.
Pseudarchaster fallax Perrier. Astrogonium fallax Per., 1885. Exp. Trav. et Talism., p. 347, pl. xiii, fig. 4, pl. xxv, fig. 4, 1894. Archaster Parelii (pars) Verrill, Rep. U. S. Comm. Fish and Fisheries for 1883, vol. xi, p. 543, pl. xiii, fig. 37, 1885.
PLATE XXX. FIGURES 2, 2a. This was formerly considered by me a variety of the preceding with narrow dorsal radial areas. More recently I have compared our specimens with one of the types of P. fallax Per., in the Mus. of Comp. Zoology. ‘They agree with the latter in all respects.
A. E. Verrill— Revision Genera and Species of Starfishes. 191
The species can be distinguished by the larger and more massive marginal plates, which encroach farther upon the dorsal surface, and by the very narrow abactinal areas on the rays. The granulation of the actinal plates is also closer and the adambulacral spines are shorter than in P. intermedius, but the two are very closely related. ‘There is an odd apical spine on the jaws. Actinal fascioles are generally present.
Pseudarchaster (?) hispidus Ver., sp. nov. PLATE XXX. FIGURE 5.
Pentagonal with moderately long rays. Radii as 1: 2.
About twenty-five marginal plates, above and below ; these are rectangular, broader than long, not oblique. The upper ones extend only a short distance on the dorsal surface, and are only a little convex. _ They are covered with numerous very small, slender spinules; those on the middle are erect, but those on the margins form fascioles of very slender spinules.
Inferior marginal plates extend far within the margin; they are spinulated much like the upper ones, but the spinules are larger, longer, tapered, acute, arranged obliquely and divergently in four or five rows, not counting the marginal fascioles ; usually none of the median ones are distinctly larger than the rest, but sometimes, on a few plates, one or two of the distal ones are somewhat larger and longer.
Abactinal paxillz are relatively large, rounded, and nearly uni- form in size. There is a somewhat distinct median row on the rays. About six rows occur opposite the third pair of marginal plates ; they are reduced to three rows near to the end of the rays, and to one median row between the last three plates. They are uniformly covered with small, sharp, elongated, divergent spinules, often thirty or more in a group. Of these, twenty or more may be marginal and a little smaller than the others, the adjacent ones interlocking so as to conceal the papule. These appear to be small and few. The plates are round, elevated, convex or somewhat clavate, well sepa- rated,
Interradial actinal regions are of moderate size, triangular, with the outer plates extending out to about the seventh pair of inter- ambulacral plates. They are in rows parallel with the adambula- crals and not separated by radiating grooves. They are rather large, roundish, covered with rather long, divergent, acute spinules, often nine to twelve on the larger ones.
192 A. E Verrill— Revision Genera and Species of Starfishes.
Jaws a little prominent, bearing a large number of slender mar- ginal spines and very numerous similar ventral ones, in about two crowded rows on each half. The apical spines are only a little larger than the marginal ones.
Adambulacral plates on the middle of the rays bear about four or five relatively very long and very slender, terete spinules on the convex marginal edge ; one to three on the actinal surface, of simi- lar size and form, and four or five divergent ones on the outer mar- gin, that are shorter, but of the same form. No pedicellariz were seen. ‘The abactinal and ventral plates and paxille are much larger and fewer than in any species of Plutonaster or Dytaster, of similar — size, and the appearance is decidedly hispid under a lens, owing to the elongated and acute spinules of the whole surface. Greater radius, 12™™; lesser, 6™™. :
Taken by the Blake Expedition in the West Indies, in 600 fathoms, and by the U. 8. Fish Commission Steamer Albatross.
The specimens of this species that I have examined are doubtless immature, but they differ decidedly from the young of the other known American species. It is not a typical Pseudarchaster.
The specimen from the Blake Expedition was mixed with speci- mens labelled as Plutonaster intermedius by Perrier.
Pseudarchaster granuliferus Ver., sp. noy. PLaTE XXX. Ficures 6, 6a.
Form stellate with a broad disk and deeply emarginate sides, the rays wide at base and tapering rapidly to acute tips. ,
Radii as 1:2.20. Greater radius, 22™™; lesser, 10™™.
The marginal plates are large and thick, encroaching considerably on both sides of the disk, producing a rather thick rounded margin. The upper ones are closely covered with polygonal granules; the lower ones are closely covered with small, uniform, closely appressed, ovate, subsquamiform granules or granule-like spinules, without any larger median ones.
The abactinal plates are small, rounded, elevated, and covered with a polygonal group of prismatic granules, about five to eight forming the central cluster. Papular pores are regularly arranged, and placed singly between the basal radial plates. The madreporic plate is small and irregular,
Actinal plates are crowded and covered with spaced polygonal granules, without any spinules. On each area there are three to five special pectinate fascioles of small size, one of which is opposite
A. E. Verrill—Revision Genera and Species of Starfishes. 193
the dentary suture. The adambulacral plates have a curved, pal- mate, strongly projecting furrow-series of five or six unequal blunt spinules; the two or three median ones are larger and more slender, compressed ; the two distal ones are shorter, stouter, flattened or spatulate, but on the distal half of the rays they are all slender. The actinal side is covered with shorter, obtuse or clavate spinules, either clustered or in two or three irregular rows, but without any larger central spines.
The jaws are not prominent ; there is an odd apical spine on each jaw, distinctly larger and longer than the rest ; the furrow margin bears ten to twelve small obtuse spinelets, like those of the adambu- lacrals ; the distal ones are rather larger than the others. On the sutural margin there are about ten shorter blunt or clavate spinules, a little larger than the actinal granules ; another small intermediate row of similar ones covers the actinal surface.
Taken by the Albatross at station 2751, in example (No. 18,448a).
fathoms. One
Pseudarchaster concinnus Verrill.
Pseudarchaster concinnus Verrill, Proc, U.S. Nat. Mus., vol. xvii, p. 250, 1894. Amer, Jour. Sci., xlix, p. 132, 1895.
PLATE XXX. FiGurReEs 3, da, 3b.
Taken at 3 stations between N. lat. 41° 28’ 30” and 41° 07’; in 1188 to 1791 fathoms.
In this species the genital pores are large and usually plainly visible, without preparation. They are situated far apart on the dorsal surface, about opposite the second pairs of dorsal marginal plates; they are separated by about nine radial rows of abactinal interradial plates ; about six plates intervene between the pore and the marginal plate.
The jaws often have an odd apical oral spine, only a little longer and larger than those adjacent. In other cases no such spine is found, the apical spines being all paired. This variation may occur on the different jaws of the same specimen.
In this species there is less distinction between the smaller and larger spines on the lower marginal plates, there being many inter- mediate in size, and the largest not very large, in three or four irregular rows. The adambulacral and dentary spines are shorter, those of the actinal side of the jaws being much less conspicuous. The larger central spine of the actinal plates is also less prominent.
Trans. Conn. Acap., Vou. X. Avaust, 1899.
13
194 A. EL Verrill—Revision Genera and Species of Starfishes.
Pseudarchaster ordinatus Ver., sp. nov. PuateE XXX. Ficures 4, 4a, 4b.
A large species, having a broad disk, with the sides regularly incurved, and rather long, tapered, subacute rays, closely resembling P. concinnus in form and in the character of the upper side, but more spinose below.
Radii about as 1:2.8. Greater radius, 47 to 50™™; lesser, 17-162"
The abactinal paxille are regularly arranged, and evenly granu- lated, with very small, crowded, slightly elongated, round granules, — of which twenty to thirty may occupy the central part ; those around the margin are longer and divergent.
Upper marginal plates about 64 on each side of the body, much higher than long, encroaching considerably on the disk, sloping upward so as to form a bevelled margin. They are rather closely _ and finely granulated, like the abactinal plates. They have the narrow sutural grooves fasciolated.
Lower marginals similar to the upper in size and shape, but cov- ered with small, acute, unequal spinules, and with one or two median vertical rows of larger appressed spines, of which there may be 8 to 10 or more on the larger plates.
The actinal plates mostly have a long, rather slender, acute central — spine, surrounded by several small acute, erect spines. Many of those of the principal series have pectinate fascioles between them. Sometimes as many as 16 of these special fascioles occur on each interradial area.
The adambulacral plates bear a palmate furrow-series of seven or eight slender, divergent, nearly equal spines; one or two larger central spines on the actinal side; and an outer marginal curved row of several small acute spinules.
The jaws usually have an odd apical spine considerably larger than the rest, but it may be lacking on some jaws; there are about eight or nine spines in the furrow series, rather longer and larger than those farther out ; and about seven to nine rather larger and longer actinal spines on each dentary plate, so that the jaws appear very spinose. .
The genital pores are small but easily visible; they are situated opposite to the second pairs of dorsal marginal plates.
Taken by the Albatross in the Gulf of Mexico, at station 2396, in 335 fathoms (No. 18,438) ; also at station 2376, in 324 fathoms. |
x
A, E. Verrill—Revision Genera and Species of’ Starfishes. 195
This species has a much thinner disk and more slender and more rapidly tapered rays with less massive plates that P. intermedius, or even P. concinnus.
The following additional species have been recorded from the East Atlantic :
Pseudarchaster annectens (Per.).
Astrogonium annectens Per., Exp. Trav. et Talism., p. 343, pl. xxiii, fig. 5, pl.
xxiv, fig. 1, 1894.
Gulf of Gascony, 900 meters; station 213, 1888, 1384 meters. This is very closely related to P. intermedius Sla.
Pseudarchaster hystrix (Per.).
Astrogonium hystrix Per., Exp. Trav. et Talism., p. 345, pl. xxiii, fig. 3, pl. xxiv, fig. 2, 1894.
Coast of Morocco, 840 meters, one example. Very closely related to the preceding.
Pseudarchaster necator (Per.).
Astrogonium necator Per., Exp. Trav. et Talism., p. 350, pl. xxiii, figs. la, 10,
1894.
Off the Azores, 1257 meters, one example.
Pseudarchaster Aphrodite (Per.).
Astrogonium Aphrodite Per., Exp. Trav. et Talism., pp. 342, 354, pl. xxi, fig, 2, pl. xiii, fig. 2, 1894.
Coast of Sahara, 1090 meters.
Aphroditaster gracilis Sla. Aphroditaster gracilis Sla., Voy. Chall., xxx, p. 117, pl. xvii, figs. 1, 2, pl. xviii, figs. 7, 8, 1889. Astrogonium gracile Per., Exp. Tray. et Talism., pp. 542, 354, 1894. Off the Azores, 1000 fathoms.
Pseudarchaster tessellatus Sla. is from off the Cape of Good Hope.
P. Patagonicus (Per.) is from Patagonia, 283 meters.
P. discus Sla. is from Messier Channel, west coast South America, 147 fathoms.
A few species have been described from the Indo-Pacific region :
196 A. E Verrilli—Revision Genera and Species of Starfishes.
P. mosaicus Alcock and Wood Mason, is from the Andaman Sea, India, in 188 to 220 fathoms.
P. roseus (= Mediaster roseus Alcock, 1893, p. 98) is from the Laccadive Sea, in 740 fathoms.
Paragonaster subtilis Perrier. ;
Goniopecten subtilis Perrier, Bull. Mus. Comp. Zo@l., ix, p. 26, 1881. Mém.
Etoiles de Mer, p. 258, pl. v, figs. 3, 4, 1884.
Goniopecten subtilis Sladen, Voy. Chall., xxx, p. 726, 1889.
Paragonaster subtilis Perrier, Exp. Tray. et Talism., p. 358, 1894.
The type of this species, from station 31, Blake Expedition, I have ~ compared with specimens of P. formosus Ver., of similar size.
The two are very closely related, but in P. subtilis the adambu- lacral plates usually have, on the actinal surface, a rather long and stoutish acute central spinule; and the outer marginal spinules are also larger than those of formosus. On the proximal plates there. are usually 6 or 7 furrow spinules and 8 or 9 on the actinal surface, all of which are stouter than in jformosus. The spinules on the actinal surface of the dentary plates are also more numerous, larger and more divergent than in formosus; they form four rather irregu- — lar rows. The larger actinal paxille have 14 to 16 marginal granules, with 4 to 6 larger central ones.
Possibly a large series of specimens would compel us to unite the two, as only varietal forms of one species.
Four other Atlantic species of this genus have been described. They are as follows: ‘
P. formosus (Ver.) 1884, p. 383); 1894, p. 257; 1895, p. 187. Off East Coast of United States, 1396 to 2021 fathoms.
P. strictus Per., , 1894, p. 363, pl. xxv, fig. 3. i East Atlantic, 3665 meters.
P. elongatus (Per.), 1885. 1894, p. 362, pl. xxi, fig. 3, pl. xxiv, fig. 4.
Off the Azores, 2115 to 4060 meters.
Perrier suggests that this may be only a variety of P. subtilis and that P. strictus may be the young of the same species.
P. cylindratus Sladen, Voy. Chall., xxx, p. 314, pl. li, figs. 3, 4, pl. liii, figs. 3, 4, 1889.
Off Cape Verde Islands, in 1850 fathoms. Closely related to P. Sormosus.
A, E. Verrill— Revision Genera and Species of Starfishes. 197
Rosaster Alexandri Perrier.
Pentagonaster Alexandri Perrier, Bull. Mus, Comp. Zoél., ix, p. 22, 1881. Nouy. Arch. du Mus., vi, p. 238, pl. vi, figs. 3-8, 1884. Rosaster Alexandri Pen. , Exp. Sci. Tray. et Talism., Echinod., p. 387, 1894.
This species has rounded, columnar, paxilliform abactinal plates, covered, like the marginal and actinal plates, with small spinules. Most of the upper marginal plates of the rays are in contact medially.
The genus Fosaster is evidently very distinct from Paragonaster. Perrier states that it has no pedicellariz of any kind, but some of the larger specimens that I have examined have had a small number of simple pedicellariz on the actinal plates.
The larger examples have two long, slender spines on the actinal side of the adambulacral plates.
It was taken at several stations by the Blake, in 84 to 1930 fathoms, and by the Albatross at a number of stations in the West Indies and Gulf of Mexico, in 182 to 980 fathoms.
INCERTA SEDES.
Hoplaster Perrier. Type, H. spinosus Per. Hoplaster Perrier, 1882, Rapport, p. 32. Exped. Trav. et Talism., p. 323, 1894.
Form pentagonal with short rays. Marginal plates well-developed, not numerous, spinulated. An odd marginal interradial above and below. Abactinal and actinal plates angular, crowded, closely united, covered with a group of elongated spinules. No pedicel- larie observed. Adambulacral plates with three or four spinules in the furrow-series and an irregular group of spinules on the actinal surface. Jaws without a recurved spine.
The relations of this genus are doubtful. Perrier placed it next to Gnathaster, on account of the odd marginal plate, etc. (See p. 202). It may, perhaps, belong to Goniasteride, or be allied to Lasz- aster. he details of its skeleton are not known.
Hoplaster spinosus Perrier, 1882, Rapport, p. 32. Exped. Tray. et Talism., p. 324, pl. xiv, figs. 2a, 2b, 1894.
Off the Azores, etc., 2995 to 3307 meters. Only small examples are known.
198 A. BE. Verrill—Revision Genera and Species of Starfishes.
Hoplaster lepidus (Sladen). Pentagonaster lepidus Sladen, Voy. Chall., xxx, p. 275, pl. lvii, figs. 1-4, 1889. _
This species agrees so closely, in all structural characters, with the type of the genus, that there can be little doubt that they belong to one genus. In fact, the present species might even prove to be an older state of the former, to judge from the descriptions. They” both came from the same region and similar depths.
Off the Azores, 1000 fathoms.
Lasiaster Sladen. Type, L. villosus Sladen.
General form as in Zosia and Goniaster. Marginal plates well- developed, in both series, paired. No odd interradial. Marginal, actinal, and abactinal plates covered with groups of small, acute spinules.
The general appearance of this genus is similar to Hoplaster, with — which it may, possibly, be related, though no odd interradials are present. M. Sladen refers the genus to the Gymnasteride.
Lasiaster hispidus (Sars) Sladen.
Goniaster hispidus M. Sars, Fauna litt. Norveg., iii, p. 72, pl. 8, figs. 24, 25, 1877.
Pentagonaster hispidus Perrier, Nouv. Arch. du Mus. d’hist. Nat., Ser. 2, vol. i, p. 84, 1878. Danielssen and Koren, Asteroidea, Norske Nordhays-Exped. Zool., xi, p. 58, pl. xv, fig. 6, 1884.
Lasiaster hispidus Sladen, op. cit., p. 372, 1889.
Arctic coasts of Europe, especially in the Drontheim Fjord, in
deep water.
The larger specimens are 72™™ in diameter.
Revision of the Classification of the orders Valvata and Pazxillosa of Perrier, and especially of the Archasteride.
Archasteridz Sladen, Voy. Chall., xxx, pp. 1-4, 1889. Perrier, Exp. Tray. et Talism., pp. 2387-252, 1894.
In a former article (1894, pp. 266-269) I endeavored to show that this extensive group is probably not a natural family.
This opinion has been confirmed to some extent by the subsequent publication of Perrier’s report, quoted above, in which he discussed, at considerable length, the characters of this “ family,” as contrasted with Pentagonasteride. Te enumerated seven principal characters by which the two families are distinguished. It is sufficient to state
€ Ae
A, E. Verrill—Revision Genera and Species of Starfishes. 199
here that every one of these seven characters fails in certain cases, and that nearly all of them may occur in each family, so that there is no certain means of deciding in which family certain genera should be placed. Perrier, himself, admits something of the kind, bunt holds that the preponderance of the characters ought to deter- mine the family in each ease.
The recent discovery of new genera has so increased the excep- tional cases, by revealing forms that’ are more or less completely intermediate between the two groups, that it has become difficult to define them in any satisfactory manner.
The two principal writers who have recently discussed the classi- fication of these starfishes, Sladen and Perrier, have differed con- siderably as to the limits and characters of each group. Thus Sladen included the Odontasteride (as Gnathaster) and the genera Mimaster and Leptogonaster in the Pentagonasteride, but Perrier put all these in the Archasteride. On the other hand, Sladen puts Pseudarchaster and Aphroditaster in the Archasteridze, but Perrier transfers them to the Pentagonasteride.
These are well-known genera that have been thoroughly studied by both writers, therefore we must conclude that the two so-called families are not really well defined, natural groups, otherwise such able investigators could hardly disagree to such an extent.
This question would be of less importance were it not for the fact that in the more general classification of Perrier, these two “ fami- lies” belong to two distinct orders. The Archasteride he places in the order Paxillosa (op. cit., pp. 28, 29); the Pentagonasteride in the order Valvata.
The fact that the two so-called families run together, without definite limitations, would necessarily imply that these two “ orders ” are also badly limited or unnatural groups.
Almost the only special character by which the two groups can be distinguished, as limited by Perrier, will be the character of the pedicellariz, which, however, are often lacking in both groups.
But the papilliform pedicellariz of the Paxillosa, with two to four or more valves, apparently formed from modified spinules or gran- ules, are also found in the Valvata. Sometimes such pedicellarize are found associated with larger valvular pedicellariz on the same specimen, in the genus Nymphaster and other genera, while well- formed, though small, bivalve pedicellarix often occur on certain of the antarctic Gnathasterine, and on other species referred to Paxil- losa.
200 A. E. Verrill—Revision Genera and Species of Starfishes.
Therefore, if this feature is to be the criterion, the Paxillosa should only include such groups as never have true bivalve pedicel- lariw. The existence of paxilliform plates on the dorsal surface can- not be made an important character, for they occur in typical forms of Valvata. The development of terminal suckers on the ambulacral feet varies much in both groups, and depends mainly on the nature . of the bottom inhabited.
The Paxillosa would be a more natural group if limited to the Porcellanasteride, Astropectinide, and the genus Archaster, while the rest of the Archasteride (Sla.) might go into the Valvata (sens ext.) However, it seems to me a more natural atrangement to con- sider these groups as the two suborders of one order, equivalent in rank to the three others proposed by Perrier. For this order Sladen’s name, Phanerozona, might well be used, in a slightly restricted sense, the Asterinide and Gymnasteride being excluded.
The classification of the order as now proposed would be as —
follows:
Order PHANEROZONA Sladen (rest.). Suborder I.—Valvata Perrier (sens ext.).
Family 1.—Lincx1p Perrier. Family Il.—Prnracrerotip Gray (restr.). Family I1].—Anrueneip& Per. (restr.). Family [V.—GontastErip& Forbes (restr.). Subfamily I.—Goniasterine V., nov.=Pentagonasterine Sla. (pars). Subfamily II.—Goniodiscine Sla. Subfamily Il].—Mediasterine V., nov. Subfamily TV.—Pseudarchasterine Sla. Subfamily V.—Hippasteriine V., nov. Family V.—OponrTastERip& Ver. noy.=Gnathasterine Per.( pars). Family VI.—PiLuronasTERID& Ver. Subfamily I.—Mimasterine Sla. Subfamily If.—Plutonasterine Sla. Subfamily II.—Pontasterine Ver., 1894. Family VII.—Gonroprctinip& V., nov. Family VII.—Benrnorectinip& V.=Benthopectinine Ver., 1894.
>
A. E. Verrill— Revision Genera and Species of Starfishes. 201
Suborder Il.—Paxillosa Per. (sens. restr.).
Family IX.—PorcreLLanastERID& Sla.
Family X.—ArcuHastTEeRID# Vig. (restr. to Archaster). Family XI.—Astropectinip® Gray (restr.).
Family XII.—Luipup V., nov.=Ludiine Sla.
It will be noticed than in the above arrangement the Archasteridz of Perrier, 1894, has been divided into five distinct families (Families V, VI, VII, VIII and X). The larger number of genera are placed in the Plutonasteride, which includes three groups that appear to be of subfamily rank.
The synonymy given will sufficiently indicate the limits of the groups in most cases.
The new family, Goniopectinide, is proposed to include Gonzo- pecten Per. (restr.), type G. demonstrans, together with an allied new deep sea genus Prionaster Ver., type P. elegans, with odd inter- radial marginal plates and a corresponding odd row of actinal plates. It is from the West Indies. The genus Craspidaster Sla., which I have not seen, probably belongs to the same family. In this group the adambulacral, actinal, and marginal plates are surrounded by . special spinules united together by a web, so as to form very spe- cialized fasciolated grooves. It is related to the Astropectinide, as well as to Pontasterine.
The family Benthopectinide includes, so far determined, only the genus Benthopecten Ver.= Pararchaster Sladen.
The family Odontasteridex is proposed for Odontaster, Gnathaster, and allied forms, having one or two large recurved spines on the jaws, and also odd interradial marginal plates. It is equivalent to Gnathasterine Per., minus Hop/laster.
Archasteride is restricted to the typical genus Arehaster, which is believed to be closely allied to the Astropectinide.
Family ODONTASTERIDZ Ver., nov. Gnathasterine (pars) Perrier, Exp. Trav. et Talism., pp. 244, 251, 1894.
Form either pentagonal or stellate with a broad disk. Marginal plates well-developed.
Jaws, each with a single, recurved, more or less hyaline median spine, or with two such spines, side by side. In the latter case one
202 A. E. Verrill—Revision Genera and Species of Starfishes.
of these spines arises from near the apex of each dentary plate. Both conditions sometimes occur, abnormally, on the same specimen. An odd interradial marginal plate, above and below, on each side. Abactinal surface covered with more or less paxilliform plates,
parapaxille or protopaxille, with intervening large papular pores— on the radial areas. The abactinal plates may bear clusters of more —
or less elongated spines, or a group of small granules. They usually form obliquely transverse lines on the rays, not always regular.
Actinal plates angular, covered either with spines or granules. Small simple pedicellariz sometimes occur on the actinal or abactinal plates. They may have two, three, or four papilliform blades.
Adambulacral plates usually bear elongated spinules arranged in three or four pairs of small transverse rows, generally only two or three of the furrow-series are on each plate ; sometimes only one. Dentary plates usually have elongated, acute marginal and apical spines. hey are sometimes closely united along the median suture; in other cases (Odontaster), they are separated by a space covered only by membrane.
The marginal plates are covered either with spinules or granules; sometimes the upper ones are granulated and the lower spinulose, like the corresponding disk-plates; they usually have deep fas- ciolated sutures.
Perrier, 1894, instituted a sub-family under the name Grathaster- ine to include the present group, together with some other forms (Hoplaster) in which no recurved jaw-spines occur. He based the group more particularly on the odd interradial marginal plate. But the latter character seems to me to be of less importance, for I have found it abnormally present in various species of Zosia and allied genera, (See under Pyrenaster dentatus, p. 167, above). Moreover in Benthopecten, which normally has an odd interradial plate, I have found it replaced by two plates, on some of the margins (see p. 218 below and pl. xxx, figs. 7, 7a).
Therefore, I have taken the presence of the recurved dentary spines as the special feature of the group.*
As the name @nathaster is nearly a synonym of Odontaster, and may, therefore, be dropped from the system by future authors, it seems desirable to change the name of the group to Odontasteride.
Perrier has divided the group into three genera: Gnathaster, Asterodon, and Goniodon.
* A similar recurved tooth is found in certain species of Pterasteride@.
A, E. Verrili— Revision Genera and Species of Starfishes. 203
Asterodon Per. has a pair of dentary spines on each jaw.* Type, A. singularis (M. and T.). Peru and Chili.
Goniodon Per. has some of the distal marginal plates enlarged. A pair of recurved spines on each jaw. Type, G. dilitatus Per., New Zealand.
Gnathaster Sladen (restr.) has the marginal plates regularly de- creasing. One recurved spine on each jaw. Type, G. pedicellaris Per.= G. meridionalis (t. Leip.), Cape Horn.
As thus limited and defined Gnuathodon is identical with Odontas- ter Ver., of earlier date. The type cited, however Odontaster. :
When Sladen established his genus Gnathodon he included in it
, 1s not a typical
all the known forms belonging to the three divisions proposed by
Perrier. He did not designate any particular species as the type.
His personal studies and detailed descriptions and figures were de- voted to three Antarctic species, all of which would go in the genus, as restricted by Perrier.
Bell, 1893, combined all the known forms under the name Odon- taster Ver., which is the earliest generic name in the group. The latter was based by me on a single species (0. hispidus), which is one of a group of species having very spinulose paxille and plates, and is apparently not strictly congeneric with O. meridionalis Smith (= 0. pedicellaris Per.), which Perrier cites as the type of Gnatho- don, 1894.
It seems to me unnecessary, therefore, to consider Gnathodon (Perrier, restr.) a synonym of Odontaster.
Since Odontaster was originally established for a more restricted group, Sladen’s name was not originally truly synonymous with it. Therefore it may well be restricted to one of the other subdivisions included in it by him, as Perrier has done imperfectly.
Another division may be established for G@. elongatus Sl., and G. miliaris Gray, in which the abactinal skeleton consists of pseudo- paxille, or low rounded plates covered with granules, while granules also cover the actinal and marginal plates, thus giving them nearly the same appearance as species of Zosia and allied genera, for which, indeed, some of them were formerly mistaken and described (under Astrogonium by Gray, and others; and under Pentagonaster by Perrier).
* Leipoldt, 1895, figures an abnormal specimen of A. singularis in which two jaws have each but one median tooth (pl. xxi, fig. 7c).
204 A. EB. Verrill—Revision Genera and Species of Starfishes.
Acodontaster Ver., gen. nov. Type, G. elongatus Sladen.
Gnathaster (pars) Sladen, Voy. Chall., xxx, p. 285, 1889. Perrier (pars), p. 244 (1894).
Odontaster (pars) Bell, Proc. Zodl. Soc. London, p. 261, 1893. Leipoldt, Zeit. wissenschaft. Zoél., lix, p. 614, 1895.
One odd median, recurved, hyaline spine* on each jaw, or angle of - the mouth-frame. The two dentary plates are closely united along — the suture. ;
Actinal, marginal and abactinal plates are covered with granules or short granule-like spinules.
Abactinal plates have the character of pseudopaxille, or are not truly paxilliform, nor much elevated. They form obliquely trans- verse rows on the rays. Papular pores are large in the radial areas.
The marginal plates decrease regularly in size distally. The adam- bulacral spines are arranged in several series; two of each series usually are situated on each plate. ;
Actinal plates form series in two directions. Adambulacral plates usually bear only two spines in the furrow-series. Pedicellariz are not found in the type. The distribution is Antarctic.
The following species belong to this group.
Acodontaster elongatus (Sladen, 1889). Off Marion I.; off Heard I.; off Kerguelen IL. etc., 50-150 fathoms,
Acodontaster miliaris (Gray, 1847). New Zealand.
Gnathaster Sladen (restr.). Type, G. meridionalis.
Gnathaster (pars) Sladen, Voy. Chall., xxx, p. 285, 1889. Perrier (pars), Exp. Trav. et Talism., p. 244, 1874.
Odontaster (pars) Bell, Proc. Zoél. Soe. London, p. 261, 1893. Leipoldt, Arch, wissen. Zodl., lix, p. 614, 1895.
A single hyaline recurved spine (movable?) on each jaw. The ; two dentary plates consolidated at the suture.
Abactinal plates are elevated, convex or capitate, and with radial basal processes. They are covered with a group of short spinules or with prismatic granules. They extend to the apical plate.
Dorsal marginal plates not very large, covered with granules or with small short spinules, like the disk. Ventral marginal plates and actinal plates covered with granules or minute spinules.
* According to Sladen’s description these spines, in his species, are more closely united to the jaw than in Odontaster, and would hardly be movable. In the latter they are only attached by their bases, at the apex of the jaw, and are movable.
A. E. Verrill—Revision Genera and Species of Starfishes. 205
Adambulacral plates are narrow and usually have only two furrow spines; several other pairs are borne on the actinal side of each plate.
G. pilulatus Sladen also belongs to this restricted group.
G. pedicellaris Per., from Cape Horn, is placed as a synonym of meridionalis Smith by Leipoldt, as are, also, G. Grayi (Bell) and G. pilulatus Sladen. All the species are Antarctic.
Odontaster Verrill.
Odontaster Verrill, Amer. Journ. Science, xx, p. 402, 1880. Proc. U. S. Nat. Mus., xvii, p. 262, 1894. Amer. Journ. Sci., xlix, p. 136, 1897.
Gnathaster Sladen (pars), Voy. Challenger, vol. xxx, Asteroidea, p. 285, 1889. Perrier (pars), Exp. Tray. et Talism., p. 244, 1894.
Odontaster Bell (pars), Proc. Zod]. Soe. London, p. 260, 1893.
A single, odd, hyaline, recurved movable spine on the apex of each jaw. Dentary plates large, separated by an open, fusiform space covered by membrane. Abactinal surface covered with ele- vated, convex or clavate paxilliform plates, or parapaxille, which usually bear clusters of elongated spinules, like true paxille ; their bases are stellate; upper marginal plates are usually finely spinulated,
Lower marginal plates and actinal plates are covered with acute, more or less elongated spinules.
Papular pores are generally large and placed singly in the angles around the radial paxille. The radial abactinal plates form more or less evident obliquely transverse rows and extend nearly or quite to the apical plate.
The odd interradial marginal plate is usually triangular or wedge- shaped. Simple pedicellariz occur rarely.
The adambulacral plates usually bear several rows of spines; usually three or four in the furrow-series, rarely but two.
The species, so far as known, are from the North Atlantic.
The open suture between the dentary plates of the jaws; the movable hyaline spine, attached only by its base, at the apex of the ~ jaw, together with the very spinose character of the abactinal pax- ille and marginal plates, separate this genus from its allies. The marginal plates are also larger than in most of the other groups, and the adambulacral plates bear usually three or four spines in the furrow-series.
A reexamination of the numerous specimens of this genus for- merly collected by the U. S. Fish Commission Steamer Albatross, off our coast, convinced me, several years ago, that two species were comprised under the name of O. Aispidus in our earlier lists, and
206 A, FE. Verrill—Revision Genera and Species of Starfishes.
probably in the collections sent to various museums by the U. §. Fish Commission and National Museum. ‘Two examples of another new species was also discovered in the same collections. I have, therefore, thought it desirable to prepare comparative descriptions of these two new species, and to give morphological figures of the three forms, for comparison.
Odontaster hispidus Verrill.
Odontaster hispidus Verrill, Amer. Journ. Sci., vol. xx, p. 402, 1880. Proc. U. S. Nat. Mus., vol. xvii, p. 263, 1894. Amer. Journ. Sci., vol. xlix, p. 136, 1895.
Puate XXIX. Ficures 3, 3a.
Form depressed, stellate, with a rather broad disk. Radii vary in proportion from 1:2 to 1:3. The rays taper regularly and are subacute.
The marginal plates are only moderately developed and do not. encroach much on the disk, either above or below. In large exam- ples there are about 37 to 39 on a side, in each series. They are convex and separated by wide and rather deep sutural grooves. The upper and lower nearly coincide. The upper ones are squarish, with rounded angles; the lower ones, along the disk margin, are higher than long. The odd interradial one is somewhat wedge-— shaped, and only a little smaller than those adjacent to it.
The marginal plates of both series are densely covered with small elongated, divergent spinules which over-arch and partly conceal the sutural furrows. The spinules on the upper plates are slender and acute. Those on the lower plates, especially those on the actinal side, are longer and much stouter, terete and tapered, sub- acute or acute. When the spinules are removed the marginal plates are covered with small hemispherical elevations, where the spinules were attached. Those of the upper plates are smaller and more crowded.
The abactinal plates are round at top, convex, well separated ; those of the radial areas and center of the disk are elevated, with a somewhat capitate round top, which is covered by a dense cluster of slender, elongated, acute divergent spinules.
Between most of the radial plates, over a large area, there are moderately large papular pores, about six around each plate, placed singly in most cases.
Smaller pores are scattered over the center of the disk, but they are absent from the small interradial areas and from the distal part of the rays.
S's
A, E, Verrill— Revision Genera and Species of Starfishes. 207
The actinal plates, when denuded of spines, are numerous, decidedly convex, with deep sutural grooves between them; their surfaces are covered with uneven irregular elevations, where the spines were attached. They are arranged in about five rows parallel to each ambulacral furrow. ‘The first row extends nearly to the tip of the ray; its plates are larger and rather more square than those of the next row. The interradial plates become small, rounded and crowded. The actinal plates all bear dense groups of rather stout, elongated, tapered, mostly acute or subacute spinules, essentially like those of the lower marginal plates.
The adambulacral plates are transversely oblong, rather narrower than the adjacent actinal plates, and have, like the latter, a tuber- culated surface. Each one, proximally, bears two, or more often three, unequal spinules of the furrow-series, but more distally they
bear only two, nearly equal ones. On the actinal side each plate
bears about four or five quite similar spines, which sometimes seem to stand, more or less distinctly, in pairs. These spines, like those of the furrow-series, are essentially like those of the actinal plates, in size and form.
The jaws are rather large, rhombic; the two dentary plates are separated by a rather wide sutural furrow covered with membrane ; they are covered with spines on the margin and actinal side, like those of the adambulacral plates. The median recurved spine is large, somewhat compressed; the distal part is hyaline and very acute.
A large example has the greater radius 55™™; lesser, 16™™. Another has the greater radius 42™™"; lesser, 14™™.
This species has been taken by the U. S. Fish Commission at many localities, from off Martha’s Vineyard to Florida, in 43 to 480 fathoms and more.
It is easily distinguished by the small marginal plates and stout actinal spinules.
Regularly 4-rayed and 6-rayed specimens have been taken.
Odontaster setosus Ver., sp. nov. Puate XXIX. Fiaures 1—I1ec, 2.
Form depressed, stellate, with a broad disk. Sides regularly in-
0 : as : curved. Radii about as 1:2, somewhat variable. The marginal plates
are pretty well-developed and encroach considerably upon the disk, above and below. They are transversely oblong, distinctly higher
208 A. EL Verrill—Revision Generu and Species of Starfishes.
than broad, and separated by deep sutural grooves. They are de- cidedly larger than in O. hispidus and not so square. The upper and lower ones correspond closely in size and position, so that the sutural grooves are continuous. Large examples have about 35 in each series on each side of the body.
In some specimens as many as five or six of the distal dorsal mar- | ginal plates are in contact medially ; in others all are separated, the abactinal plates reaching even the apical plate.
The marginal plates, above and below, are thickly covered with large numbers of small, slender, acute spinules, those near the mar- gins divergent and forming fascioles. The spinules of the upper plates are rather smaller and more numerous than those of the lower ones, but there is no such difference in character as in O. hispidus, When the spinules are removed the plates are thickly covered with minute tubercles.
The abactinal radial plates are well separated, small, paxilliform — with a rounded, convex or capitate top, covered with a cluster of slender, acute, setiform, divergent spinules.
The papular pores are large and conspicuous and occupy large areas; they are placed singly. The actinal plates are rather numer- ous, rhombic, finely tuberculated, arranged in three or four rows parallel with the ambulacra; the first series extends to about the seventh marginal plate. They are covered with dense clusters of slender, acute, setiform spinules, like those of the upper surface but longer.
The adambulacral plates are transversely oblong, narrower than the adjacent actinal ones. They bear each three or four slender furrow spinules in a nearly regular row, and a dense group of 10 to 12 or more, somewhat longer, slender spinules on the actinal side. The latter are similar to the actinal spinules, but rather larger and less acute.
The dentary plates bear marginal and actinal spines, similar to those of the adambulacral plates, but those at the apex are shorter, prismatic and blunt, while there are usually two or three near the sides that are larger than the rest and somewhat curved. ‘The plates are separated by wide sutural grooves.
The recurved spines are compressed and often somewhat curved ; the distal end is hyaline, suddenly narrowed or acuminate, and usually very acute.
One of the larger specimens has the greater radius 37™™ ; lesser, 18™™, Another has the greater radius 32™"; lesser, 17™™.
A, E. Verrill— Revision Genera and Species of Starfishes, 209
This species was taken by the U. 8S. Fish Commission Steamer Albatross, at many stations, from off Martha’s Vineyard to the Carolina coasts, in 56 to 400 fathoms or more. It was often asso- ciated with O. hispidus.
In form and general appearance it resembles O. hispidus, but is easily distinguished by the higher marginal plates, and especially by the slender setiform spinules of all the plates on the under side.
Odontaster robustus Ver., sp. nov.
Puate XXIX. Ficures 4, 4a.
Form broadly stellate, with short, rapidly tapered rays and thick margins. The sides are regularly incurved. Radii about 1:15.
The marginal plates are larger and thicker than in the two preced- ing species. Those of the two series correspond closely in size and position. There are 27 of each series on each side of the body, in the type. They encroach considerably upon the disk, both above and below, and rise distinctly above the abactinal plates, thus form- ing a conspicuous margin. They are transversely oblong, about twice as high as broad. About four pairs of the distal dorsal plates are in contact medially.
The odd interradial plates are small and wedge-shaped, and do not reach the marginal sutural groove, but in this groove, opposite the odd interradials, there may be a small, odd, ovate plate. This is lacking, or very small, on two of the margins of the type. The sutural grooves are narrow and deep, with marginal fascioles of small slender spinules. Both series of marginal plates are thickly covered with small, slender, setiform spinules, those of the lower series somewhat larger and longer than those of the upper ones.
The abactinal plates are small, round, well separated, paxilliform. Those of the radial areas have a rather high column, somewhat capi- tate, with the top somewhat convex and covered with a divergent cluster of small, slender, acute, setiform spinules.
The papular pores are conspicuous and occupy five large radial and a disconnected central area; those in the central parts of each area are much larger than those at the edges; about six surround each plate.
The actinal plates are squarish and form about four rows parallel with the ambulacra; they are separated by rather wide grooves, and each bears a thick group of elongated, slender, setiform spinules.
Trans. Conn. Acap., Vou. X. Avcust, 1899,
14
210 A. E. Verrill—Revision Genera and Species of Starfishes.
The adambulacral plates are rather narrower than the adjacent actinals. Each bears four or five slender spinules in the furrow series; these are terete and rather larger than the actinal spinules ; on the actinal side there is a group of 12 to 16 slender spinules, those next the furrow series about the same as the latter in size and form; the outer ones are rather smaller.
The dentary plates are separated by wide open sutures; their mar- ginal and surface spinules are like those of the actinal plates.
The recurved spines are conspicuous and not much compressed, with regularly tapered, very acute, hyaline tips.
The type specimen has the greater radii 33 to 35™™; lesser, about 74, seo :
The type (No. 9758) was taken in 1881, by the Albatross, off Martha’s Vineyard, at station 994, in 368 fathoms, mud. A smaller specimen (No. 18423) was taken in 1885, at station 2586, in 328 fathoms, in the same region. ;
This species is easily distinguished from the two preceding by the thicker margin and disk, shorter rays, larger and fewer marginal plates, more numerous adambulacral spinules, ete.
Family PLUTONASTERIDZ.
Plutonasterince (sub-family) Sladen, op. cit., pp. 2, 60, 1889. Perrier, Exp. Tray. et Talism., p. 251, 1894.
This group appears to be sufficiently distinct to be regarded as a family. The great group called the family Archasteridw by Sladen and by Perrier is so heterogeneous that it cannot be definitely de- fined, as already explained by me. (See p. 199.)
In the present group the form is stellate, the rays often long and tapered. The abactinal plates are usually very numerous, in the form of columnar parapaxille or protopaxille, covered with small divergent spinules. They generally have no very definite arrange- ment and the median radial series is often not distinguishable.
The marginal plates are generally well developed and paired, but are sometimes small. They often bear one or more acute spines.
The actinal plates are imbricated and generally form rows run- ning from the ambulacral to the marginal plates.
The pedicellariz, when present, are usually of simple structure, with two to four papilliform blades. See pl. xxvu, fig. 6. They seem to be lacking in many species. Supra-ambulacral plates are present.
Nearly all the species of this group are from the deep sea. None are littoral. For the subdivisions see p. 200.
A, FE. Verrili— Revision Genera and Species of Starfishes. 211
Plutonaster Agassizii Verrill.
Archaster Agassizii Verrill, Amer. Journal Sci., vol. xx, p. 403, 1880. Plutonaster rigidus Sladen, op. cit., p. 91, pl. xiv, figs. 3, 4; pl. xv, figs. 3, 4, 1889 ; also var. semiarmatus, op. cit., p. 94. Plutonaster bifrons (pars) Sladen, op. cit., p. 88, 1889 (very young example). Plutonaster Agassizii Verrill, Proc. Nat. Mus., vol. xvii, p. 248, 1894. Amer. Journ. Sci., xlix, p. 151, 1895. PuaTE XXVII. Figure 6.
This species is intimately related to P. bifrons aud other forms that have been described from the East Atlantic. Probably several of these species will have to be united eventually. It is also very closely allied. to P. intermedius (Per.) of the West Indies, with which I have compared it. From the latter it appears, however, to be distinct.
Our specimens occasionally have one or two small pedicellariz on the actinal side near the jaws (see pl. xxvu, fig. 6). They have three or four simple papilliform blades, and are very similar to the ordinary form found on Dytaster.
Perrier and Sladen both state that no pedicellariz are found in this genus.
Four-rayed and six-rayed specimens occasionally occur. ,
Taken at 103 stations between N. lat. 41° 53’ and 35° 45/ 23"; in 182 to 1700 fathoms. Most common in 300 to 1200 fathoms.
Plutonaster efflorescens Perrier.
Archaster efflorescens Perrier, Etoiles de Mer, p. 255, 1887.
Plutonaster efflorescens Perrier, Trav. et Talism., p. 322, 1894.
The type of this species from station 29, 955 fathoms, Blake Ex- pedition, when examined by me in 1896, was in a very poor state of preservation ; all the rays but one were broken off and the granules of the disk were largely rubbed off.
It is a young specimen. The radii are only 17"™ and 5°5™™, so that the adult specific characters are not developed.
The rays are relatively large for so young a specimen, slender, tapered, and narrow distally, being more like those of Dytaster than Plutonaster. ‘
Marginal plates are 34 to 36 on each side of the body, small, squarish, not oblique ; the upper ones extend a little on the actinal side of the disk. They are covered with minute raised spinules, which are not crowded; along the interradial margins each usually bears a slender tapered, conical spine, about as long as the breadth of the plate. The marginal plates are convex with a distinct groove between them.
212 A. B Verrill— Revision Genera and Species of Starfishes.
The lower marginal plates extend much farther back from the margin than the dorsal ones, but they have the same fine spinulation; there are distinct marginal fascioles between them ; most of the distal and some of the proximal plates also have a central spine, like those of the dorsal series, but rather longer.
The apical plate is relatively large, elongated, ovate, with a large proximal notch.
The abactinal paxille are numerous, very small, round, nearly uni- form in size; when rubbed the plates are convex and elevated, well separated; three or four rows continue even to the apical plate. There is no distinct median dorsal series; each plate bears a group of four to eight (usually six) very small, slightly elongated, divergent spinules, forming regular stellate clusters, without any evident larger central spine.
The actinal plates are rather numerous, forming triangular areas; they are similar to the abactinal plates, but rather larger; each usually bears six to eight small, rough, divergent spinules, in a stellate group. They extend to about opposite the fourth or fifth adambulacral plates, and do not form evident radial rows.
The adambulacral plates are relatively large ; the inner or furrow margin is convex and along the middle portion of the groove of each plate four or five slender, elongated, rough, terete furrow-spines on its convex edge, and six to eight shorter and smaller, divergent spinules on its actinal surface, forming two or three irregular trans- verse rows, or else an irregular roundish cluster.
The dentary plates are rather prominent, sub-carinate, and are covered with numerous small, slender spinules on the actinal surface; on the furrow margin there aré numerous small slender spinules and about six larger, convergent, apical ones. No pedicellariz could be found.
Although this is evidently the young of some large species, it differs decidedly from the young of Dytaster insignis and D. gran- dis, of the same size, with which I have compared it directly. It has shorter rays; single marginal spines; much smaller and more finely spinulose dorsal paxillee ; more numerous actinal plates, with finer and more numerous spinules; more numerous spinules on the dentary plates; more nearly equal and regular and more slender furrow spines on the adambulacral plates; the edge of the latter is less prominent, so that the furrow-series is less broken. In these characters it agrees better with Plutonaster than with Dytaster.
It should probably be referred to Plutonaster, as has been done by Perrier (1894), but it does not agree with the young of either of the adult forms known from the American coasts.
A, E. Verrili— Revision Genera and Species of Starfishes. 213
Perrier (1894) seems inclined to unite this with pulcher (Per.), though he points out a number of differences. The latter is also quite young. Their identity seems to me very doubtful, after a comparison of the types.
Family GONIOPECTINIDZ Ver., nov.
Stellate with elongated rays; marginal, adambulacral and actinal plates bordered with peculiar pectinate spinules united by a web- like membrane, and thus forming specialized, continuous fascioles. Surface of the marginal plates usually smooth or with a few scat- tered granules, sometimes entirely granulated, usually covered with a thin membrane. .
Marginal plates large, regularly paired; the sutures corresponding above and below ; sometimes they are spinose. There may be an odd interradial marginal plate in each series (Prionaster).
Abactinal plates are paxilliform or columnar and covered with small spinules. They are arranged in oblique transverse rows on the rays. Actinal plates form radial series, usually double (single in Craspidaster), running from the adambulacral to the marginal plates, with deep fasciolated grooves between them, continuous with the fasciolated grooves between the marginal plates.
The adambulacral plates project over the ambulacral furrows, forming constrictions; they bear a curved or angular series of fur- row spinules united by a basal web.
The jaws are rather large and very prominent, with an open suture. They bear two or more enlarged apical spines, and more or less numerous smaller spinules on the actinal side.
Craspidaster Sladen appears to belong to this family.
Goniopecten demonstrans Perrier. Goniopecten demonstrans Per., 1881, p. 24. Etoiles de Mer, p. 249, pl. iv, fig. 5, 1884. Exp. Trav. et Talism., p. 295, 1894. Puate XXVIII. Figure 5.
The genus Goniopecten, as originally defined by Perrier, included Plutonaster and other forms now regarded as very distinct genera.
But later (1894, p. 294) he restricted the genus to the single type, G. demonstrans. This was also done by me independently, in 1894 (p. 249).
The genus is very peculiar in appearance, owing to the smooth plates and curiously fasciolated sutural grooves.
214 A. E. Verrilim Revision Genera and Species of Starfishes.
It is, however, very much like the new genus Prionaster in appear- ance, but it has a fasciolated sutural furrow running from the suture of the jaws to the suture between the first pair of marginal plates.
It has no odd interradial marginal plates, which are present in Prionaster. The dorsal marginal plates do not bear spines as in | the latter. It has a large madreporic plate with fine radial gyri.
The dorsal nephridial pore is surrounded by a large number of very small paxillee, which form a low central prominence (probably much more elevated in the young). The papular pores are numer- ous, small, arranged regularly, about six around each paxilla, over
-most of the disk and on wide basal radial areas, but even in the basal regions they are lacking along the three or four median radial rows of paxillze and do not reach the ends of the rays. The paxille are very numerous on the disk, smaller centrally ; they are high, with rounded or elliptical tops, and covered with a group of very small, short, blunt spinules, of which one to three are central. Distally the paxille become narrow-elliptical and very small. On base of the rays they form obliquely transverse rows. The marginal plates are mostly smooth except around thé mar- gins, and covered with thin membrane; distally on the rays they bear minute scattered granules. Around their margins there is a regular rim, formed by the regular row of webbed spinules, which project over the edges of the sutural grooves. In these grooves there are several rows of much finer slender spinules. The apical plate is rather large, obconic, unarmed. 2 The actinal plates are large, angular, of various forms, not numer- ous ; they extend out to about the 12th adambulacral plate, there being but a single row of small ones beyond the 7th. They form, proximally, double series, the two united rows corresponding to each marginal plate, but with from 14 to 24 adambulacrals. The actinal plates are flat and most of them bear from 1 to 3 minute’ scattered spinules, besides the marginal fasciolated row. ;
The adambulacral plates proximally have an oblique, angular fur- row-series of 9 or 10 slender, divergent spines, webbed together at base. Farther out, about the middle of the ray, the series becomes more regularly convex and more prominent, with 10 to 12 smaller and more slender spinules, which sometimes, in dry specimens, nearly meet across the furrow, leaving large roundish or elliptical spaces for the passage of the adambulacral feet, which are very large and tapered, but without any sucker. The actinal margin of the adain- bulacral plates has one or more series of small, stout, divergent spinules, webbed together and fasciolated.
A. EF. Verrili— Revision Genera and Species of Starfishes. 215
The jaws are very prominent, elliptical in outline. There are two or four tapered apical spines, much larger than the rest; the furrow- series is convex and contains 9 or 10 small spinules, like those of the adambulacrals. The elevated actinal surface is covered with many small, acute, spaced spinules, in three or four irregular rows.
No pedicellariz could be found.
A large specimen has the larger radius, 115™™; lesser, 21™™.
Taken by the Blake Exped. in 358 fathoms, off Santa Cruz, etc., and by the Albatross in the West Indies and Gulf of Mexico, at several localities, in 335 to 347 fathoms.
I have compared the Albatross specimens with the types of Perrier from the Blake Expedition.
Prionaster Ver., gen. nov. Stellate with long, tapered, squarish rays, high at base. Abactinal paxille are arranged on the rays, in obliquely trans- “verse rows, about four rows to each marginal plate ; they are small, high, rounded or elliptical, with a terminal cluster of small spinules. . Marginal plates large, high, not encroaching much on the disk, those on the interradial regions much higher than those of the rays. There is an odd interradial plate above and below, similar to the others in size and shape. The upper and lower plates correspond accurately even to the end of the ray. The proximal upper ones mostly have a central, acute, movable spine near the upper end. The lower ones may also bear a small spine. All are margined by a very regular series of small pectinate spinules, webbed together. Some of them may bear groups of minute granules.
The actinal plates are not numerous, flat, covered with thin mem- brane and with a few small scattered spinules, and a marginal series webbed together, so as to form fascioles. They are arranged in ‘double series ; the series are separated by the fascioles, but the two rows of a series are not. An odd interradial series, with two rows of plates, runs from the jaw and first adambulacral plates to the odd interradial marginal plate. (See Plate xxvu, figure 4a.) The other series correspond each to a marginal plate, but have no regular rela- tion to the adambulacrals.
The jaws are very prominent, with a large sutural groove. They have each two large apical spines and a row of sutural spinules. The adambulacral plates are large and project far over the furrow, so that the spines meet across it. They have a furrow series of numerous small spinules, webbed together ; their lateral and outer margins have smaller webbed spinules.
216 A. EL Verrill—R—evision Genera and Species of Starfishes.
Prionaster elegans Ver., sp. noy. PLATE XXVII. Ficures 4, 4a, 4b, 4c.
Disk small; sides high and vertical, evenly incurved ; rays high and nearly square at base, tapering regularly to the slender tips.
Radii as 1:5. Greater radius, 70" ; lesser, 14™™.
The marginal plates are oblong and much higher than long on the disk, but gradually become squarish on the rays. The upper and lower are exactly coincident, so that the vertical sutures are contin- uous. Their sides are nearly perpendicular and they encroach only a short distance on the disk, but at the middle of the rays each series is about as wide as the actinal area ; distally, near the tips of the rays, they are separated only by a single row of very small paxille. The distal plates bear groups of small spaced granules near the upper end. Each of the upper ones, except on the distal third of the rays, bears a small, movable, tapered, acute spine at the ~ Bs upper angle ; those at the base of the rays are longer than the rest. - Some of the lower marginal plates of the rays have a similar, but i smaller, spine at the lower angle and near the distal edge of the — plate ; most of the interradials have also a small cluster of minute granules near the lower end. All the marginal plates are bordered by a very regular and even series of small spinules webbed together to their tips. Those of the upper plates are much more numerous,
finer and closer, and evenly pectinate ; they nearly touch across the grooves. Those of the lower plates stand a little apart and are more divergent, about half as many in the same space as on the upper plates, and very similar to those between the actinal plates. The apical plate is large, prominent, oblong, with the inner end acute- angled.
The actinal plates are irregular in size and form, but mostly have ~ curved outlines ; they are partially concealed by a thin membrane, and many of them bear a very small subcentral spinule. All are bordered on that side next the fasciolated grooves by a row of appressed, slender, webbed spinules, which nearly or quite meet across the grooves; the latter are continuous with the grooves between the marginal plates and with those between and back of the adambulacrals. The actinal areas are not large and extend to about the eighth adambulacral. The median odd series consists of two closely united rows of about six each, the distal ones becoming very small. The next series contains a row of five plates and one of three similar plates; this series corresponds to the second and third adambulacrals. The next series, corresponding nearly with the
A. E. Verrill— Revision Genera and Species of Starfishes., 217
fourth adambulacral, has three plates in one series and two in the other. Beyond this the plates are few and irregular.
The adambulacral plates are broad and roundish, the proximal ones quite oblique ; their furrow edge projects over the furrow and bears a row of 10 to 12 small, slender, acute spinules, which are somewhat divergent and are webbed together for about half their length ; they meet or interlock across the furrow, leaving rounded or ovate open spaces between them for the passage of the large and tapered ambulacral feet. ‘These spinules become very small and slender dis- tally, but still meet across the furrow. On the outer and lateral margins of the plates there is also a series of divergent, webbed, fasciolated spinules like those of the actinal plates.
The jaws are oblong and very prominent on the actinal side ; each half has an actinal sutural row of very small spinules, and some additional ones on the surface. The furrow-series contains about ten spinules, increasing in length toward the apex of the jaw, where there are two much larger and longer, acute oral spines.
The madreporic plate is rather large, with fine gyri. The dorsal nephridial pore is situated in the center of alow elevation composed of very small, round paxille.
Taken by the Albatross at station 2401, in the Gulf of Mexico, in 142 fathoms. (No. 18,428.)
Family BENTHOPECTINIDZ Ver., nov. _Benthopectinine Ver., Proc. Nat. Mus., xvii, p. 245, 1894.
Benthopecten spinosus Verrill.
Benthopecten spinosus Verrill, Amer. Journal Sci., vol. xxviii, p. 218, 1884 ; Explorations made by the Albatross in 1883, in Annual Report, U.S. Comm. of Fish and Fisheries, pp. 519 [47], 548 [41], 1885. Proc. Nat. Mus., vol. Xvii, p. 245, 1894. Amer. Journ. Sci., xlix, p. 129, 1895.
Pararchaster semisquamatus var. occidentalis Sladen, Voyage of the Challen- ger, vol. xxx, p. 10, 1889.
Pararchaser armatus Sladen, op. cit., p. 19, pl. 1, figs. 5, 6; pl. 4, figs. 5, 6, 1889.
PLATE XXX. FiGurREs 7, 7a.
This species was taken north of Cape Hatteras, at 62 stations, between N. lat. 42° 47’ and 35° 10’, in 721 to 2021 fathoms, by the U.S. Fish Commission. Most common in 1200 to 1600 fathoms.
It was also taken in the Gulf of Mexico, station 2380 and station 2381, in 1430 and 1330 fathoms, and off Jamaica, station 2127, 1639 fathoms.
218 A, E. Verrili— Revision Genera and Species of Starfishes.
One specimen of this species (No. 15,570) is remarkable for having, on one of the interradial margins, a pair of plates in place of the usual odd median plate.
Figure 7, plate xxx, represents the abnormal segment of this spe-
cimen, with the jaw and corresponding pair of marginal plates —
(7m, m), and figure 7a represents a normal segment and jaw of the ©
same specimen, with the odd marginal plate, (m).
Family ASTROPECTINIDZ: Gray.
Blakiaster conicus Perrier.
Blakiaster conicus Per., 1881, p. 28. Etoiles de Mer, p. 265, pl. ix, fig. 2,.
1884. Leptoptychaster conicus Per., Exp. Trav. et Talism,, pp. 242, 248, 1894.
PLATE XXVIII. Ficure 7.
Perrier, in his later report, has united this genus with Leptopty- chaster, but it seems to me sufficiently distinct, though doubtless they are closely allied. In this genus the actinal plates are not arranged in distinct radial series, nor do they have such well devel- oped fascioles between them. On the contrary, they have a rather irregular, crowded, tesselated arrangement, the plates being round- ish or polygonal, pretty closely united, without deep, sutural, fas- ciolated furrows. The marginal plates, also, have only rudimentary fascioles. The jaws are stout and evenly convex, instead of thin and carinate. The dorsal paxille are larger, rounded, and more regular,
There is a distinct dorsal nephridial pore or “anus.” The dorsal papule are large, five or six around each plate, except on the distal half of the ray and on the small interradial areas.
The lower marginal plates have three or four larger and longer spines on the border.
There are also, on some of our specimens from off Havana, a.
number of pedicellariw. Those on the actinal and adambulacral plates have four to six convergent papilliform blades, similar to the surrounding spinules, but rather stouter and blunter (see pl. xxvu, fig. 7). Similar ones, but smaller, with three or four blades, occur on the marginal and abactinal plates.
West Indies and Gulf of Mexico, 92 to 175 fathoms.
A, EF. Verrill—Revision Genera and Species of Starfishes. 219
Sideriaster Ver., gen. noy.
Form broadly stellate with a very large disk ; dorsal surface con- vex, and capable of inflation, closely covered with uniform, stellate paxille. Upper marginal plates small, entirely lateral.
Interradial actinal areas are large, with numerous plates, the distal ones extending to the distal third of the rays. They are arranged in single radial series, each series usually corresponding to an adam- bulacral plate and most of them to a marginal plate, but some of the series are short and do not reach the margin, there being more adambulacral than marginal plates proximally, but distally they generally correspond in number, though there are sometimes, locally, two marginals to one adambulacral. These plates are covered with granules, and have divergent, fasciolated spinules along their radial margins, thus forming fasciolated grooves that are coincident with those between the marginal plates.
The abactinal paxille are large, closely arranged, and nearly uniform in size and shape, regularly stellate, with short, even spinules.
The madreporic plate is very large, round, flat, fully exposed, and has very numerous, thin, radiating gyri.
Papular pores are very numerous and are arranged singly, about six around each plate over the whole of the disk and rays, even close to the ends.
There is no distinct dorsal nephridial pore visible, nor do the cen- tral plates differ in size from those of the disk in general.
Marginal plates are small and not prominent. The upper ones are entirely confined to the margin, and are granulated, without spines. The lower ones form the lower part of the margin, but extend also on the disk below; they are spinules with a median row of larger spines.
The adambulacral plates have a prominent furrow angle, on which there is a large, median, odd compressed spine ; at each side of this there are, in the furrow-series, two or three erect flattened spines ; a stout spine occurs on the center of the actinal side, with a single or double row of shorter flat spines back of it.
The jaws are large, not very prominent, covered with numerous short, blunt spinules and having furrow spinules like those of the adambulacral plates.
This remarkable genus seems to be very distinct from all known forms, but clearly belongs to the Astropectinide. Its very broad convex disk and large actinal interradial areas and small marginal plates are exceptional ; and so is the very large madreporic plate.
220 A. EL Verrilli— Revision Genera and Species of Starfishes.
Sideriaster grandis Ver., sp. noy. PLaTE XXX. Ficures 8, 8a, 8b.
Large, regularly five-rayed, with a broad, somewhat inflated disk, regularly and broadly incurved at the sides. The rays are rather - long and rapidly tapered. Radii as 1:3.4. Greater radii, 133-138™™; lesser, 40™™,
The margin is formed mostly by the upper plates, which do not extend at all upon the upper side. They are small and short, those on the interradial margins shortest and highest, at least four times as high as long. They are covered with coarse rounded granules, and bordered with fascioles of slender spinules. The lower marginals are of the same length and extend on the under side considerably. They are covered closely with small appressed, flattened spinules, largest centrally, grading laterally to the marginal fasciolated spin-— ules. On the middle of each plate there is a vertical row of about four stout, tapered, more or less flattened, acute spines.
The abactinal paxillz are round and high, remarkably uniform in size, arranged on the rays in imperfect, transverse, oblique rows. They bear a round, rosette-like cluster of rather coarse, short, clavate or capitate, divergent spinules, of which one is usually central, with 6 or 7 in a circle around it, while about 15 to 18 form the marginal row, interlocking with those of the adjacent plates, so as to conceal the papular pores. The latter are rather large and regularly arranged over the whole disk and nearly to the ends of the rays, usually six around each paxilla. The bases of the paxille appear stellate.
The madreporic plate is remarkably large and flat or slightly con- cave, with very numerous and thin radiating gyri.
The actinal plates are granulated nearly like the upper marginal plates. Other under parts have been described above under the generic description.
Pedicellariz occur in small numbers on the adambulacral plates and on the first row of actinals. They have two or three short, stout, flattened, spinuliform blades, similar in size to the adjacent spinules.
One specimen (No. 10,877) was taken by the Albatross at station 2378, in Gulf of Mexico, in 68 fathoms.
A. E. Verrilli— Revision Genera and Species of Starfishes. 221
Family PTERASTERIDZ Per.
Hexaster obscurus Perrier. Hewxaster obscurus Per., Mem. Soc. Zool. France, iv, p. 267, 1891. Res. Camp. Sci., xi, p. 41, pl. iii, figs. 1, 1a, 1896. Pteraster (Temnaster) hexactis Verrill, Proc. Nat. Mus., vol. xvii, p. 175, 1894. Temnaster hexactis Verrill, Amer. Journ. Sci., xlix, p. 202, 1895.
There can be no doubt that the genus and species described by Perrier is the same as that described by me. Both were from the same region and nearly the same depth.
The original description by Perrier (1891) was overlooked by me when I described the species in 1894.
Only one specimen was taken by the Albatross, in 57 fathoms, at station 2433, N. lat. 43° 05’; W. long. 50° 43’, off Newfoundland.
It was taken by the Hirondelle, off Newfoundland, in 155 meters.
Hymenaster regalis Ver., 1894; var. Agassizii nov.
Large, swollen, polygonal, with short rays, and with concave in- terradial areas. The interradial margins are prolonged into a broad, soft web. Abactinal pore small, regular, five-angled.
Adambulacral spines three in a series, appearing rather stout and club-shaped at the tip, in the alcoholic preparation. The whole under side of the body is covered with a thick fleshy membrane.
The dorsal surface is covered with prominent, regularly arranged, well spaced, slender, acute spines.
Color, in alcohol: under side deep pink or rose-color; upper side somewhat paler pink. This species was taken by the Blake Expedi- tion, off Martha’s Vineyard, N. lat. 41° 24’ 45”, in 1242 fathoms, 1880.
Family ASTERINIDZ Gray. Marginaster austerus Ver., sp. nov.
Pentagonal with five short, triangular, subacute rays ; interradial margin incurved. Dorsal surface thickly covered with small and short, rough spinules, on crowded indistinct plates,
Papule rather large, solitary between the plates, generally diffused, even down to the margin; a regular row between the upper and lower marginal plates and just above the lower ones. Ten primary calicinal plates of the disk are larger than the rest and distinct from them ; the interradial most so. Dorsal nephridial pore distinct, sur- rounded by spinules.
222 A, EL Verrilli—Revision Genera and Species of Starfishes.
Upper marginal plates small, irregular in form and arrangement, scarcely distinct from the abactinal plates, except close to the end of the rays, and without marginal ciusters of spinules. Lower mar- ginal plates prominent, transversely oblong, depressed, the sharp outer edge bearing a regular horizontal row of four to six rough, blunt spinules, usually four or five-on the proximal and five or six on ~ the distal ones; on the upper side of the same plates there is a secondary row of the same number of much smaller and shorter spinules.
Actinal plates evident, irregular in size and form, the smaller dis- tal ones roundish; the more central ones elliptical, transversely elon- gated, and bearing about three crescentric rows of spinules; one to three bear a single, small, central spine in each area. In one speci- men there are two of these spines. The adambulacral plates, near the mouth, bear a single slender inner spine, on the edge of the groove, and two stouter ones, side by side, on the actinal surface; in the middle part of the groove the spines are placed obliquely, and distally the three spines gradually come to stand nearly in a single transverse row, and they also become longer and more crowded.
Taken in the West Indies by the Blake Expedition and by the Albatross, in fathoms.
Family STICHASTERIDZ Perrier, 1885.
Stichasteridce Sladen, 1889, p. 480. Perrier 1894, p. 128. 1896, pp. 25-27. Verrill, 1895, «p. 206.
.
Stephanasterias Verrill, 1871. Type, S. albula.
Stephanasterias Ver., Bull. Essex Inst., iii, p. 5, 1871. Expl. of Casco Bay, Proc. Am. Assoc. Adv. Sci. for 1873, pp. 356, 359, 364, 1874. Check List Invert., 1879. Expl. Albatross, 1883, p. 540, 1885.
Nemaster Perrier, Exp. Trav. et Talism., pp. 129, 181, 133, 1894. Camp. Scientif. ?Hirondelle, p. 27, 1896. :
Stichaster (pars) Verrill, 1866, p. 851. Perrier, p. 347, 1875. Sladen, p. 432, 1889.
Perrier, in adopting this generic division, proposed by me in 1871,_ changed the name to Nanaster, on the ground that Stephanasterias was preoccupied by Stephanaster Ayres. The latter name was well known to me when I used the former, but I regard the two names as perfectly distinct: the one being based on Asterias ; the other on Aster, The genus, if adopted, should therefore be called Stephanasterias. It is certainly very closely related to typical St?-
chaster.
A, E. Verrill— Revision Genera and Species of Starfishes, 223
The type species, S. a/bula, is common in moderately deep water (1-229 fathoms) from Greenland to Cape Hatteras. It has once been taken by the Albatross, in 1253 fathoms (station 2726, N. lat. 36° 34’), unless some mistake was made in the labelling. It has also been dredged by the U.S. Fish Commission off the coast of South Carolina and apparently in the West Indies. At least I have not yet been able to find any satisfactory characters for distinguishing the West Indian form S. gracilis (Per. as Asterias, 1884) from the northern one. It also has a wide range on the European coasts.
Note.
The specimens above described and discussed belong to several collections ;
I. The general collection of the Peabody Museum, Yale University, of which I have had personal charge for many yeats.
II. The Museum of Comparative Zoology of Harvard University, where I have had opportunities to examine especially the collections made in the West Indies by the U. S. Coast Survey steamer ‘‘ Blake,” during several expeditions under the supervision of Mr. Alexander Agassiz. This col- lection is of particular importance for it contains the types described by Perrier, from these expeditions. My thanks are due to Mr. Walter Faxon for his kindness in affording me facilities for this study.
Ill. The very extensive collections made by the U. S. Fish Commission, under my supervision, from 1871 to 1887, off the north-eastern coasts of North America. A large part of this collection is now in the U.S. National Museum, but a duplicate series is in the Yale Museum.
IV. A very interesting collection of deep-sea species dredged in the West Indies and Gulf of Mexico by the U. S. Fish Comm. steamer ‘‘ Albatross” in 1884-1886, and sent to me from the U. S. Nat. Museum for identification and study. This collection contains most of the new species described by Perrier, and some additional new forms.
V. Asmall but interesting collection made in the Bahamas and off Cuba by an expedition from the University of Iowa, and sent to me for study.
The two last named collections will be reported upon by me in detail in subsequent articles.
Much of the value of this article will be due to the unusually accurate enlarged drawings of the structural details of many of the genera and species discussed. These have all been made by my son, Mr. A. H. Verrill, and reproduced in facsimile by photolithography.
224. A. FE. Verrilli— Revision Genera and Species of Starfishes.
BIBLIOGRAPHY.
List of the principal works quoted.
The following partial list of works is intended to include, espe-
cially, those that are of importance in connection with the generic
synonymy and those that relate to the deep-sea species of the Atlantic, though others are included for various reasons. Most of the older descriptive works are omitted. For convenience, a special list of papers by the writer, relating to starfishes, has been added.
733.—Linck. De Stellis Marinis. Not binomial. 1836.—Agassiz, Louis. Prodrome d’une Monographie des Echinodermes. Mem. de la Societé des Sciences de Neufchatel, vol. i. 1889.—Forbes, Edw. Mem. Werner. Soc., vol. viii. 1840.—Gray, J. E. Synopsis of the Genera and Species of the class Hypostoma,
Annals and Magazine of Natural History, vol. vi, Nov. and Dec.,
1840.
1840.—Miiller and Troschel, in Wiegmann’s Archiv., vi, Bd. I.
1841.—Miiller and Troschel, Monatsb. Berlin Akad., 1841.
1841.—Forbes, Edw. British Starfishes, 8vo, with cuts.
1842.—Miiller and Troschel. System der Asteriden, 4to, 12 plates.
1847 (a).—Gray, J. E. Proceedings of the Zodlogical Soc. of London, Part xv.
1847 (b).—Gray, J. E. Annals and Magazine, Nat. Hist., vol. xx.
1857.—Stimpson, Wm. Crustacea and Echinodermata of the Pacific Shores of North America. Part I. Journ. Boston Soc. Nat. Hist., vol. vi, p. 444, 6 plates. (Starfishes are on pl. xxiii.)
1859.—Lutken, Chr. Bidrag til Kundskab. om de ved Kysterne af,Mellem-og Syd-Amerika levende Arter af Séstjerner. Vidensk. Meddel., Kjoben- hayn.
1859.—Mobius, K. Neue Seesterne des Hamburger und Kieler Museums, 4to, 4 pl., Hamburg.
1862.—Dujardin and Hupé. Echinodermes, in Suites a Buffon. (Largely a trans- lation of Miill. and Trosch., Syst. Aster.)
1866.—Gray, J. E. Synopsis of the species of Starfishes in the British Museum,
4to, 16 plates. London.
1865-1867.—Von Martens, E. Ueber Ostasiatische Seesterne. Arch. fur Na- turges., xxi, p. 345; xxii, p. 57; xxx, p. 106.
1869.—Perrier, Edmond. Recherches sur les Pédicellaires et les Ambulacres dés Astéries et des Oursins, Ann. Sci. Nat., Ser. V, vol. xii, p. 177,
1871.—Lutken, Chr. Fortsatte kritiske og beskrivende Bidrag til Kundskab om Sostjernerne (Asteriderne), III. Vidensk. Meddel. naturhist. Forening i Kjobenhavn, Nr. 15-19, 1871, 88 pages, 2 plates. No. I of this series was published in 1859 ; No. II, in 1864.
1875 (a).—Perrier, Edm. Classif. et la Synonym. des Stellerides. Comptes rendus, 1875, p. 127.
A, FE, Verrili— Revision Genera and Species of Starfishes. 225
1875 (b).—Perrier, Edmond. Revision des Stellerides du Museum. Archives de Zool. Expérim. et Gen., vol. iv, pp. 265-450.
1876.—Perrier, Edmond. Revis. des Stell. du Museum. Arch. Zodl. Expér. et Gen., vol. v, p. 1; p. 209.
1877.—Agassiz, Alexander. North American Starfishes, 4to, with fine plates.
; (Structural details.) Mem. Mus. Comp. Zodl., vol. v, No. 1.
1878.—Perrier, Edm. Etude sur la répartition Geographique des Astérides. Nouv. Archiv. du Mus., Ser. II, vol. i, pp. 1-108.
1878.—Viguier, C. Anat. Comp. du Squelette des Stellérides. Arch. Zool. Expér. et Gen., vol. vii, pp. 83-250, with plates.
1880.—Perrier, Edm. Comptes rendus, pp. 436-8. (Brief descr. of several new starfishes from the Blake Exped.)
1881 (a).—Perrier, Edm. The same, pp. 59-61. (Additional starfishes from the Blake Exped., briefly described.)
1881 (6).—Perrier, Edmond. Note préliminaire surles Etoiles de mer, rec. durant _les dragages du Blake. Bulletin Mus. Comp. Zodl., vol. ix. '1884.—Perrier, Edm. Memoire sur les Etoiles de mer rec. dans la mer des
Antilles et le Golfe du Mexique. Nouv. Archiv. du Museum d’Hist. Nat:, Ser. II, vol. vi.
1884.—Danielssen and Koren, Asteroidea, Den Norske Nordhays-Expd., 1876-1878, Zool., xi, 15 plates. Christiania. -
1885.—Perrier, Edm. Note prelim. sur les Echinod. dragués par le Travailleur et le Talisman. Ann. Sci. Nat., Paris, 1885.
1885.—Sladen, W. Percy. Rep. on the Scientific Results of the Voy. of H. M. S. Challenger, Narrative of the Cruise, vol. i, Part om.
1889.—Sladen, W. Percy. Report on the Scientific Results of the Voyage of H. M.S. Challenger, Zodlogy, vol. xxx, Part t1. Report on the Asteroidea.
1891 (a).—Perrier, Edm. Sur les Stellérides rec. dans le Golfe de Gascogne, aux Acores, et a Terre-Neuve, pendant les camp. sci. du yacht l’Hiron- delle. Comptes rendus, Acad. des Sciences, May, 1891.
1891 (6).—Perrier, Edm. Stellérides nouv. prov. des camp. du yacht l’Hirondelle. Mem. Soc. Zool. de France, vol. iv, p. 258.
1891 (c).—Perrier, Edm. Mission Sci. du Cap Horn, vol. vi, Zoélogie. Part mm Echinodermes.
1893.—Alcock, A. Natural History Notes from H. M. Indian Marine Survey Steamer Investigator. Ser. II. No. 7. An Account of the Collec- tion of Deep-sea Asteroidea. Ann. and Mag. Nat. Hist., Ser. 6, vol. xi. Feb., 1893. Three plates.
1894.—Perrier, Edm. Expéditions Scientif. du Travailleur et du Talisman, Part I, Echinodermes.
1895.—Leipoldt, F. Asteroidea der Vettor-Pisani Exped. Zeitsch. fur wissen- schaft. Zoél., lix, pp. 547-654. 2 plates.
1896.—Perrier, Edmond. Résultats des Campagnes Scientif. par Albert I, Prince
2 souv. de Monaco. Fas. xi. Contrib. a l’etude des Stellérides de VAtlant. Nord. 4to, iv plates. Monaco,
Trans. Conn. Acap., Vou. X. AvGust, 1899. 15
226 A. EL Verrill—Revision Genera and Species of Starfishes.
Partial list of papers by A. E. Verrill, relating to Starfishes, wholly or in part.
1866.—Polyps and Echinoderms of New England, with descr. of new species. Proc. Boston Soc. Nat. Hist., vol. x, pp. 333-357.
1867 (a).—Notes on the Echinoderms of Panama and the West Coast of America, with descriptions of New Genera and Species. Trans. Conn. Acad., i, pp. 251-322. pe
1867 (6).—On the Geographical distribution of the Echinoderms of the West Coast of America, and Comparison of the tropical Echinoderm Faunz of the East and West Coasts of America. Trans. Conn. Acad., i, pp. 323-339.
1868 (a).—Notice of the Corals and Echinoderms collected by Prof. C. F. Hartt, at the Abrolhos Reefs, Province of Bahia, Brazil, 1867. Trans. Conn. Acad., i, pp. 351-371, 1 pl..
1868 (b).—Notice of a collection of Echinoderms from La Paz, Lower California with descriptions of a new genus. Trans. Conn. Acad., i, pp. 371-376.
1868 (c).—Supplementary note on Echinoderms of the west coast of America.
Trans. Conn. Acad., i, p. 376.
1871 (a).—Descriptions of Starfishes and Ophiurans from the Atlantic coasts of - America and Africa. Amer. Jour. Science, vol. ii, pp. 130-133.
1871 (b).—Additional observations on Echinoderms, chiefly from the Pacific coast of America. Trans. Conn. Acad., i, pp. 568-593, 1 pl.
1871 (c).—On the Echinoderm-Fauna of the Gulf of California and Cape St. Lucas. Trans. Conn. Acad., i, pp. 593-596.
1871 (d).—Marine Fauna of Eastport, Maine. Bulletin Essex Inst., Salem, Mass., vol. ili, pp. 2-6. ;
1872.—Radiata from the Coast of North Carolina. Brief Cont. to Zoél., No. 22. Amer. Journ. Sci., vol. ili, p. 435.
1874 (a).—Report upon the Invertebrate Animals of Vineyard Sound and adjacent waters, with an account of the physical characters of the region, in Annual Report of the U. S. Commissioner of Fish and Fisheries, vol. i, for 1871 and 1872. Washington, D. C., 1874. (The Crustacea by S. I. Smith), pp. 295-478, 88 plates, and a map. (A separate edition with new pagination was published by the authors in 1874.)
1874 (b).—Explorations of Casco Bay by the U. S. Fish Commission in 1873. Proce. Amer. Assoc, for Advancement of Science, Portland Meeting, 1873, pp. 3840-395, 6 plates. ;
1876.—Note on some of the Starfishes of the New England Coast. Amer. Journ. Sci., xi, pp. 416-420 (Critical).
1878 (a).—Notice of Recent Additions to the Marine Fauna of the eastern coast of North America, No. 1, Brief Cont., No. 38, bis. Amer. Journ. Sci., xvi, p. 207, Sept., 1878.
1878 (b).—The same, No. 2. Amer. Jour. Sci., vol. xvi, p. 371.
1879 (a).—Notice of Recent Additions to the Marine Invertebrata of the North- eastern Coast of America, with descriptions of new Genera and Species, and critical remarks on others, Pt. I, Annelida, Gephyrza, Nemertina, Echinodermata, etc. Proc. U. S. Nat. Mus., vol. viii, pp. 165-206, (Starfish on pp. 201-208.) ;
A, E, Verrill—Revision Genera and Species of Starfishes. 227
1879 (b).—Anthozoa and Echinoderms (three new starfishes) in Report Prog. Geol. Surv. Canada, for 1878, by J. F. Whiteaves.
1879 (c).—Preliminary Check List of the Marine Invertebrata of the Atlantic Coast from Cape Cod to the Gulf of St. Lawrence, pp. 1-32. New Haven, 1879. Supplement i, 1881. Supplement ii, 1882.
1880.— Notice of the remarkable Marine Fauna occupying the outer banks off the Southern Coast of New England. No. 1. Brief Contr. to Zodlogy, No. xlvii. Amer. Jour. Science, vol. xx, pp. 390-403, (Several new starfishes. )
1882 (a).—Notice of the remarkable Marine Fauna occupying the outer banks off the Southern Coast of New England. No. 3. Brief Contr. to Zoology, No. xlviii. Amer. Jour. Science, vol. xxiii, pp. 185-142. The same, No. 4, pp. 216-225. The same, No. 7, vol. xxiv, p. 360, Nov.
1883.—Recent explorations in the region of the Gulf Stream, off the Eastern Coast of the United States, by the U. S. Fish Commission. Science, pp, 448-446.
1884 (a).—Notice of the remarkable Marine Fauna occupying the outer banks off the Southern Coast of New England. No.9. Brief Contrib. to Zoology No. lv. Amer. Jour. Science, vol. xxviii, pp. 213-220, Sept. (New Echinoderms and Anthozoa.)
1884 (b).—The same, No. 10, vol. xxviii, p. 8378. (Four new starfishes.)
1884 (c).—Notice of the remarkable Marine Fauna occupying the outer Banks off the Southern Coast of New England, and of some additions to the Fauna of Vineyard Sound. Ann. Report U. S. Comm. Fish and Fisheries, vol. x, p. 658.
1885 (a).—The same, No. 11. Brief Cont., No. lvii, vol. xxix, pp. 149-107. (Three new starfishes. )
1885 (b).—Result of the Explorations made by the steamer ‘‘ Albatross” off the Northern Coast of the United States, in 1883, in Annual Report U. S. Comm. Fish and Fisheries for 1883, pp. 503-699, 44 plates. Starfishes are on plates xiii to xix.
1894.—Descriptions of new species of Starfishes and Ophiurans, with a Revision of certain species formerly described. Proc. U. S. Nat. Mus., vol. xvii, pp. 245-297.
1895.—Distribution of the Echinoderms of Northeastern America. (Brief Cont. to Zodl., Nos. 58 and 59.) Amer. Jour. Sci., vol. xlix, pp. 127-141 ; pp. 197-212.
oN Tae ae
Acodontaster, 204. | Eugoniaster, 172. elongatus, 204. | investigatoris, 173. miliaris, 204.
Anthenea, 149. | Gnathaster, 201, 202, 203, 204, 205.
Antheniaster, 173. | elongatus, 208. sarissa, 174. Grayi, 205.
Antheneide, 200. meridionalis, 203, 204.
Anthenoides, 173. miliaris, 203. sarissa, 174. | pedicellaris, 208, 205.
Aphroditaster, 189. pilulatus, 205. gracilis, 195. Gnathasterine, 200, 201, 202.
Archaster Agassizii, 211. _Goniaster, 147, 148, 150. Bairdii, 181. Africanus, 156, 158. efflorescens, 211. | Americanus, 151. Parelii, 190. cuspidatus, 147.
Archasteride, 198, 201. granularis, 162.
Asterias granularis, 162. hispidus, 198.
Asterinidae, 221. Lamarckii, 157.
Asterodon, 203. semilunatus, 151. singularis, 203. _Goniasteride, 145, 200.
Astrogoniine, 187. _Goniasterinz, 200.
Astrogonium, 149, 150, 157, 189. Goniodiscus, 149. annectens, 195. cuspidatus, 149. Aphrodite, 195. pedicellaris, 182. australe, 161, 2354. Goniodiscine, 200. - fallax, 190. | | Goniodon, 203. gracile, 195. dilitatus, 203. granulare, 149, 162, 234. | | Goniopecten, 218. hystrix, 195. demonstrans, 213. Lamarckii, 157. intermedius, 213. necator, 195. subtilis, 196.
_Astropectinide, 201, 218. | Goniopectinide, 200, 201, 248.
Benthopecten, 201, 202. | Hexaster obscurus, 221. spinosus, 217. | Hippasteria, 148.
Benthopectinide, 200, 201, 217. Caribea, 174.
Benthopectinine, 200, 217. Europea, 148,
Blakiaster conicus, 218. * Magellanica, 175.
phrygiana, 148, 175.
Calliaster Childreni, 149. planus, 146.
Ceramaster, 161. | Hippasteriinz, 174, 200.
Cladaster, 175. Hoplaster, 197, 202. rudis, 176. | lepidus, 198.
Craspidaster, 213. | spinosus, 197.
Ctenodiscus corniculatus, 146. | Hosia flavescens, 149
Cribrella oculata, 146. spinulosa, 149.
Hymenaster Agassizii, 221.
Dorigona, 146, 184, 185. regalis, 221. arenata, 186.
Jacqueti, 186. Iconaster, 185. longimana, 185. Isaster, 178. prehensilis, 186. Bairdii, 181. Reevesii, 185.
subspinosa, 185. Lasiaster, 198. ternalis, 185. _ hispidus, 198.
Dytaster grandis, 212. Leptoptychaster conicus, 218. insignis, 212. 'Linckiide, 200.
A. E. Verrill— Revision Genera and Species of Starfishes.
Litonotaster, 171. intermedius, 172.
Luidiide, 201.
Ludiine, 201.
Marginaster austerus, 221. Mediaster, 177, 178. zequalis, 179. Agassizii, 181. arcuatus, 159, 183. Bairdii, 181. Japonicus, 159, 183. Patagonicus, 159, 184. pedicellaris, 182. roseus, 184, 196. stellatus, 181. Mediasterinz, 177, 200. Mimasterine, 200.
Nanaster, 222.
_ Nereidaster, 186. symbolicus, 187. bipunctus, 187.
Nymphaster, 177, 184, 199. albidus, 186. arenatus, 186. basilicus, 186. Jacqueti, 186. prehensilis, 186. protentus, 186. subspinosus, 185. symbolicus, 186. ternalis, 184, 185, 189.
Odontaster, 203, 204, 205. hispidus, 203, 205, 206. meridionalis, 203. pedicellaris, 203. robustus, 209. setosus, 207.
Odontasteridx, 200, 201.
Ogmaster capella, 185.
Paragonaster, 188, 189, 196. cylindratus, 196. elongatus, 196. formosus, 196. strictus, 196. subtilis, 196.
Pararchaster, 201. armatus, 217. occidentalis, 217. semisquamatus, 217.
Peltaster, 167, 168. hebes, 169. planus, 170.
_ Paxillosa, 199, 200, 201.
Pentaceros obtusangulus, 148, 234.
Pentacerotidx, 200.
Pentagonaster, 147, 148, 150, 157, 184.
abnormalis, 158, 234. affinis, 168. Alexandri, 197.
| |
|
|
| | | | /
229
Pentagonaster arcuatus, 183. arenatus, 186. balteatus, 162. Bourgeti, 158. capella, 185. concinnus, 162. dentatus, 159, 167. Deplasi, 161. Dubeni, 158. gibbosus, 159. granularis, 162. Gunnii, 158. hispidus, 198. intermedins, 159, 172. investigatoris, 173. Japonicus, 183. Lamarckii, 157. lepidus, 159, 198. longimanus, 185. Miilleri, 185. Patagonicus, 184. parvus, 153, 155. planus, 170. pulchellus, 158. ternalis, 185. subspinosus, 185.
Pentagonasteride, 145, 198, 199.
Pentagonasterine, 200.
Phaneraster, 150. semilunatus, 150.
Phanerozona, 200.
Plinthaster, 161. compta, 163. nitida, 165.
Plutonaster, 211. Agassizii, 211. bifrons, 211. efflorescens, 211. intermedius, 211. pulcher, 213. rigidus, 211. semiarmatus (var.), 211.
Plutonasteride, 200, 210.
Plutonasterine, 200, 210.
Pontasterinz, 200.
Porcellanasteridz, 201.
Prionaster, 215. elegans, 216.
Pseudarchaster, 189. annectens, 195. Aphrodite, 195. concinnus, 193. discus, 189, 195. fallax, 190. granuliferus, 192. hispidus, 191. hystrix, 195. insignis, 190. intermedius, 190. mosaicus, 196. necator, 195. ordinatus, 194. Patagonicus, 195.
230 A. EB. Verrill—Revision Genera and Species of Starfishes.
Pseudarchaster roseus, 184, 196. tessellatus, 195.
Pseudarchasterine, 187, 189, 200.
Pseudoreaster, 148. obtusangulus, 148. Pteraster hexactis, 221. Pterasteride, 202, 221. Pyrenaster, 166. affinis, 168. dentatus, 167.
Rosaster Alexandri, 189, 197.
Sideriaster, 219. grandis, 220. Stephanaster, 148, 157, 222. elegans, 157. Stephanasterias, 222. albula, 222, 223. gracilis, 223. Stichaster, 222. Stichasteride, 222.
Temnaster hexactis, 221. Tosia, 148, 158, 160. astrologorum, 161.
Tosia aurata, 161. australis, 148, 160. compta, 161, 163. Deplasi, 161. eximia, 161. grandis, 161. granularis, 161, 162. Greenei, 161. Grenadensis, 162. Gosselini, 162. heesitans, 162. magnifica, 161. mamuillata, 162. mirabilis, 161. nitida, 161, 166. Perrieri, 161, 165. placenta, 161. pulvinus, 162. rubra, 160. simplex, 161. tubercularis, 160. tuberculata, 161. Vincenti, 162.
Valvata, 199, 200.
A. E, Verrili— Revision Genera and Species of Starfishes. 231
EXPLANATION OF PLATES.
All the figures have been drawn from nature by Mr. A. H. Verrill except Plates xxiva and xxv, which are from photographs.
PLATE XXIV.
Figure 1.—Mediaster Bairdii Ver. Type. Dorsal side of the distal part of one of the rays with the granules removed. x8.
Figure 2.—The same specimen. Group of abactinal paxille from the base of a ray with the spinnles removed, showing the papular pores. x8.
Figure 3.—The same. Actinal side of aray. x4.
Figure 4.—The same. Abactinal side of the middle portion of aray. x3.
Figure 5.—The same. Group of adambulacral and adjacent actinal plates. x95.
Figure 6.—The same. Group of paxille and papular pores from the abactinal
side of the disk. Some of the paxille have pedicellarie. x12.
Figure 7.—The same from the ray, without pedicellarie. x8.
Figures 8, 9.—The same. Smaller abactinal paxille.
Figure 10.—Mediaster equalis Stimp. Actinal side of the basal part of a ray. x 6.
Figure 11.—The same specimen. Abactinal side of the basal partofaray. x6.
Figure 12.—The same specimen. Group of abactinal paxillz and papular pores from the base of a ray, with granules partly removed. x8.
PLATE XXIVa.
Figure 1.—Goniaster Americanus Ver. Original type, dorsal side. From a photograph. Somewhat reduced. Figure 2.—The same specimen. Ventral side. Somewhat reduced.
, PLATE XXV.
Figure 1.—Goniaster Africanus Ver. Original type. From a photograph. About natural size. . Figure 2.—The same specimen. Ventral side. About natural size.
PLATE: XX VI.
Figure 1.—Goniaster Americanus Ver. Original type. Group of plates of the abactinal radial areas at the base of a ray, showing the two kinds of plates, covered with granules, and the papular pores. The central plate has
* three spatulate pedicellarie. Much enlarged.
. Figure 2.—The same specimen. A single actinal plate having a pedicellaria (p) with curved blades. The blades have been removed to show the pits into which they fold when fully opened. Much enlarged.
232 A. FE. Verrill— Revision Genera and Species of Starfishes,
Figure 3.—The same species. Group of abactinal plates from the central part of the disk of a smaller specimen, showing several pedicellariz () in dif- ferent positions. x about 24.
Figure 4.—The same specimen. Dorsal side of the distal part of aray. +514.
Figure 5.—The same specimen. Actinal side of one of the jaws and adjacent parts. xdl¢,
Figure 6.—The same species, A young specimen (No. 18,459), like that described ~
as P. parvus by Perrier. Dorsal surface. x 7.
Figure 7.—Nymphaster ternalis (Per.). Dorsal side of a part of the disk and
base of aray. The granules have been partly removed from some of the marginal plates, and entirely from some of the abactinal ones. x91.
Figure 8.—Mediaster Bairdii Ver. Part of the abactinal system of plates from
the base of a ray and a part of the disk, seen from the inner side, showing the bases of the paxille with the radiating ossicles that connect them together, and the intervening papular pores. On the right are some of the interradial plates without papular pores between them. x 914.
Figure 8a.—The same. Abactinal paxille from the papular region.
RA XexeValn
Figure 1.—Tosia (Plinthaster) nitida Ver. Type. Dorsal view of a part of the disk andaray. x51.
Figure 1a.—The same specimen. Actinal side. x6.
Figure 1b.—The same specimen. Abactinal plates of a papular radial area. x 6.
Figure 2.—Tosia (Plinthaster) compta Ver. Type. Actinal side of the base of a ray. x51g.-
Figure 3.—Pyrenaster dentatus (Per.). Oral region of specimen No, 18,433. x 514.
Figure 3a.—The same. Dorsal surface of one of the rays of No. 18,483. Variety with wider abactinal radial areas. x51. ;
Figure 3b.—The same species. A younger specimen (No. 7079). Under side of a part of the disk showing the actinal and adambulacral plates. Two of the actinal plates have a bivalve pedicellaria. x91.
Figure 4.—Prionaster elegans Ver. Type. Under side of basal part of a ray. x51.
Figure 4a.—The same specimen. One of the jaws and adjacent actinal and adambulacral plates, showing the odd interradial series of actinal plates (m) and the first of the paired series on each side. x 51s.
Figure 4b.—The same specimen. Distal part of the upper side of a ray, with the apical plate. x 7.
Figure 4c.—The same. One of the dorsal paxille. Much enlarged.
Figure 5.—Goniopecten demonstrans Per. Under side of part of a ray, middle portion, showing adambulacral and marginal plates in contact. x51. Figure 6.—Plutonaster Agassizii Ver. One of the jaws and adjacent parts of a specimen having pedicellarie. One of these, with four blades, is situated
on the first actinal plate in line with the dentary suture. x 7.
Figure 7.—Blakiaster conicus Per. Part of the actinal surface, showing two
of the pedicellariz (p) with spiniform blades. x 14.
~
A. E. Verrill— Revision Genera and Species of Starfishes, 233
PuaTeE XXVIII.
Figure 1.—Hippasteria Caribea Ver. Type. Actinal side of a part of the disk and base of aray. x6.
Figure la.—The same. One of the adambulacral plates, more enlarged.
Figure 2.—Cladaster rudis Ver. Type. Dorsal side of a part of the disk and one of the rays. x6.
Figure 2a.—The same. Actinal side. x6.
Figure 2b, 2c.—The same. Profile views of one of the spatulate pedicellarie, much enlarged.
Figure 3.—Peltaster planus Ver. Original type. Dorsal side of the distal part ofaray. x9dl4.
Figure 3a.—The same specimen. A group of plates from the basal part of the ray ; c, c, median plates. x 51g.
Figure 4.—Peltaster hebes Ver. Type. Actinal side of the basal part of a ray. xd.
Figure 5.—Litonotaster intermedius (Per.) Ver. Dorsal side of aray. x6.
Figure 5a.—The same. Denuded group of plates from the small papular area ; c, ¢, median row.
Figure 5b.—The same. Actinal side of the base of the ray and part of disk. x 8.
PLATE X XIX.
Figure 1.—Odontaster setosus Ver. Under side of the base of a ray and part of the disk. x91.
Figure la.—The same. Under side of a portion of the disk with the spinules removed to show the plates ; (m) lower marginal plates; b, b, actinal; a, a, adambulacral plates. x 7.
Figure 1b.—The same. One of the jaws and adjacent parts, showing the recurved odd apical spine (s) of the jaw. 91s.
Figure 1c.—The same. A few paxillze from one of the basal radial areas of the upper side. x about 5.
Figure 2.—Odontaster setosus Ver. Part of the interradial area of the disk with three adjacent upper marginal plates. The middle plate (m) is the odd interradial. x 7.
Figure 3.—Odontaster hispidus Ver. Part of the abactinal and upper marginal plates (m) at the base of a ray. The larger spines (v) of the lower marginal plates are also visible. x 7.
Figure 8a.—The same. Adambulacral plates (a, a) and spines, and first row of actinal plates (b, b). The spines have been removed from most of the plates ; f, ambulacral furrow. x7.
Figure 4.—Odontaster robustus Ver. Type. Actinal side of the base of a ray. x 914.
Figure 4a.—The same. Abactinal side of aray. x 6.
234 A. E. Verrill—Revision Genera and Species of Starfishes.
PLATE XXX.
Figure 1.—Pseudarchaster intermedius Sla. Actinal side of a part of the disk andaray. x9dlg.
Figure la.—The same. One of the adambulacral plates more enlarged.
Figure 1b.—The same. Abactinal side of the basal part of a ray with most of the granules removed. x51, 3
Figure 2.—Pseudarchaster fallax (Per.). Abactinal side of a part of the disk and a ray, from which most of the granules have been removed. x51.
Figure 2a.—The same. One of the jaws and adjacent plates. x 51g.
Figure 2b,—The same. One of the abactinal paxille. Much enlarged.
Figure 3.—Pseudarchaster concinnus Ver. Type. Actinal side of the basal part of a ray, showing three pectinate fascioles (p). x51.
Figure 3a.—The same specimen. One of the jaws and the adjacent plates show- ing three pairs of pectinate fascioles. x 514,
Figure 36.—The same specimen. A group of abactinal paxille and papular pores from the basal radial region of a ray, including the median row (ce, c). x about 6.
Figure 4.—Pseudarchaster ordinatus Ver. Type. Actinal side of part of the — basal area of a ray and disk, showing four fascioles. x 51¢.
Figure 4a.—The same specimen. A jaw and the adjacent parts, showing seven pectinate fascioles (p).
Figure 4b.—The same specimen. A group of abactinal plates and papular pores» from the base of a ray, including the median row of plates (ce, c).
Figure 5.—Pseudarchaster hispidus Ver. Actinal side of the basal part of a ray- Sle
Figure 6.—Pseudarchaster granuliferus Ver. Type. Actinal side of a part of the disk and basal part of a ray, showing a single pectinate fasciole (p). Figure 6a.—The same specimen. <A group of abactinal paxille with granules removed. x6. a
Figure 7.—Benthopecten spinosus Ver. Abnormal specimen. One of the jaws (j) and interradial area, with lower marginal plates, showing a pair (m, m) in place of a single odd interradial plate. x 7.
Figure 7a.—The same specimen. A similar view of another jaw, having the normal odd interradial plate (m). x 7
Figure 8.—Sideriaster grandis Ver. Type. Actinal side of a part of the middle of aray. x4.
Figure 8a.—The same. One of the adambulacral plates more enlarged. x 7.
Figure 8b.—The same specimen. One of the abactinal paxille. x51.
ERRATA,
Page 146, foot note, for planus, read plana.
Page 148, line 24, for obtusangida, read obtusangulus. Page 149, line 21, for granularis read granulare. Page 158, line 4, for abnormale, read abnormalis. Page 161, line 7, for australis, read australe.
V.—On THE DEVELOPMENT OF THE PitipIUM OF CERTAIN NEMERTEANS. By Westry R. Cog.
As is well known, the embryology of the nemerteans presents two distinct types of development—the indirect and the direct. In the indirect type the cleavage of the egg results in the formation of a free-swimming larva of complex structure known as the pilidium. Invaginations of the body-walls of this pilidium give rise to a young nemertean possessing the form of the adult. In the direct type there is no such differentiated larva, the segmentation of the egg resulting directly in the formation of a worm resembling the parents.
Intermediate between these two extreme types is a third form where there is a comparatively thin external, ciliated skin, inside which the young worm attains a rapid development to the adult form. Such an intermediate form is as yet known only in a single species of Lineus (L. viridis = gesserensis) common both on the shores of New England and in Europe. These larve are known as Desor’s larve.
The direct type has been found to occur both in the Mesonemer- teans and in Metanemerteans, and from its wide distribution we may conclude that it is common toa much greater number of species than is the indirect type. In a portion of these, however, the outer ciliated layer of ectoderm is thrown off (as in many other Platyhelminths) after a second ciliated layer has formed beneath it. This occurs in Cephalothrix galathew, according to Dieck (11). Biirger (5) is of the opinion that this is likewise true in Prosorochmus. In other species the outer covering of the larva, as Salensky (27) found in Monopora, passes directly into that of the adult. In only a few species of nemerteans, however, have we any knowledge whatever in regard to the embryology. Of these the development is but very superficially known in all except a half dozen species. Salensky has described the direct development of Monopora vivipara (27). The same type of development has been studied by Birger (5) in Pros- orochmus, which is likewise a viviparous species. In addition to these Lebedinsky (20) has recently given a detailed account of the direct mode of development in Drepanophorus spectabilis and Tetras- temma vermiculus.
Among the nemerteans of New England I have noticed that the direct type of development occurs in Amphiporus ochraceus, A. virescens and Cephalothria linearis.
-
236 W. R. Coe—Development of the
The indirect type, in which there is a pilidium formed, is known in still fewer species, and these belong to the Heteronemerteans, in most of which cephalic slits are developed. Metschnikoff (25) found that this type occurs in Lineus lacteus of Europe. There have like- wise been several species of pilidium described from Europe, and three or more from America, but with one or two exceptions it has - never been known to what species of nemerteans they belonged. Joh. Miiller as early as 1854 (26) found that Pilidiwm gyrans gave rise to a young nemertean which he considered identical with Micrura fasciolata.
C. B. Wilson (32) has raised the young pilidium from the eggs of Cerebratulus lacteus. The young pilidium of this same species has been known to me for several years. The pilidia of Cerebratulus leidyt and of Micrura ceca* may easily be reared from the eggs during the months of July and August. It is from these two species, as well as from Cerebratulus marginatus, from Naples, that material | for the present paper has been obtained.
The deposition of the eggs and the early development of Lineus viridis were well described by Desor (10) in 1848 under the name of Nemertes obscura. The ciliated larva which he describes, and which represents an intermediate stage between the nemerteans with direct development and those in which a pilidium is formed, has since been known as Desor’s larva. The complete embryology of this species has now become well known through the researches of M. Schultze (29), V. Beneden (3), MacIntosh (22), Barrois (2), Hubrecht (16), and Arnold (1). Birger, in his splendid monograph on the nemer- teans (5), gives a clear summary of both the direct and indirect types of development.
On the development of the pilidium very little has yet been pub* lished, although several authors have described more or less fully the development of the young nemertean within the pilidium. Joh. Miller in 1847 gave a good description of a pilidium which he recog- nized as being a larval form of some animal, and which he named P. gyrans. In 1854 (26), after further investigation, he discovered that a young nemertean developed within the pilidium. This nemer- tean he supposed to be the young of M. fasciolata. Gegenbaur in 1854 (12) and Krohn in 1858 (19) discuss further the relations of the young nemertean to the pilidium. Leuckart and Pagenstecher (21) describe the structure of P. gyrans and P. auriculatum, and give brief notes in regard to the development of the young nemer-
* Verrill (31).
Pilidium of Certain Nemerteans. 237
tean. In 1869 Biitschli (6) published a further description of P. gyrans and gave a detailed account of the development of the nemertean. Salensky (28) has described very carefully the structure of the mature pilidium, and the manner in which the nemertean devel- ops from it. His observations on the structure of the pilidium include good descriptions of the mesenchyme, the muscular system and, more particularly, of the highly developed nervous system with which he finds the embryo to be endowed. The development of the nemertean from the pilidium is also described in detail and with great clearness by Biirger (4, 5). Wilson ($3), Fewkes, and others have published descriptions of several peculiar forms of pilidia. Verrill (30) also figures two or more species without definite descrip- tions. The development of the muscular system from the ameboid cells of the larval mesoderm has been described by C. B. Wilson (32) for Cerebratulus lacteus.
The only description of the development of the pilidium from the egg, however, is that given by Metschnikoff (24, 26) from Lineus (Micrura?) lacteus. The regular and equal cleavage of the egg gives rise to a typical blastula with a comparatively large segmentation cavity. The ventral cells (entoderm) soon become columnar, while those on the dorsal surface (ectoderm) remain more flattened. At the same time the cells of the whole blastula become covered with cilia, and a large flagellum appears at the upper pole. A regular invagination of the entoderm occurs after the separation from it of a few mesoderm cells. The cavity of the invaginating entoderm bends posteriorly to form the digestive canal. With the appearance of lappets the gastrula can be recognized as a pilidium. A series of infoldings of the walls of this pilidium gives rise to the body of the nemertean.
Fertilization. A detailed account of the maturation and fertilization of the eggs of C. marginatus has been published recently in the Zoologischen
Jahrbiichern (8). Soon after the egg comes in contact with the
water, a pair of very minute asters appear beside the nuclear mem- brane, as in other eggs. Their ultimate origin was not determined. A typical polar spindle is formed, and the sixteen ring-like chromo- somes divide, whether the egg has been fertilized or not. The process will go no further, however, until after fertilization, when the polar bodies are formed as usual. The first polar body occasionally divides into two. The egg is oriented even before deposition: for the asters
238 W. R. Coe—Development of the
of the maturation spindle appear on that side of the eccentrically- placed germinal vesicle which lies nearest the surface of the egg ;_ the polar bodies are formed on the same side; the spermatozoon usually enters at the opposite pole; and the first cleavage plane passes through the region of the polar bodies.
After the entrance of the spermatozoon its head thickens up, and. a delicate aster with distinct centrosome appears in its immediate vicinity. The centrosome, and later the aster, divides into two, with the formation of a delicate spindle. The spindle is soon ruptured and the two resulting asters may remain together near thé sperm- head; may separate widely from it and from each other; or one of them may wander off to a considerable distance, while the other remains in the immediate vicinity of the sperm-nucleus.
The centrosome remaining in the egg disappears after the forma- tion of the second polar body, as do both of the sperm-asters some- what later. The egg is thus left without visible centrosome, as’ commonly occurs in eggs of other animals. In this respect they agree most closely with Kostanecki and Wierzejski’s account of the mollusk Physa (18), and with Child’s description (7) of the process in the annelid Arenicola. The radiations of the sperm-asters, however, remain long after the disappearance of the centrosomes, and in most cases may be recognized even after the formation of the cleavage- — asters.
From the evidence obtained from a few eggs in which the sperm- centrosomes did not disappear as soon as usual, I am of the opinion that these centrosomes do not actually end their existence with the disappearance of the sperm-asters, and that they are identical with those which later appear in the pair of cleavage-asters.
The centrosomes of the cleavage-asters are surrounded by distinct centrospheres which increase enormously in size with a very slight increase in the size of the centrosomes. These centrospheres are not artefacts, for they may be seen in the living egg. The centrosomes divide very early. In the anaphase they separate somewhat, and about each a delicate aster is formed quite within the body of the centrosphere. These little asters extend their rays outward into the reticulum of the egg and eventually form the asters of the second cleavage, somewhat as Griffin has described in Thalassema (18). Here, as in the later stages, there is certain proof of the existence of the centrosome from one cell-generation to another. Its persist- ence does not necessarily indicate, however, that it has any right to claim for itself a place among the essential and permanent organs of the cell.
Pilidium of Certain Nemerteans. 239
Cleavage.
The eggs of many species of nemerteans will rarely be deposited in confinement, but will develop readily when artificially fertilized. On the southern coast of New England the eggs of Cerebratulus lacteus may be thus fertilized during the months of March and April; those of C. leidyi, Micrura ceca, Cephalothria linearis, Hunemertes carcin- ophila, and several species of Amphiporus and Tetrastemma during July and August. The eggs of Lineus socialis are mature in mid- winter, and those of L. viridis (gesserensis) from February to June. At Naples an abundance of the eggs of Cerebratulus marginatus may be obtained in March and April. Those of the first three species, C. lacteus, C. leidyi, M. ceca, as well as C. marginatus, develop readily to pilidium-forms which are characteristic of the different species, and which will live for two weeks or more in confinement. In none of these species, however, did I find it possible to keep the pilidia alive until the development of the young nemertean.
The eggs of CU. leidyi and M. ceca contain but little yolk, and are beautifully transparent. Those of C. dacteus and C. marginatus, on the other hand, contain a large quantity of yolk, are larger in size, and much less transparent. The pilidia, however, are equally trans- parent in all. The phenomena of cleavage and gastrulation are very similar in all four species, although the presence or absence of yolk modifies them to some extent.
It is, perhaps, advisable to follow the development in a single species and compare the others with this. Let us choose for this purpose Micrura cceca, the eggs of which furnish an almost ideal example of the regular, spiral type of cleavage.
The ripe egg of this nemertean measures about .09™™ in diameter. The vitelline membrane is extremely delicate, and can only be seen at the point where the polar bodies are formed. It is therefore very easily ruptured, and the polar bodies are lost by even a slight disturb- ance. The first cleavage takes place in the usual manner and divides the egg into two apparently equal cells (Fig. 1, Plate xxx1), Ata temperature of 20° C. this occurs about one hour and ten minutes after fertilization. The centrosomes of the first cleavage spindle appear very early, as stated above, and the asters which are to form the spindles of the second cleavage appear even before the comple- tion of the first cleavage (cf. 8). The first cleavage usually passes nearly through the point of extrusion of the polar bodies. A slight “ Zwischenkérper” is formed, but this soon disappears. The two blastomeres separate widely at first and remain connected only by a
240 W. R. Coe—Development of the
narrow bridge of protoplasm. Later they become closely pressed together (Fig. 1). 7
The spindles of the second cleavage occupy, as is usual, positions — nearly at right angles to that of the first, and the second division is — likewise vertical. The four resulting blastomeres are again almost — exactly equal in size. They are also apparently similar in regard to- their constituent protoplasm, for each of the four cells may be con- sidered as being made up of one quarter of the finely granular protoplasm which was collected at the animal pole during the pro-_ cess of fertilization, and of one quarter of the deutoplasm which — segregated towards the vegetal pole at the same time. There is no reason for believing, however, that the cells are actually alike as regards their ultimate constitution, for, as we shall see, it is proba- ble that one of the cells has quite a different sphere of activity than any of the others. The second cleavage does not take place exactly at right angles to the first, so that two of the cells lie at a slightly | higher level than the other two. This is one of the first indications of the spiral nature of the cleavage, which becomes so conspicuous in the later stages. In the Mollusk Crepidula Conklin (9) makes the very interesting observation that the obliquity of the cleavage is manifest even before the first cleavage.
Immediately after the completion of the second division the result- — ing blastomeres are almost perfectly spherical in form, so that they touch only on very small surfaces (Fig. 2, Pl. xxx1). Eight or ten minutes later, however, they have drawn so closely together that the surfaces of contact almost meet in the center, and leave only a very small segmentation cavity (Fig. 3). It sometimes happens that this cavity 1s entirely obliterated above, in which case the cells B and PY commonly come in direct contact and close up the space, while A and’C do not meet at all. The four cells form a nearly perfect square, one side of which is about equal to the diameter of the unsegmented egg.
When the spindles of the third cleavage are forming, each one is directed upward and somewhat to the right, showing that the third division is to be right-handed. After the separation of the eight blastomeres all are again of nearly spherical shape. The upper four are slightly, though Gisuinetly, paneer than the others,* and are more
*In CO. letdyi and in C. lacteus such is likewise the case. In Testy aster on the other hand, the upper four cells of the eight-celled stage are described by Lebedinsky (20) as being smaller than the lower four. In other nemerteans all of the eight cells are said to be of equal size.
Pilidium of Certain Nemerteans. 241
widely separated from each other. Indeed, they do not usually come even in contact at first (Fig. 4). Each lies above, slightly out- side, and to the right (following the hands of a watch) of its sister cell. It is interesting to note how these cells, at the time of their origin only indirectly connected, enter into the most intimate union in the later stages.
As the upper cells draw together they move so far to the right that they finally occupy positions almost, though not quite, in the intervals of the cells below (Fig. 6). This drawing together and rotation of the blastomeres of a segmenting egg is doubtless mainly due to the general physical property of surface tension. As stated by Wilson (34), the spiral type of cleavage “owes its peculiarities entirely to mechanical conditions, the blastomeres assuming the posi-
tion of greatest economy of space, precisely like soap-bubbles or other elastic bodies.” It is not unlikely that these purely physical conditions are aided by a slight mutual attraction exerted by the living protoplasm of the blastomeres.
Seen from the side (Fig. 5) the upper cells in front lie above and to the left of their sister cells. Even in the eight-celled stage there is a perceptible segmentation cavity in the center.
At the fourth cleavage the upper four cells of the eight-celled stage divide obliquely downward and to the right, while the lower four divide obliquely upward and to the left. This cleavage is there- fore typically left-handed. The sixteen cells become arranged in four zones (Fig. 7), the cells of each zone forming a nearly perfect square, and alternating with those above and below.
The fifth cleavage is right-handed, the sixth left-handed, and so on, conforming to the regular spiral type. In the 16-cell stage the segmentation cavity is more than equal in diameter to one of the blastomeres, and in the later stages increases rapidly in size. In the sixth cleavage some of the thirty-two cells divide earlier than others, so that we have several distinct stages between the 32- and the 64-celled stage.
Gastrulation.
About nine hours after fertilization cilia appear on the outer surface of all the cells, and the embryo begins to swim. With the appear- ‘ance of cilia a marked differentiation of the cells on the opposite sides of the blastula is noticeable, and the embryo rotates about a single axis. The rate of rotation is about two or three revolutions per second. The cells of the upper pole (corresponding with the side of Trans. Conn. Acap., Vou. X. Avuaust, 1899, 16
242 W. R&R. Coe—Development of the
the egg where the polar bodies were extruded) become more drawn
out and flattened (Fig. 1, Pl. xxx11), while those of the opposite pole — become more columnar in shape and extend far inward into the seg-—
mentation cavity. The blastula flattens out and the lower half begins a regular invagination,
Before invagination, however, a few cells which are to form the-
larval mesoderm are separated off from the endoderm cells and make
their way into the segmentation cavity (Fig. 1, Pl. xxx). The actual
origin of these cells is difficult to make out in WZ. ceca. In C. lacteus
and C. marginatus, where the blastomeres are not of so nearly the
same size, the origin of the mesoderm is more easily determined and will be discussed in a separate paragraph. Suflice it to say here that it appears to arise from two somewhat distinct, though closely con- nected, sources—from the divisions of a large, posterior pole cell, as in annelids, and from some of the endoderm cells. These latter seem
to be indiscriminately pushed inward and set free in the segmentation —
cavity. It is possible that this appearance is misleading and that all of the mesoderm is actually derived from a single polar cell. The derivatives of the polar mesoderm cell always lie primarily in the lower wall of the blastula between the ectoderm and the entoderm. After the cells enter the segmentation cavity they lie upon the ento-
derm, and after gastrulation arrange themselves around the mouth ~
of ‘the gastrula (Fig. 4, Pl. xxxtt).
As gastrulation proceeds the bilateral nature of the gastrula becomes very evident, not only by the arrangement of the super- ficial cells, but also by the marked backward inclination of the invaginating layer of entoderm (Fig. 1, Pl. xxxi1), In those species which contain but little yolk (JZ ceca, C. leidyi) the gastrulation is perfectly regular and the entoderm-cells retain nearly the same rela- tions with each other that they held in the blastula. Where yolk is abundant (C. lacteus, C. marginatus) the arrangement of the cells is broken up by the invaginating process, and they assume new posi- tions in the enteron.
Development of the Pilidium.
By the end of the first day the cells of the dorsal surface of the embryo become much more flattened and their boundaries much less evident. Most of the yolk-globules originally contained in the cells have meanwhile been absorbed and the ectoderm becomes almost perfectly transparent. At the time of the absorption of the yolk many of the cells contain one or more large, clear vacuoles which
-
C—O ee ee ee ee
Pe ee
sds
Pilidium of Certain Nemerteans. 243
later disappear. By the flattening of the cells the size of the embryo is greatly increased, measuring about .11™" in diameter, and the ectoderm is more widely separated from the enteron (Fig. 2, PI. xxx), At the same time the ectoderm at the two lower, lateral edges of the embryo extends downward to form the side-lobes, or lappets, of the pilidium. With the further growth of the pilidium the lappets become more or less highly developed, according to the peculiarities of the particular species to which it belongs (Figs. 5, 6, i xxxir).
The cells at the extreme upper pole of the embryo, instead of flattening out, become more crowded and columnar, and form the apical plate (Figs. 1, 2, Pl. xxxm1). This plate is at first covered with numerous long cilia which continue to increase in length and
gradually fuse together to form a few, very long flagella. As the pilidium grows older this fusion goes on until in many cases only a single large flagellum remains.
At the end of the first day the digestive tract (Fig. 2, Pl. xxxm) becomes differentiated into two distinct cavities—the esophagus and the intestine, the peculiarities of which will be described in detail below.
: The Pilidium of Micrura ceca.
At the age of six or eight days the pilidium of this species meas- ures about .14—.15™™ in length, and just about the same in a vertical direction from the apical plate to the bottom of the lappets. The pilidia usually swim near the surface of the water, with the flagellum directed upwards. A large proportion of all those contained in a vessel are usually collected in a single small space which may, or may not, be on the side of the vessel turned toward the light.
Like other pelagic embryos the pilidium is almost perfectly trans- parent. In general form it is balloon- or helmet-shaped with pecu- liarly short lappets (Fig. 6, Pl. xxx1). This pilidium differs from most other species in being of much greater diameter a short dis- tance below the apical plate than it is on a level with the mouth. Seen from above, the outline of the body is oval or elliptical, with the longer diameter directed antero-posteriorly. The dorsal and lateral surfaces (which make up the so-called umbrellar surface)
‘are formed from very thin five- or six-sided cells, each with a small, disk-shaped nucleus scarcely more than one-eighth the diameter of the cell. There are not usually more than sixty to eighty of these cells on the whole umbrellar surface exclusive of the lappets. Their
244 W. R. Coe—Development of the
boundaries (Fig. 6) can be made out only after treatment with suita- ble reagents.
In the lappets the cells become much thicker and are correspond- ingly more numerous. On the free edges of these organs they are closely pressed together and become highly columnar. All the ectoderm cells are covered with cilia, the length of which corre-. sponds more or less closely with the thickness of the cell on which they are borne. Thus the very thin cells of the dorsal surface have very short, scattered cilia, but as the columnar cells at the edges of the lappets are approached, the cilia become correspondingly longer and more closely packed together.
The apical plate is sharply marked, and is usually provided with a single flagellum nearly as long as the body. There are commonly several shorter and more slender flagella in addition to the primary one (Fig. 6, Pl. xxxir).
The mouth is highly distensible, and provided with thickened lips and buccal folds as described below. It is triangular in form and is situated slightly in front of the middle of the subumbrellar surface.
The esophagus is remarkably spacious and extends well upwards towards the apical plate. The intestine is comparatively small. It usually contains a quantity of small particles—probably food-ma- terial—which are kept in rapid motion by the long cilia lining the cavity.
The food of the pilidium probably consists of small pelagic larve, infusorians, diatoms, etc.—in short, of any minute organisms which ~ can be carried into the intestine by the ciliary motion.
On the sides of the body the network of highly refractive muscua- lar fibers with their clear nuclei are always conspicuous. They will be described in detail below.
Near the borders of the lappets and below the intestine a large number of mesenchyme cells are formed. Some of these are con- spicuous from their decidedly yellowish color. Clusters of such yel- lowish cells doubtless correspond to the ‘ golden-yellow spots around the margins” of the pilidium figured by Verrill (30, Fig. 6, Pl. 39). This is no evidence that the species are identical, however, for a brown or yellow color is found in such cells in several other species.
The pilidium of C. leidyi (Fig. 5, Pl. xxxi1) differs from that described above mainly in the greater extent to which the lappets are developed in the former. In general form it is broader at the base, and not so much swollen above as in M. ceca, The size. and general appearance otherwise are so closely similar that in certain
Pilidium of Certain Nemerteans. 245
states of contraction it would be difficult, especially in the earlier stages, to determine to which of the two species a given individual belonged.
The pilidium of (. /acteus is easily distinguished from the above by its larger size and much more highly developed lappets. A detailed description of this species is promised in a forthcoming paper by Prof. C. B. Wilson, who has already published a single figure of it (32).
The Pilidium of Cerebratulus marginatus.
At Naples, in 1896, the eggs of this species could be artificially fertilized during the latter part of March and throughout the month of April. At the end of seven days the pilidium (Fig. 5, Pl. xxxv)
‘is about .27™™ in length, and .21™™ from the apical plate to the lower edge of the moderately extended lappet. It is therefore nearly seven times as large as the pilidium of MZ. ceca or C. leidyi of the same age. The eggs themselves are also much larger and contain a large amount of yolk. Development is consequently much slower than in the two other species, the eggs of which contain but a small quantity of yolk. The general course of development, however, is similar in all.
The segmentation cavity in the blastula is smaller than in I. ceca and (©. leidyi. Instead of being flattened, the blastula is much higher than broad* The cells on the under surface become remarka- bly long and encroach greatly upon the segmentation cavity (Fig. 3, Pl. xxxur). Preparatory to invagination the entoderm cells become so closely pressed together at their lower ends that they gradually assume a conical or pear-shaped form, with their larger ends project- ing far into the segmentation cavity. As the process goes on some of them become pushed inward, separated entirely from the neigh- boring cells, and float freely in the fluid of the segmentation cavity. They thus become mesoderm-cells, and apparently supplement the few primary mesoderm-cells which seem to have an origin similar to that of the mesoderm in the annelids.
One of these primary mesoderm-cells is seen in Fig. 3, Pl. xxxmu, 7, in its original position at the lower, posterior border of the blastula. Three other mesoderm cells are shown in red above the entoderm. To the left of the primary mesoderm-cell are seen two very small
*The gastrula of this species is figured by Birger (5, Figs. 1, 2, Taf. 30) from sketches by Hubrecht.
246 W. R. Coe—Development of the
cells which apparently correspond to “ posterior entoblast” cells in” the annelid embryo. A further discussion of the origin of the mesoderm will be given below.
The process of invagination is somewhat different than in M. ceca. The pear-shaped endoderm cells near the middle of the lower surface become pressed further into the segmentation cavity so that a slight~ ventral depression is left (Fig. 4, Pl. xxxim). This marks the begin- ning of the invagination. The individual cells slip away from the outer surface, but remain closely in contact. As those in the center — pass inward they draw in their irregular, tapering, lower extremities | and arrange themselves about a central cavity—the archenteron. This cavity is very narrow and communicates with the exterior by a small, circular mouth (Fig. 4). As the process goes on it becomes more like the typical invagination described above for MW. cea.
The intestinal canal is early differentiated into two regions as in the two species above described, and is likewise provided with a simi- lar intestinal valve (Fig. 6, Pl. xxxi). The esophagus remains very narrow for several days, but later becomes much enlarged. It never develops, however, to the great extent characteristic for the two other species. Buccal folds are present as thickenings of the pos- terior esophagal walls. |
The muscular system develops as in the other species, and is — arranged in much the same way. The lateral bands to the lappets consist of more numerous fibers which are not so regularly arranged. The two bundles running from the apical plate to the anterior border of the esophagus are well marked in this species. '
During the formation of the gastrula the ectoderm becomes very thin, and the apical plate forms as in WM, ceca. There is usually but a single flagellum. This is nearly as long as the body.
The mesenchyme cells are several times more numerous and larger than in the other species, but have the same general arrangement (Figs. 3-6, Pl. xxxmr). Some of those which find their way into the lappets later acquire a reddish tinge. There is also a slightly red- dish tinge to the marginal cells of the lappets.
This may, perhaps, be looked upon as an indication that this pilid- ium possibly develops into the Pilidium gyrans of Miller. The time of the year when both may be found probably agrees. More- over the geographical distribution of P. gyyrans corresponds more or less closely with that of C. marginatus. The great abundance of C. marginatus, and the enormous number of eggs produced, makes the chances of obtaining pilidia of this particular species greater than is the case with almost any other species.
Pilidium of Certain Nemerteans. 247
The Digestive Tract.
At the end of twenty-four hours the wall of the enteron in JZ. ceca and C. leidyi becomes divided into two distinct regions—an anterior, buccal cavity, or esophagus, composed of flattened cells, and a posterior blind sack, or intestine, of columnar cells provided with very long cilia (Fig. 2, Pl. xxxm1; Fig. 1, Pl. xxxrv). Later cell- processes grow out to form a sort of intestinal valve which separates the two cavities completely. These processes (Fig. 3, Pl. xxxiv) are produced by the half-dozen cells which lie immediately between the two cavities. They are devoid of cilia, and closely resemble the blunt pseudopodia of an ameba. Ordinarily they meet in the center, so that the two cavities of the digestive tract are not in direct com- munication.
The cells of the valve do not actually fuse together, however, as may be seen by placing a little finely powdered carmine in the water with the embryos. The particles of the carmine are readily taken into the mouth and collected at the posterior end of the esophagus. Here they are churned round and round against the intestinal valve. When a considerable quantity has been collected, the pseudopod- like ends of the cells forming the valve separate, as in a true sphinc- ter, and the carmine passes into the intestine. There it is rapidly revolved for a time by the motions of the long cilia lining the cavity.
Hubrecht (16) and Arnold (1) find in Desor’s larva that at a cer- tain stage in the development the esophagus is for a long time completely cut off from communication with the intestine by a solid growth of cells. At a later period these cells separate, and communi- cation is again established. It seems probable that this mass of cells is represented in the pilidium in a rudimentary way by the ameboid cell-processes of the intestinal valve.
Eventually the particles of carmine, or of undigested food-material, are thrown out of the mouth, but not at the same region where they entered ; for these particles always enter at the anterior end of the mouth and leave at the posterior.
The mechanism by which the in-coming is separated from the out- going current consists of a thickened fold of large cells covered with especially strong cilia. This fold, which we may call the buccal ridge, extends forwards and downwards on each side (fig. 3, Pl. XXXIV; Fig. 1, Pl. xxxv, a) to the border of the mouth-opening, where it is continuous with the lateral lips.
These oblique folds, which are so greatly developed in M. ceca and C. leidyi, are represented in C. lacteus and marginatus only by slight thickenings of the posterior wall of the buccal cavity.
248 W. R. Coe—Development of the
During the first two days only a few particles of powdered car- — mine will be ingested, but after this time it is devoured in large quantities.
The mouth of the pilidium develops directly out of the gastrula- mouth, although the true blastopore is pushed inward and marks the opening of the esophagus into the intestine. In the early gas- trula the opening becomes oval, and this shape is retained in the pilidium,
On the sides of the mouth the epithelium of the esophagus is sepa- rated from that of the under surface of the pilidium by a pair of thickened folds, or lips (Fig. 2, Pl. xxx1v), covered with especially strong cilia. . These lips are united in front, but are widely separated posteriorly and become continuous with the pair of buccal ridges which reach inwards towards the intestine. The relation of these parts is well shown in Fig. 3, Pl. xxxiv. In this figure one of the lips — is seen between the lines converging at m, and represents the extent of the mouth proper. It is only between these lips that water or food-particles can enter the esophagus. The lip is continued posteriorly into the buccal ridge, a, which finally becomes lost in the wall of the esophagus. It is between the two buccal ridges that water with substances in suspension passes out from the esophagus to the exterior.
In Fig. 1, Pl. xxxiv, which represents an earlier stage, no such buceal folds are present, and consequently there is no definite cur- rent of water passing through the esophagus. At this stage very few, if any, particles of carmine are taken into the digestive canal.
When the buccal ridges have become established, a current of water carrying food-particles, etc. is constantly passing into the esophagus through the mouth proper, and out again between the posterior buccal ridges, In passing through the esophagus the solid material is collected by the cilia and passed into the intestine, as was described above.
The esophagus of the pilidium is found by Salensky (28), Butsebli (6) and others to develop directly into the esophagus of the adult nemertean. Likewise from the intestine the corresponding part of the adult némertean develops. Hubrecht (16) finds that this is likewise true of Desor’s larva. He also considers the esophagus to be composed of entoderm, and that the blastopore becomes the mouth of the nemertean. Arnold (1), on the other hand, believes that the esophagus is formed by an invagination of “secondary ” ectoderm, by which the blastopore is pushed inward and is repre-
Pilidium of Certain Nemerteans. 249
sented by the opening of the esophagus into the intestine. This becomes closed for a long time, as found by Hubrecht.
In the nemerteans with direct development, both the esophagus and the rectum are formed by special invaginations of the ectoderm, and are both widely separated from the blastopore.
The development of the intestinal tract in the pilidium, however, offers little positive evidence as to whether the esophagus is ectoder- mic in origin, because in this form the whole digestive tract (except the rectum) is formed by an almost continuous invagination. From a comparison with other forms, as well as from the histological pecu- liarities of the esophagus of the adult nemertean, it seems highly probable, as urged by Biirger in his splendid monograph, that this organ is here likewise of an ectodermic origin, A strong point in favor of this is the fact that the intestinal valve is indicated at a _ very early period. The histological differences likewise manifest themselves very early. As Biirger also states, such a supposition is necessary in order that the nephridial invaginations in different groups of nemerteans may not appear to originate in different layers of the body.
We may thus conclude that the process of invagination involves not only all of the entoderm cells on the lower pole of the blastula, but also some of the surrounding ectoderm cells by a continuous process of infolding. The blastopore would be pushed inward, and would be marked by the intestinal valve, or an homologous thicken- ing of the epithelium.
The whole digestive tract is capable of movement to a consider- able extent independent of that of the body-walls. In life the form of its parts, especially of the esophagus, is continually changing. This movement, which is sometimes vigorous, is accomplished by the contraction of the fibers of the few muscular cells which le directly beneath the epithelium of the digestive canal, and of other fibers which connect this canal with the other parts of the body.
Histologically the two portions of the digestive tract show marked differences in structure. Both in MW. ceca and in C. leidyi the esophagus is very large (Fig. 3, Pl. xxxiv) and is bordered by extremely thin epithelium. In the earlier stages of the pilidium the cells of the esophagus were as large as those of the intestine. The esophagus later increases to several times its original size with- out any increase whatever in the number of cells composing it, so that each cell must cover a large surface and becomes correspond- ingly thin. InC. marginatus (Fig. 6, Pl. xxx1m) and also in C. lacteus
250 W. R. Coe—Development of the
the esophagus is comparatively much smaller and its epithelium i is” correspondingly thicker.
The cilia on the esophagal cells are for the most part short and scattered. Near the intestine the cells become much thicker and the cilia longer and more numerous.
The cells making up the blind intestinal sack are always more or- less columnar in form, and all are provided with very long and strong cilia. By means of these any food-particles which may pass into the intestine are churned rapidly round and round. In MZ. ceca — and C,. letdyi the intestinal cells are more highly columnar in the earlier stages (Fig. 2, Pl. xxx) than in the later (Fig. 3, Pl. xxxry). In C. marginatus (Fig. 6, Pl. xxx) and in C. lacteus the cells are much more elongated, and their oval nuclei comparatively smaller than in the other two species.
In that portion of the cell which lies nearest the lumen of the — intestine are often found numerous small granules, and sometimes, — vacuoles of secretion (Fig. 1, Pl. xxxv). This secretion is doubtless aspecial digestive fluid which is poured out into the canal when food is present. In C. marginatus such granules or vacuoles often appear even before the pilidium is fully formed.
In many specimens we find that a few of the intestinal cells stain quite differently than most of the others. Two of these cells are ~ shown in Fig. 6, Pl. xxx. Their protoplasm is more granular and is also vacuolated. They have likewise a much greater affinity for stains. Both Salensky (28) and Biirger (5) describe such cells. The former suggests that they may be specialized cells of the nerv- ous system, while Biirger considers them to be differentiated gland-cells. There is little doubt that the latter view is correct, especially as the contents of the cells often resemble very closely those of some of the gland-cells in the intestine of the adult.
The Apical Plate.
In the course of gastrulation, when the cells of the upper surface of the body begin to flatten out, it is observed that a small cluster of cells at the very apex do not take part in the flattening process, but retain their columnar shape. These furnish the “ Anlage” of the future apical plate. The cells divide longitudinally and thereby become much smaller than the neighboring cells, although they remain as long or longer than at first. They thus become highly columnar, and are conspicuous in sections, because of their less gran- ular protoplasm, which shows a special affinity for stains (Fig. 4, Pl.
Pilidium of Certain Nemerteans. 251
xxxu). Their position is also marked by the depression, or circular pit, which eventually appears at this point. This pit seems to be well developed in all species of pilidia, In general it increases in depth with the increase in age of the individual. It is thus very much deeper in mature pilidia found free in the water than in the young specimens raised in the aquarium.
After the formation of the muscular system strong fibers pass directly from the apical plate to the anterior, lower borders of the body and to the esophagus. By the contraction of these muscles the pit is deepened, and the plate is withdrawn far below the general surface-of the body. The movement of the flagellum is probably also aided by these muscles.
The cilia are longer on the apical plate than elsewhere on the _ body, and some of them increase so much in size that they come to be recognized as flagella. The number of such flagella is at first considerable (Figs. 1, 2, Pl. xxxmz), but they rapidly fuse together to such an extent that only a few are present in the early pilidium (Fig. 1, Pl. xxx1v). These increase in length until they become half as long as the diameter of the body. Fusion goes on as the pilidium increases in size. At the age of three or four days the number has diminished to one or two long, whip-like flagella with several shorter ones (Figs. 5, 6, Pl. xxx; Figs. 2,3, Pl. xxxiv). It usually happens that these eventually fuse together, and the pilidium is provided with a single, thickened flagellum, the length of which is in M. ewea and C. leidyi about equal to that of the body.
When but a single flagellum is present its compound nature can be recognized at the point where it spreads out on the cells of the apical plate, for a considerable number of cells furnish it support. Biitschli (6) was the first to recognize that the flagellum was a con-* solidation of several more slender ones. Aceto-carmine often serves to separate the original constituents.
In C. marginatus there is usually but a single flagellum as early as the gastrular stage. In addition to this several rudimentary ones may be present (Fig. 6, Pl. xxxm1; Fig. 5, Pl. xxxv). Wilson figures (32) several flagella for the pilidium of C. /acteus. I think a single one, however, is much more common in all stages after the gastrula.
It occasionally happens that the flagella of the apical plate are arranged in two distinct clusters, as Wilson shows in his figure of C. lacteus. Likewise the apical plate itself may be divided into two, In a few instances two distinct and widely separated plates were present—one at the apex as usual, and the other about half way
252 W. R. Coe—Development of the
between this one and the lower border of the pilidium. Both were fully developed, and a distinct set of muscular fibers ran from each of them to the lappets. All such cases must, of course, be looked upon as abnormalities.
Salensky mentions (28) that some of the cells of the apical plate are continued internally into slender processes which he considers to ~ be nerve-fibers. J was quite unable to distinguish any such nerve- fibers from the numerous muscle-fibers which have their centers of attachment among the cells of the plate. The relations of the cells of the plate with these muscular fibers and the cells of which they are outgrowths are shown in Fig. 6, Pl. xxxv. As will be seen from the figure the -muscle-fibers do not come off directly from the plate- cells themselves, but rather they are fastened into the intercellular cementing substance. Whether Salensky mistook these fibers for nerve-fibers it is impossible to decide.
Both from its position and general appearance the apical plate of | the pilidium is naturally looked upon as homologous with that of the trochophore-larva of Annelids. In both groups this plate usually bears a tuft of long cilia or a single flagellum. From it muscles run similarly to the digestive tract and to the sides of the body. This organ in the trochophore, as is well known, represents the “ Anlage” of the superior esophagal ganglion of the adult. The sensory nature of this plate in the pilidium has, for these reasons, been generally conceded. On this account it is very disappointing to be unable to find (by the use of methylene blue, or otherwise) any trace of differ- entiated nervous structures in the young pilidia under consideration.
The Lappets.
At the end of the first day the ectoderm of the lower, lateral edges of the embryo becomes extended to form the side-lobes, or lappets, of the pilidium. These increase in size very slowly in JZ. cwea and C. leidyi, and at the end of two weeks are, in comparison with those of many other species, only slightly developed (Fig. 6, Pl. xxx11). It seems probable that this rudimentary condition of the lappets remains thoughout the life of the pilidium, since they are here decidedly longer than those described for the mature Pilidium auri- eulatum of Europe.
In (C. leidyi the lappets are developed to a greater extent than in M. ceca, as is seen from a comparison of Figs. 5 and 6, Pl. xxxu.*
* It should be noted that the magnification of the two figures in not the same. The pilidia of both species are of nearly equal size.
Pilidium of Certain Nemerteans. 253
In C. marginatus (Fig. 5, Pl. xxxv), these organs develop much more rapidly and to a much greater extent. Their size is still greater in C. dacteus, as may be seen from Wilson’s figure (32).
Their epithelial cells are thicker and correspondingly more numer- ous than over the rest of the body. The free edge of each lappet is thickened and covered with especially strong cilia.
The muscular system is more highly developed here than in any other, part of the body excepting that immediately surrounding the apical plate. In life the shape of the lobes is continually changing, and violent contractions are not infrequently observed. The dis- tribution of the muscular fibers will be described below. In the optical sections shown in Figs. 2 and 4, Pl. xxxry, the relations of the muscular cells and fibers to the epithelium are indicated.
Besides the cells and fibers of the muscular system the cavity of the lappets contains a considerable number of mesenchyme cells freely suspended in the slightly gelatinous fluid which fills the whole cavity of the body. These mesenchyme cells are irregular in form, sometimes showing ameboid movements, and are often conspicuous from their yellowish or brownish color.
The Nervous System.
Salensky (28) has described for the pilidium a highly developed and complex system of nerve-cells and nerve-fibers. The nervous structures develop, as does the muscular system, out of the amceboid cells of the larval mesenchyme. Both bipolar and multipolar cells are said to be present, and are provided with numerous, branching processes.
Salensky considers that the cells of the apical plate are highly sensory, and finds that many of them are continued internally into very fine nerve-fibers. These run in company with the muscular fibers to the anterior border of the esophagus and to the lappets. He likewise finds a band of nerve-cells and nerve-fibers, constituting what he calls a nerve-ring, reaching completely around the margins of the’pilidium and extending to the borders of the lappets. At the anterior, upper border of each lappet he finds a ganglionated swell- ing in the nerve-ring. This he considers the central organ of the nervous system. Finally, he considers that some of the cells lining the intestine belong to the category of nerve-cells.
These last-mentioned are obviously nothing but glandular cells which are partially filled with a deeply-staining secretion, as de-
254 W. R. Coe—Development of the
scribed above, and have no sensory function whatever. On the nature of the other nervous structures which Salensky describes the pilidia here recorded throw no light whatever. In no case was I able to demonstrate that any supposedly nervous elements did not actually belong to the muscular system. In this respect my results agree with those of others who have worked upon similar pelagic embryos of other groups.
The Larval Mesenchyme.
The mesodermic structures of the pilidium are of two sorts, but the cells from which they arise are at first indistinguishable from each other. They later become widely different and form the mus- cular system, described below, and the larval mesenchyme. The mesenchyme cells float freely about in the body-cavity and doubtless secrete a small quantity of transparent, gelatinous substance which, with a large amount of water, fills up the whole cavity.
The cells of the mesenchyme have already been well described by Metschnikoff (25), Salensky (28), C. B. Wilson (82) and others. It is from these cells that the musculature of the adult nemertean de- velops. Hubrecht (16) states that in Desor’s larva the adult nery- ous system is also derived from the larval mesenchyme. Biirger, on — the other hand, feels confident that such is not actually the case, but that in Desor’s larva, as in the pilidium, the nervous system arises directly from the ectoderm (8, p. 475). The development of the muscle-cells of the pilidium from the undifferentiated ameboid cells of the mesoderm has recently been described by C. B. Wilson.*
Some of the mesenchyme cells become collected in clusters near the edge of the lappets and below the intestine. These often con- tain a number of clear vacuoles the contents of which perhaps con- tribute to the formation of the transparent, jelly-like substance which fills the whole body. Others are equally conspicuous because of their decidedly yellowish color. They are, on the whole, not very different from the original mesoderm cells. They are oval or ame- boid in shape, and are much larger than the muscle-cells. They have little to do with the formation of the pilidium and are held in reserve until the body of the young nemertean begins to develop.
* It is doubtless due to a typographical error that Wilson (32, p. 20) states that ‘‘the mesenchyme first appears as isolated cells derived from the ectoderm, as ob- served by Metschnikoff.” Metschnikoff (25, p. 53) says that they almost certainly arise from the entoderm.
Pilidium of Certain Nemerteans. 255
The Muscular System.
In the living pilidium the muscles appear as clear, highly refrac- tive fibers, which are most commonly slightly curved. In optical cross-section each fiber appears as a bright dot. They run in a more or less irregular course from the muscle-cells to their attachment in the epithelium, or to an anastomosis with another fiber. When the embryo is killed they contract forcibly, thereby reducing greatly its size. This is in part accomplished by forcing water out of the digestive canal.
In contracting the fiber not only shortens in length, but also becomes wavy. and twisted upon itself, or coiled up in a spiral like the stem of a Vorticella. Fig. 3, Pl. xxxv, shows the fibers in a living pilidium, while Fig. 5, Pl. xxxiv represents similar fibers after
the embryo has been killed with corrosive sublimate.
The nuclei of the muscle-cells are mostly clear, oval bodies about .003"™" in length. Around the nucleus is a very small quantity of perfectly clear protoplasm, from which two, three, or more contrac- tile fibers extend in various directions.
The origin of the muscles from undifferentiated mesenchyme is easily foliowed because of the transparency of the embryos. The cells of the mesenchyme, as stated above, appear in the segmenta- tion cavity in the earliest stages of gastrulation. Some of them seem to originate directly from the entoderm, while others are derived from a pair of primary mesoderm cells set apart as such in the blas- tula. Never have I seen any indication that any of them came from the ectoderm, as Hubrecht (16) describes in Desor’s larva.*
At the time of their first appearance the mesenchyme cells are very large and few in number. During gastrulation they multiply rapidly, as may be seen from the many karyokinetiec figures which they contain. At the time of appearance of the lappets which dis- tinguish the pilidium from the gastrula their number is very consid- erable. Some of them become irregular in shape, as described by Metschnikoff (25), and as Wilson (32) has pointed out in (C. Jacteus. If watched carefully, some of these irregular cells will be seen to send out ameboid processes, and exhibit a great deal of indepen- dent motion. They not only change their shape, but actually move about in the cavity of the body by means of their pseudopod-like processes.
* Arnold (1) has recently reinvestigated the development of Desor’s larva, and finds that the mesoderm originates on both sides of the blastopore at the point where entoderm and ectoderm come together.
256 W. R&R. Coe—Development of the
They later collect in more or less constant numbers in certain definite localities, where they send out especially long processes and — attach themselves permanently to the epithelium. When they are © no longer freely movable in the segmentation cavity their processes grow out into long fibers. They are then recognized as muscle- elements. In the early pilidium we find a dozen or more such cells . immediately beneath the apical plate. Others fasten themselves firmly to the walls of the body and to the digestive tract, and a con- siderable number to the lower borders of the lappets.
Fibers now grow out in certain definite directions. At the end of two or three days, they have elongated and anastomosed with each other to such an extent that they form a practically continuous sys- tem of interdependent muscles. Some of these eventually become arranged in several definite muscular bands, while others form an irregular network connecting these bands, as Salensky (28), Biitschli (6), and Wilson ($2) have deseribed for other species. The mus-— culature in Df. ceca and C. leidyi consists (a) of two principal mus- cular bands extending from the apical plate to the two lappets ; (0) of a pair of bands extending longitudinally along the floor of the pilidium at the sides of the mouth ; (c) of a few fibers connecting the apical plate with the anterior border of the esophagus; (d) of a
few fibers with numerous branches lying on the anterior and poste-
rior faces of the embryo; (¢) of a few surrounding the digestive tract and serving to connect this organ with the body-walls.
Of these bands the lateral pair connecting the apical plate with the lappets is by far the most extensive. In close proximity to the crowded, columnar cells of the apical plate lie about six or eight muscle-cells on each side. From each of these cells proceed one or two long, slender, branched fibers which pass close beneath the body-epithelium downwards until they either anastomose with fibers from similar cells in the lappets or become attached directly among the epithelial cells at the lower margin of these lobes (Fig. 2, Pl. xxxiv; Fig. 3, Pl. xxxv). One or two other fibers from each cell pass among the columnar cells of the plate and serve to anchor the muscle-cell in place. From the dorsal surface the arrangement of these cells and fibers is shown in Fig. 2, Pl. xxxv. Those fibers running nearly parallel down the right and left sides constitute the lateral bands.
The muscle-cells in the lappets send similar processes toward the apical plate. These, in part, anastomose with the fibers from the apical plate. Thus there are from eight to sixteem fibers running
w=
Pilidium of Certain Nemerteans. 257
more or less parallel between the apical plate and the lower borders of the lappets. The cells in the lappets likewise send off longitudinal fibers along the floor of the lobe, as well as towards the borders of the mouth. This complex system of muscles enables the lappets to be moved readily in any direction.
Figure 3, Pl. xxxv, represents somewhat diagrammatically the arrangement of the muscle-cells and fibers which are seen in a side
view of a pilidium. The nearly parallel vertical fibers constitute
one of the lateral muscle-bands. The relation of the muscle-cells to the cells of the apical plate is shown in Fig. 6, Pl. xxxv. As seen in the figure, the muscular fibers are attached between the epithelial cells rather than directly to their faces. Likewise the arrangement of the muscle-cells and mesenchyme-cells in the lappets, and the
relation of the fibers to the epithelial cells is shown in Fig. 4, Pl.
XXXIV. On the anterior and posterior faces of the pilidium we find com- paratively few fibers, and these are much branched. They run
- mostly horizontally or obliquely instead of vertically, as on the
lateral faces. They anastomose freely with each other, with the fibers from the apical plate, and with others passing inward to the digestive canal. Those on the anterior face are represented in Fig. 4 bl, XXXV.
In the mature pilidium one of the largest and most conspicuous muscle-bundles is that running from the apical plate to each side of the anterior border of the esophagus and thence to the lappet. This muscle is well developed in C. marginatus, where it consists of several strong fibers. It is represented in C. leidyi and M. ceca merely by a very few-delicate and widely separated fibers. Two of these fibers are represented in Fig. 3, Pl. xxxiv. They serve both to hold the esophagus in place and to withdraw the apical plate. They doubtless also aid in rocking the plate and thus moving the flagellum.
From the region of the mouth a series of scattered muscular fibers runs radially in all directions. A few are fastened to the anterior and posterior margins of the body, but most of them pass into the lappets, at the edges of which they are attached. Besides these there is a horizontal bundle running longitudinally along the floor of the body-cavity on each side of the mouth. One of these muscle- bands is represented in dotted lines in Fig. 3, Pl. xxxv, (6). Their fibers are fastened at the anterior and posterior margins of the body, and anastomose freely with other fibers.
Trans. Conn. Acap., VoL. X. Aveust, 1899,
17
258 W. R. Coe—Development of the
Surrounding the whole alimentary canal and connected with the other muscular bands by frequent anastomoses is an irregular net- work of scattered fibers which provide this organ with independent motion and also serve to hold it in place.
SHEFFIELD BIOLOGICAL LABORATORY OF YALE UNIVERSITY.
EXPLANATION OF PLATES.
PLATE XXXII.
Karly cleavagé of the egg of Micrura ceca. Camera lucida drawings of living eggs at a magnification of about 300 diameters.
Figure 1.—Egg about one hour and forty minutes after fertilization. The fully formed spindles of the second cleavage are indicated by dotted lines, Above are the two polar bodies.
Figure 2.—The same egg eight minutes later. The spindles are still evident. The polar bodies lie between the blastomeres.
Figure 3.—The same egg after fifteen minutes more. The blastomeres have drawn very closely together, leaving but a minute cavity between them. 1, 1, denote the plane of the first cleavage; 2, 2, that of the second cleavage. A, B, C, D, the four quadrants of the egg. One of the asters of the third cleavage is seen in each blastomere.
Figure 4,—The third cleavage just completed. The upper four cells are widely sepa- rated. Each lies somewhat outside and to the right (following the hands of a watch) of its sister-cell.
Figure 5.—Eight-celled stage from the side. The first polar body lies above. 3, 3, denote the plane of the third cleavage.
Figure 6.—Eight-celled stage from above after the blastomeres have drawn together. Same position as fig. 3. Polar bodies are not shown.
Figure 7.—Sixteen-celled stage from above. In the same position as fig. 3.
Figure 8.—The same stage from the side.
PLATE XXXII.
The mesoderm cells are indicated in red.
Figure 1.—Vertical section of the blastula of Micrwra ceca, nine hours after fertiliza- tion of the egg. The two large cells (one of which is in the process of division) within the segmentation cavity are mesoderm cells. x 600.
Figure 2.—Horizontal section of the gastrula of JM, ceca, at a stage somewhat later than that shown in fig. 3. The section passes along a plane at a level with 2, 2, fig. 1, Pl. xxxmm. The circle of ciliated cells in the center represents a section of the entodermie sack, In the space between the entoderm and ectoderm are seen several cells of the larval mesoderm. These cells are now much smaller and more numerous than in fig. 1. x 600.
Pilidium of Certain Nemerteans. 259
Figure 3.—A gastrula of Jf ceca, seventeen hours after fertilization. The dark ring in the center indicates the blastopore. The cell-boundaries and the nuclei are drawn as accurately as possible from an egg fixed in picro-acetic, stained witb acidulated hematoxylin and mounted in oil of cloves. The anterior surface is towards the top of the page. Cilia are not indicated. x 600.
Figure 4.—Vertical section of the gastrula of C-. ledyi, seventeen hours after fertiliza- tion. The apical plate is indicated by darker stipple. The mesoderm-cells lie at the sides of the entodermic invagination. The segmentation cavity is smaller than normal owing to the shrinkage which took place during the process of im- bedding in paraffin. One of the endoderm cells is dividing vertically. x 500.
Figure 5.—A pilidium of C. leidyi at the age of ten days. The dotted lines indicate the position and thickness of the walls of the enteric cavity; 0, cavity of esopha- gus; 2, cavity of intestine. x 250.
Figure 6.—Pilidium of M. ceca at the eighth day. The nucleion the surface are those of the ectoderm cells, the boundaries of which are indicated by faint, dotted lines. The coarse, dotted line indicates the position of the enteron, as in fig. 5. The mouth is indicated by the heaviest dotted line (cf. Pl. xxxtv, fig. 3). Below the mouth is seen one of the very short lappets with its strong cilia and numer- ous nuclei. x 400.
PLATE XXXII.
Tn all the figures the cells of the larval mesoderm are indicated in red.
Figure 1.—Median, optical, sagittal section of gastrula of M. ceca fifteen hours after fertilization. The cells of the apical plate with their developing flagella are in- dicated by the darker stipple. The cilia, which cover the whole surface, are not shown. x600. Corrosive sublimate, acidulated hematoxylin.
Figure 2.—A similar section of the very early pilidium of WZ. ceca twenty-four hours after fertilization. Below the section is a line which indicates the extent to which the lappets have developed. The boundaries of the cells are merely indi- cated. Slightly diagrammatic. ~ 600.
Figure 3.—Median vertical section of blastula of C. marginatus. At the right of the elongated entoderm cells is a larger cell, m, in process of division. This cell appears to be homologous with one of the primary mesoblast cells of the annelid embryo. Immediately to the left of m are two minute cells which bear a strik- ing resemblance to the annelid “‘ posterior entoblast” cells. Picro-acetic. x 350.
Figure 4,—Early gastrula of C. marginatus. The cells of the apical plate are indi- eated by darker stipple. The blastopore is very narrow. The entoderm cells have not yet become arranged to form the intestinal canal. Picro-acetic. x 300.
Figure 5.—Median, sagittal section of late gastrula of C. marginatus. Picro-acetic. x 350.
Figure 6.—Median, sagittal section of the embryo of C. marginatus at the beginning of the pilidium-stage. The extent of the lappets is indicated by the line (with cilia) below the section. On the dorsal side the section must be considered as being of great thickness, for the drawimg represents not only a section of the apical plate, but also a surface view of half of the depression above it from which the flagellum issues. The two darker cells in the intestinal wall are gland-cells,
_ The intestinal sphincter (devoid of cilia) is partially contracted. Slightly dia- grammatic. Picro-acetic. x 350.
260 W. &. Coe—Development of the
PLATE XXXIV. ~
The mesodermic structures are printed in red.
Figure 1.—Optical, median, sagittal section of the early pilidium of C. leidyi, thirty- — six hours after fertilization. To the section the right lappet has been added, and likewise the right half of the apical depression. The nuclei and cell-boundaries x. have been drawn from an actual section. The mesoderm cells are arranging themselves in their definite positions. x 376.
Figure 2.—Optical, transverse section of the pilidium of Jf ceca at the age of ten days. The section passes through the mouth, esophagus and apical plate (cf. fig. 3). Below the esophagus are sections of the two horizontal lips (2) with — strong cilia bordering the mouth. The lappets are contracted, and therefore thicker than in life. Just inside the epithelium (both of the ectoderm and ento- derm) are drawn in red the cells and fibers of the larval musculature. The larger, dotted, red cells are those of the mesenchyme. The single, coarse flagel- lum is seen to be made up by the consolidation of several more slender flagella. The mesoderm cells are grouped mostly in the lappets, and just beneath the apical plate. x 500. .
Figure 3.—The left half of a pilidium of Jf ceca of ten days. A median, sagittal — section is represented in stipple. In addition to this the left buccal ridge (q@) is shown in surface view at a lower level. This buccal ridge is seen to be continu- ous with the left lip at the border of the mouth. Below the lip the short lappet is shown somewhat contracted. The mouth is widely opened and extends hori- zontally between the lines converging at m. The intestinal sphincter is partially contracted. The mesodermic structures are indicated as in fig. 2. Two mesen- chyme cells with amceboid processes lie above the digestive canal, and several others without processes lie beneath the intestine. x 500.
Figure 4.—Transverse section of a lappet more highly magnified to show the relation of the mesodermic structures to the ectoderm-cells. Three muscle-cells are shown with their fibrous processes inserted between the epithelial cells. Near the angle of the lappet a single, rounded mesenchyme cell is represented. The letter b indicates the ectoderm of the lateral surface of the body; c, that of the under surface.
Figure 5.—Five partially developed muscle-cells with contracted processes after kill- ing with corrosive sublimate.
PLATE XXXV.
The mesoderm is printed in red. F
Figure 1.—Median, sagittal section of the pilidium of M. ceca at the end of the third day. This is a much earlier stage than that shown in fig, 3, Pl. xxx1. The muscular cells have attached themselves in their definite positions, and have already begun to send out fibrous contractile processes. Not all of the mesen- chyme-cells are distinguishable from the muscle-cells, The outline of .the lappet is indicated beneath the section. One of the buccal ridges (a) is indicated, as is also the extent of the mouth (m). x 500.
Pilidium of Certain Nemerteans. 261
Figure 2.—Dorsal view of pilidium of I. ceca at the age of tweive days. In the center are.the cells of the apical plate (black) with two large flagella directed upwards. To the right and left of the apical plate are seven or eight muscle- cells with their fibers running down the sides of the body towards the lappets. These fibers constitute the lateral muscular band. In front and behind the apical plate the muscle-cells are few in number, and repeatedly anastomose. The ectoderm is not shown; a, anterior, p, posterior end. Slightly diagrammatic. x 500.
Figure 3.—Pilidium of If ceca at the age of twelve days seen from the left side. The epithelium is supposed to be transparent except at the margins of the draw- ing, where it is shown in section to illustrate its thickness; a, anterior end, p, pos- terior end. At the side of the apical plate are seen six muscle-cells with nearly parallel fibers running directly into the lappet. These lie immediately beneath the surface epithelium, and constitute the lateral muscular band. A few cells on the lateral face of the embryo send out anastomosing fibers in all directions. The horizontal band of muscles, indicated by the dotted lines (6), are those which run antero-posteriorly at the sides of the mouth. Three mesenchyme cells are indicated by c, while a half-dozen others are seen between the muscle-cells in the lappets. Slightly diagrammatic. The number and position of the muscle- cells and fibers was confirmed in the living object, and also in many stained preparations. x 500.
Figure 4.—An anterior view of a pilidium of the same age as those in figs, 2 and 3.
Figure 5.—Pilidium of C. marginatus at the age of ten days. The dotted lines indi- cate the digestive tract.
Figure 6.—Section through the apical plate to show the relation of the muscle-cells to the epithelium.
LITERATURE.
1. Arnold, G., Zur Entwicklungsgeschichte des Lineus gesserensis. Trav. Soc. Imp. Nat, St. Pétersbourg, vol. xxviii, 1898. Review by O. Biirger in Zool. Cen- tralblatt, vol. vi, 1899.
2. Barrois, J., Mémoire sur |’Embryologie des Nemertes. Annals des Sci. Nat., ya, LST.
3. van Beneden, P. J., Recherches sur la faune littorale de Belgique. Mém. Acad. Sci. de Belgique, T. xxxii, 1861.
4. Burger, O., Studien zu einer Revision der Entwicklungsgeschichte der Nemer- tinen. Ber. Nat. Ges. Freiburg, Bd. viii, 1894.
5. -——-.—— Nemertinen. Monograph 22, Fauna und Flora des Golfes von Neapel. Berlin, 1895.
6. Bitschli, O., Kinige Bemerkungen zur Metamorphose des Pilidiums. Archiv. f. Naturgeschichte, 1873.
7. Child, C. M., The maturation and fertilization of the egg of Arenicola marina.
Trans. N. Y. Aead. Sci., vol. xvi, 1897. Coe, W. R., The maturation and fertilization of the eggof Cerebratulus. Zool. Jahrbiicher, Bd. xii, 1899. 9. Conklin, E. G., The Embryology of Crepidula. Journ. Morphology, vol. xiii, 1897. Desor, E., Embryology of Nemertes, ete. Boston Journ. Nat. Hist., vol. vi, 1848.
om
~ a
34. 35.
W. R. Coe—Development of the Pilidium, ete.
Dieck, G., Beitr. zur Entwicklungsgeschichte der Nemertinen. Jenaische Zeitsch. Naturwissenschaft. Bd. viii, 1874. es
Gegenbaur, C., Bemerkungen tber Pilidium, ete. Zeitsch. f. wiss. Zool., Bd. vy, 1854,
Griffin, B. B., The history of the achromatic structures in the maturation and fertilization of Thalassema. Trans. N. Y. Acad. Sci., 1896. .
Hoffmann, C. K., Entwicklungsgeschichte von Tetrastemma varicolor. Nieder-- land. Archiv. f. Zool., Bd. iii, 1877.
—-—— Zur Anatomie und Ontogenie von Malacobdella. Ibid., Bd. iv, 1877. _
Hubrecht, A. A. W., Proeve eener ontwikkelingsgeschiedenis van Lineus ob- scurus. Utrecht, 1885.
—— Contributions to the Embryology of the Nemertea. Quart. Journ. - Mic. Science, vol. xxvi, 1886. Kostanecki and Wierzejski, Ueber das Verhalten der sogenannten achromat- ischen Substanz im befruchteten Ei. Arch. f. mik, Anat., Bd. xlvii, 1896. Krohn, J., Ueber Pilidium und Actinotrocha. Miiller’s Archiy. f. Anat. u.— Physiol., 1858. - Lebedinsky, J., Beobachtungen tiber die Entwicklungsgeschichte der Nemertinen. . Arch. f. mik. Anat., vol. xlix, 1898, pp. 503-556. Nachtrag, pp. 623-650. Leuckart und Pagenstecher, Untersuchungen ueber niedere Seethiere. Miller’s Archiv. f. Anat. u. Physiol., 1858. _
MacIntosh, W. C., British Annelids, I, Nemerteans. Ray Society, 1873.
Mead, A. D., Development of marine annelids. Journ. Morphology, vol. xiii, 1897. :
Metschnikoff, E., Studien tiber die Entwickelung der Echinodermen und Nemer- tinen. Mém. Acad, Imp., St. Pétersbourg, T. xiv, 1869.
Vergleichend-embryologische Studien. Zeits. f. wiss. Zool., Bd. XXxXvii, 1882. ,
Miller, J., Ueber verschiedene Formen von Seethieren. Miiller’s Archiv. f, Anat. u. Physiol., 1854. -
Salensky, W., Recherches sur le développement du Monopora vivipara. Archives de Biologie, T. v, 1884.
——-—— Bau und Metamorphose des Pilidium. Zeits. f. wiss. Zool., Bd. xiii, 1886.
Schultze, M.; Zoologische Skizzen. Ibid., Bd. iv, 1853.
Verrill, A. E., The Marine Nemerteans of New England. These Transactions, vol. viii, 1892.
Supplement to the Marine Nemerteans and Planarians of New England. Ibid, vol. ix, 1895.
Wilson, C. B., Activities of Mesenchyme in certain larvae. Zool. Bulletin, vol. ii, 1898.
Wilson, E. B., On a new form of Pilidium. Johns Hopkins Univ. Biol. Studies, vol. ii, 1882.
———— The Cell-lineage of Nereis. Journ. Morphology, vol. vi, 1892.
———— Considerations on cell-lineage and ancestral reminiscence. Ann. New York Acad. Sei,, vol. xi, 1898.
VI.—Tue Marvuration, FerrinizaTion AND Earty DErveELop- MENT OF THE PLANARIANS. By Wit~aRpD G. VANNAME.
Tllustrated by plates XXXVI to XLI inclusive.
Tue eggs of some species of planarians, particularly those of some of the polyclads, offer many advantages for the study of the processes of maturation and fertilization. The ease with which many of these animals. may be kept in captivity, and the abundance of eggs which they produce when kept under the right conditions, make the mate- rial easy to obtain if the adult animals can be secured at the proper season, while the fact that the eggs are fertilized within the body of the adult protects them from the unnatural conditions which so often produce abnormalities and pathological changes in eggs artificially fertilized.
There are, however, difficulties in the way of preserving these eggs which make good preparations not easy to obtain. The membrane enclosing the egg is very thick and is penetrated by preserving fluids with difficulty. In addition to this they are in many cases thickly coated with mucus, which is coagulated by the reagent and offers much resistance to its penetration. These envelopes of the eggs are also apt to cause trouble in imbedding and in cutting sections.
The consequence has been that although the eggs of a number of species of planarians have been studied by different investigators, there is uncertainty and disagreement in regard to many points which can hardly be due to essential differences between the species studied, since with one exception they were all polyclads, as are also the two which are the subject of the present paper.
Mitotic figures were observed in the eggs of planarians by Selenka (21) and Lang (18) many years ago and a series of short papers was published by Van der Stricht (22 to 27 inclusive) on the egg of Thysanozoon Brocchi, but the principal contributions to our knowl- edge of the subject have appeared within the last two or three years. These are the work of Klinckowstrém (14) on the egg of Prosthe- cerceus vittatus, the two papers of Francotte (3, 4) describing and illustrating those of the same and several other polyclads, and a later work of Van der Stricht (28) on the egg of ZThysanozoon. The investigations of the two last mentioned writers especially have done a great deal to clear up the doubtful points and bring our ideas of
264 W. G. Van Name—Embryology of EBustylochus.
the development of the planarian egg into line with the observations on those of animals of other classes.
But as I have been so fortunate as to obtain some unusually good preparations of polyclad eggs, a description of them may be of interest not only on account of the additional light it may throw on the points left unsettled or passed over briefly by the previous. writers, but also as confirming many of their conclusions by means of investigations on the eggs of different genera and species.
I studied at first only the eggs of Hustylochus ellipticus (Girard) Verrill, a polyclad found abundantly in the vicinity of New Haven, Conn., during the winter and spring months, on the beach a little below high water mark, clinging to the under side of large stones. The animals may be kept alive for many weeks in sea-water, if it is occasionally changed. A few eggs were laid in January and Febru- ary, but more were produced during March, April and May.
Girard (7, 8) published an account of the embryology of this spe- cies under the name Planocera elliptica, nearly half a century ago. His figures are remarkably well drawn, and considering that he wrote at a time when the maturation, and the significance of the fertiliza- tion and cleavage of the egg were not.understood, the number of facts correctly observed by him is remarkable. No one else, so far
as Iam aware, has published anything in regard to its embryology, _
and of course not only the cytology, but much else remained as a new field for investigation.
The eggs are laid in clusters or sheets containing from one to two dozens up to several hundreds, arranged in a single layer and closely attached together with a white mucus-like secretion, which is at first very sticky and adheres to everything it touches, but gradually becomes harder. The eggs lie so close together in the cluster that the egg membranes, though not the egg itself (for there is a space between them), become more or less polygonal and no definite arrangement of the eggs can usually be made out. In Planocera nebulosa Girard, another polyclad of which I afterwards obtained eggs, they are not placed so close together, and their arrangement in a zig-zag line extending back and forth across the cluster, can readily be seen. It took an individual of this latter species less than a min- ute to lay a row of eggs across a cluster about three-sixteenths of an inch wide and back again, the animal keeping the forward end of the body stationary or advancing it slightly as row after row of eggs was laid, and slowly moving from side to side the posterior part of the body, where the reproductive openings are situated. In the case
W. G. Van Name—Embryology of Eustylochus. 265
of both species the clusters of eggs are usually deposited on the bottom or sides of the jar in which the animals are kept. Some- times they are laid on the surface of the water and float until once submerged, when they sink to the bottom.
Eggs of Hustylochus ellipticus.
The eggs average 0°080™™ in diameter. Each is enclosed in a tough membrane more than 0:100™™ in diameter, so that the egg is free to move in the clear fluid filling the intervening space. Owing to the large number of yolk globules which it contains I was able to make out nothing definite concerning the internal processes of maturation and cleavage in the entire egg either living or stained.
Development is very slow and varies greatly in the time occupied
in the different stages, so that some of the eggs in a cluster are often farther advanced than others. In other cases, however, the eggs of the same cluster are in nearly the same stage.
A few hours after the egg is laid, which occurs more often at night than during the day, it flattens out at one point and the first polar body is given off. The time that elapses before this occurs is probably largely dependent on the temperature, as Francotte found to be the case in the eggs of Leptoplana. The polar body exhibits amceboid movements both before and for some time after its com- plete separation. Fig. 30 shows the different forms assumed by the polar body in one instance, in the course of ten minutes. When the first polar body has separated the egg again becomes spherical but soon flattens again in the same region, and the second polar body separates in a similar manner. Both polar bodies are large, the diameter of the second and smaller one being about one-tenth that
.of the egg, while the first is about one-half as large again as the second, and usually assumes an elongated form. They usually be- come detached from the egg after a short time, and float around between it and the egg-membrane, but they show no signs of disin- tegration even after they have been rolled around in this space for many days by the motion of the cilia which develop on the embryo. I have observed no ameeboid movements of the egg itself.
Some twelve or fifteen hours after the egg is laid the first cleavage
“takes place, though it may occur sooner or be delayed still longer. It is a vertical and so far as I can determine an equal one. The second cleavage planes are not quite vertical but are so inclined that the somewhat smaller cells that are budded off lie in a slightly higher
“
266 W. G. Van Name—Embryology of Eustylochus.
plane than the others. They separate in the direction of a left- handed spiral. The third cleavage is horizontal and the cells are budded off in the direction of a right-handed spiral. The upper four cells (about the animal pole) are quite conspicuously smaller than the others and of about equal size in spite of the inequality of the cells from which they are derived. The two, four, and eight- ° cell stages are shown in Figs. 44, 45 and 46.
Eggs of Planocera nebulosa V errill.
In the eggs of Eustylochus I was unable to discover the sperm- centrosome and sperm-aster. In the effort to find them many lots of material, in every stage in which these structures might be looked for, were preserved in fixing fluids of different kinds. Dozens of slides were mounted, many of them showing the maturation and cleavage spindles in a beautiful state of preservation and satisfac- torily stained. These were gone over with the greatest care with a high power objective, but I failed to find any trace of the sperm- asters er anything that I had any reason to believe was the sperm- centrosome.
Still as the egg of Hustylochus ellipticus is densely packed with large yolk globules which not only stain with hematoxylin but also with plasma-stains, and are in many preparations so dark colored and densely crowded that the sperm nucleus itself is often impossible to distinguish, I was even then unwilling to accept this negative evi- dence as final, and determined to examine the eggs of some other planarian, where the conditions might be more favorable,
Through the kindness of Dr. Wesley R. Coe of New Haven, to whom I ‘am also indebted for some preparations of the eggs of Eustylochus which he had himself made some time previously, I received during the spring of this year (1899) some living individu- als of the rather rare species Planocera nebulosa Girard. These he obtained while collecting the large nemertean Cerebratulus lacteus Verrill, in the tubes or burrows of which this species usually lives, though it is occasionally found elsewhere. It can be kept in confine- ment more readily and lays even more freely than Hustylochus ellip- ticus. My specimens laid their eggs in March and April, more abundantly during the latter month.
These specimens of Planocera nebulosa were pronounced by Prof. Verrill identical with those he has called by this name in his work on the marine planarians of New England (29). In that work (p.
W. G. Van Name—Embryology of Eustylochus. 267
94) he says of this species: “I have referred this with considerable doubt to Girard’s species, for the description of the latter was very mieapre..... The generic relations of this species are somewhat Geubtiul. .... In many respects the reproductive organs are like those of Hustylochus, and it is possible that the affinities may be even greater with that genus than with Planocera.”
This question I have not had the material or the opportunity to investigate. The species is, however, a strongly characterized one, both in appearance and habits, and is readily recognizable, so that no confusion will be caused should it prove necessary to transfer it to another genus. It is an exceedingly active creature and crawls with considerable rapidity, for a planarian, while Zustylochus ellip- ticus 1s very sluggish and moves but little unless disturbed.
The eggs are larger than those of Hustylochus ellipticus, measur- ing 0090" or more, but the amount of shrinkage which occurs in preserving the eggs and in imbedding them is so variable that in sections they often appear no. larger than those of Hustylochus. They are surrounded by a membrane considerably larger than the egg, but in this species it is much thinner, the amount of mucus present about them is less, and they are not laid so close together.
The difference in the size of the eggs of the two species does not seem to affect the size of the mitotic figures, pronuclei or polar bodies, but results simply from an increased amount of cytoplasm and yolk.
The descriptions and figures in the remainder of this paper are taken from eggs of Hustylochus unless otherwise stated. Neverthe- less the resemblance between the eggs and embryos is so close that they would apply almost equally well to Planocera nebulosa. Dit- ferences of any importance between the two species have been noted. The methods of preparation used were the same in the case of both species.
Methods of Preparation.
As I have already intimated, it was necessary to depend entirely upon sections for studying the internal processes of development: In preserving the eggs a number of different reagents were tried ; saturated solution of corrosive sublimate both with and without the addition of two per cent. of glacial acetic acid or one and one-half per cent. of nitric acid, picro-acetic containing one per cent. of acetic acid, seventy per cent. alcohol with five per cent. of glacial acetic acid, Gilson’s, Hermann’s, Flemming’s and Perenyi’s fluids.
268 W. G. Van Name—Embryology of Eustylochus.
With all of these except the last three I obtained at one time or another fair preparations. The chief objection to the fluids contain- ing osmic acid is the difficulty of staining the eggs fixed with them, but in addition to this the structures were usually not so well preserved as with some of the other reagents. On the whole I found the corrosive-acetic and picro-acetic fluids the most satisfac- © tory, and nearly all of my best preparations were preserved in one or the other of these two reagents. The clusters of eggs can be handled entire, but because of the thick membrane and the coating of mucus surrounding the eggs the penetration of the killing fluid is much interfered with, and satisfactory preparations are exceptional, while a large percentage are absolutely worthless.
The appearance of the cytoplasm and yolk in the finished prepara- tion varies with the reagent used in preserving them. Most of the above mentioned fluids, especially those containing corrosive subli- mate, harden the yolk globules well, so that they preserve their spherical shape and to a considerable extent their size. They stain quite readily, both with hematoxylin and with plasma stains. The picro-acetic fluid, however, allows the yolk spheres to shrink into irregular forms and to so great an extent that the whole egg often becomes greatly contracted and of irregular outline. The yolk has less affinity for stains, and the cytoplasmic reticulum, not the yolk globules as in the corrosive preparations, becomes very conspicuous. I have endeavored to reproduce this appearance in the drawings, some of which (for example, Figs. 2, 11, 25 and 27), were drawn from picro-acetic preparations.
The masses of eggs were usually left in the fixing fluid over night, washed in seventy per cent. alcohol, dehydrated and imbedded in parafin. I did not meet with any difficulty in imbedding the eggs, nor in cutting them into sections ‘007 to ‘°011™™ in thickness, which was as thin as necessary. Many of the preparations could be cut much thinner.
Klinckowstrém, Francotte and Van der Stricht all found great difficulty in imbedding or cutting their material, but I found no unusual precautions necessary except to take care that the eggs remained in each change of alcohol or clearing fluid long enough to insure thorough penetration. For clearing and as a solvent for the paraffin I used xylol, and my success may have been partly due to the use of this medium as well as to the more favorable material. For the study of the early stages, previous to laying, the entire animals may be killed and imbedded, but as observed by Van der
W. G. Van Name—Embryology of Eustylochus. 269
Stricht, it is better to tear up the animals and to preserve the frag- ments. In this way more rapid and complete penetration, is possible. The fragments of the animal with the eggs still in them were then imbedded and sectioned in the manner deseribed above. For these early stages I found the picro-acetic fluid much the most satisfactory fixing agent.
For staining, Heidenhain’s iron-hematoxylin method was usually employed, but certain structures, such as the nuclear network of the germinal vesicle, were shown much better by staining with Delatield’s hematoxylin. With these I generally used a ground stain of Orange G or Bordeaux red, the latter proving the most satisfactory. In using these, particularly the orange, care must be taken not to stain too deeply, as the structure then becomes obscured, and it is _ difficult or impracticable to extract the stain again. It apparently makes no difference whether the sections are treated with the ground stain before or after the hematoxylin.
With saffranin, which was largely used by Van der Stricht, I had little success. When the iron-hematoxylin method is used, the yolk globules stain black, but usually a large proportion of them, or sometimes all, become decolorized in the subsequent extraction with the alum solution. If the extraction of the stain is only partial, many of the yolk globules are left with a black center, because of the tendency of the stain to dissolve away first on the outside of the globule, where it is first reached by the solution. Such cases are shown in Figs. 37, 38 and others. Such an object, especially when close to the sperm-nucleus, may easily be mistaken for a centrosome surrounded by a centrusphere. The absence of radiations and the occurrence of such objects in various points of the egg are usually sufficient to show their true nature.
The Germinal Vesicle.
The shape of the eggs contained in the ovaries is determined by the pressure of the adjacent eggs and tissues. With the exception of those of advanced development—those which are forming or have formed the first polar spindle—all contain a large round or oval germinal vesicle (Fig. 1). In sections of fully developed ovarian eggs this may be 0°02" or more in diameter, and it always Contains a large nucleolus, which, as observed by Francotte and Van der Stricht, in the early stages stains more deeply with hematoxylin than in the later. It is nearly spherical and bounded by a definite outline. At first it is homogeneous, in later stages sometimes vacuo- lated, though this may be due to defective preservation.
270 W. G. Van Name—Embryology of Eustylochus.
Owing to the irregular shape of the eggs I cannot say whether the normal position of the germinal vesicle is central or eccentric, nor whether the orientation of the egg is already determined.
The germinal vesicle contains a more or less irregular network of threads which do not stain very dark with Delafield’s hematoxylin, although with the iron-hematoxylin method they stain black. The latter method is, however, poorly adapted for showing the real struc- ture of this network and the early stages of the formation of the chromosomes of the first polar spindle, on account of the tendency to precipitate hematoxylin in excess in certain places, increasing largely the apparent amount of chromatic substance and disguis- ing the structure. This has also been noticed by Francotte and Klinckowstrém. Stained with Delafield’s hematoxylin the threads are seen to form a network thickened at the nodes and where the strands touch or unite with the nuclear membrane (Fig. 1).
In immature ovarian eggs the cytoplasm is comparatively small in amount and the germinal vesicle occupies a large proportion of the bulk. The change in size and proportions which takes place as the egg ripens is due largely to the formation of the yolk, which, by the time the egg is ready to leave the ovary, is distributed throughout the whole of the cytoplasm in the form of small globules. This comparatively uniform distribution of the yolk persists to a con- siderable degree through all the maturation and early cleavage stages, so that as a rule only the space occupied by the mitotic figures or nuclei, and sometimes their immediate vicinity, is entirely free from yolk globules.
The cytoplasm appears to consist of a delicate reticulum of threads built up of microsomes staining dark blue with hematoxylin, in the meshes of which the yolk globules lie. Besides this there appears to be a clear matrix in which both the reticulum and the yolk globules are contained. Coe (1) observed this in the egg of Cere- bratulus, but suggests that it may be an appearance due to the shrinkage of the yolk spheres. If no such matrix exists it would seem probable that the cytoplasm has really an alveolar rather than a reticular structure.
The Formation of the First Polar Spindle.
The early stages in the formation of the chromosomes proceed substantially as described by Van der Stricht (28). The observa- tions of Francotte, including those on Prostheceraeus, where Klinckowstrém failed to find a spireme stage, are also much the
an “~~
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W. G. Van Name—Embryology of Eustylochus. 271
same. Near the periphery of the germinal vesicle a part of the nuclear network can be seen to increase in thickness and stain more deeply. It continues to thicken and assumes a beaded outline, showing that it is composed of minute masses or grains of chromatin, connected together (Fig. 2). Whether this is already broken up into segments representing the future chromosomes, when it first becomes noticeably different from the rest of the threads of the network, I have not been able to discover, as it is almost certain to be cut at different points by the planes of the section. In the egg shown in Fig, 2 it is pretty evident that it has already broken up into segments. These show no signs of longitudinal cleavage in anticipation of the division of the chromatin between the egg and the first polar body. They contract into rounded or oval masses (Figs. 3a and 36) the chromosomes of the first polar spindle, while the remainder of the nuclear network gradually disappears.
By this time the asters of the future spindle have become dis-
tinguishable (Figs. 2, 3a and 36). Specimens stained by the iron- hematoxylin method show that each contains a black staining centrosome (Figs. 3a and 30). _ The question of the origin of these centrosomes is one upon which there is great difference of opinion. Klinckowstrém found the two asters originating separately and simultaneously, their centrosomes lying about :040™™ apart. Van der Stricht’s own observations con- firm this; nevertheless he expresses his belief in the claim of Francotte that he has found the centrosome single—“ nous le voyons s’allonger, puis se fendiller et se diviser ainsi en deux nouveaux cor- puscules polaires” (3, p. 12).
In regard to the place and source from which the centrosome or centrosomes take their origin there is also difference of opinion. Klinckowstrém considered them of nuclear origin, basing this belief, however, on a single instance, of which he gives a figure. Van der Stricht believes that they originate from the nuclear membrane and are therefore of nuclear origin. He finds that they afterwards separate from the membrane. Francotte finds the centrosome appearing close to the nuclear membrane, sometimes “si voisin qu’il semble faire corps avec cette membrane,” but does not believe in its nuclear origin.
I have not demonstrated any connection between the centrosomes and the nuclear membrane nor found evidence of their origin by the division of a single body, though this of course may occur before the formation of the aster rays enables us to distinguish them. By
272 W. G. Van Name—Embryology of Hustylochus.
that time they lie in the cytoplasm near the nuclear membrane, which is usually more or less indented, and at a little later stage begins to disappear at these points. They lie widely separated but not necessa- rily at opposite poles of the germinal vesicle (Figs. 2, 3a and 36), and are not connected by a central spindle. ‘The similarity of this mode
of appearance to that found by Coe (1) in the nemertean Cerebratulus
and by Griffin (12) in Thalassema is very great. The observations of Coe are of especial importance, for the preparations on which his work is based have surely never been excelled in beauty and clear- ness of detail, if indeed they have been equalled.
When the centrosomes, which stain black with the iron-hema- toxylin, first become distinguishable, they are surrounded by a more or less distinct centrosphere, which stains with cytoplasmic stains
and runs out into the few short aster rays developed at this early » stage. It may be of some significance that there is no distinct —
line of demarcation between the rays and centrosphere, although in later stages this ceases to be the case. Moreover in this early stage, the rays seem to have but little connection with the cytoplasmic reticulum, but as they become longer they branch and grade imper- ceptibly into the reticulum with which they are continuous (Figs. 4 and 5), as Wilson (31) has so clearly shown in Toxopneustes.
These phenomena suggest that the rays are at first outgrowths of the centrosphere, but grow by the continuous addition and rearrange- ment of the microsomes of the reticulum. The branching which later becomes noticeable is largely determined by the positign of the yolk globules. This would indicate a difference of origin of the basal and peripheral portions of the aster rays. The indications are, however, that the centrosphere itself is differentiated from the cytoplasm surrounding the centrosome by the influence of the latter, making the ultimate origin of both portions of the aster rays the same.
The term centrosphere, as I use it, is equivalent to the “ centrosom ”? of Boveri and to the “zone médullaire de la sphére attractive” of Van Beneden and to the “couche médullaire (de la sphére attrac- tive)” of Van der Stricht. Such a structure as the “couche” or “zone corticale de la sphére attractive” of Van Beneden, which is also described by Van der Stricht in Zhysanozoon and by Fran- cotte, I do not find in well preserved eggs, and I find no evidence of any differentiated body external to and surrounding the centrosphere (Van der Stricht’s “couche médullaire’’), except the aster rays extending out into the cytoplasm. In poorly preserved eggs I have
4
W. G. Van Name— Embryology of Hustylochus. 273
sometimes seen more or less suggestion of such a ‘couche cortical,” but cannot consider it as anything but an abnormal or artificial effect, as the best preparations do not show it. An appearance which may be of somewhat the same nature is very frequent in eggs where the penetration of the killing fluid has been slow or imper- fect. No centrosome can be found in such eggs and the centro- sphere is indefinite in outline and surrounded by a light colored layer of greater or less thickness, in which no structure can be discerned. ‘The aster rays begin outside of this. Such a condition, although unquestionably abnormal and artificial, I have represented in Figs: 24 and 28, as other structures in the egg are well shown in these specimens. It occurs both in the polar and cleavage spindles.
Francotte (4) illustrates in his Figs. 27, 29, 30 and 31, sections _ which apparently have this same defect, and it is largely upon such that he appears to base his belief that the centrosphere (he calls this with the “corpuscle central” the ‘“ centrosome’) is surrounded by a membrane in the later phases of mitosis. Such a condition is not indicated by my best preparations, though it is probably true that the outer portion of the centrosphere becomes more dense. At least it stains more deeply. (See Figs. 5, 7, 8, 12 and 13.)
That which I have termed centrosome is the “granule central” or *‘corpuscule central” of Van der Stricht and Francotte, the “centralkorn” of various writers.
Van der Stricht (28) believes that this, as well as the centrosphere (“couche médullaire ”), is formed by differentiation out of a “cen- trosome” which is at first homogeneous. Moreover he considers that although the “sphére attractive,” which includes what I have termed the centrosphere, may not always reveal itself with all the clearness that might be desired, yet it is none the less a permanent organ of the cell.
This view is certainly not supported by the subsequent history of the centrosphere in Hustylochus or Planocera. As explained below, the greater part of it is destined to form the second maturation spindle, and the centrosphere that rémains in the egg after the sec- ond polar body is expelled rapidly degenerates and disappears. The manner in which the centrosphere is formed is a much more difficult matter to determine, but the poorly defined limits of the centro- sphere in the early stages both of the polar and cleavage spindles, where it runs out into the rays with no distinct line of demarcation between the latter and the centrosphere, when compared with its distinct and definitely limited outline in the later stages of the same
Trans. Conn. Acap., Vou. X. Aveust, 1899.
18
a
274 W. G. Van Name—Embryology of EHustylochus.
spindle, suggest that it is formed by a progressive differentiation of the cytoplasm about the centrosome.
As I have already said, there is at first no central spindle connect- ing the two asters which are to become the poles of the first polar spindle. They seem to be entirely independent (Figs. 2, 3a and 30). But as the nuclear membrane, which has everywhere become very — thin and delicate, is more and more completely dissolved, the rays extend into the germinal vesicle and connect with the chromosomes and with rays of the other aster (Fig. 4 and 5). Thus a spindle is formed, the fibers of which are in the early stages very irregular and anastomose with each other in so complex a manner that they appear somewhat like a more condensed and deeply staining part of the general reticulum of the egg.
As Coe (1) has found in the egg of Cerebratulus, the spindle is thus partly at least formed from material previously or to some extent still enclosed in the nuclear membrane, and therefore undoubtedly nuclear. This is Francotte’s opinion also. Van der Stricht expresses the belief that a rudiment of the central spindle exists from the time the centrosomes first separate, but admits that no such thing is visible in the early stages. It is probable, however, that if such a structure did exist and were sufficiently substantial to persist while the centrosomes move to opposite sides of the germinal vesicle, it would at the same time be distinct enough to be visible.
The development thus far takes place usually in the ovaries or at least in the first portions of the oviducts. Francotte however men- tions cases where the egg was laid while still in the germinal vesicle stage. Such instances I have not met with. By the time the egg reaches the uterus (Fig. 5) the first polar spindle is fully formed, the chromosomes lying in or near its equator, though some of them are often more or less displaced (Fig. 36). Their form is now usually that of a ring, a cleft or opening having appeared through the center of each. This ring form is sometimes visible ata still earlier stage (Fig. 3a). The other forms which the chromosomes may assume are described below.
About the time the nuclear membrane dissolves the nucleolus dis- appears also, Occasionally there are traces of it for a little while afterwards both in Hustylochus and Planocera. Francotte observed this in Leptoplana and Prosthecereus, but Van der Stricht found that it did not persist after the disappearance of the membrane i in Thysanozoon.
By this time the aster rays have increased both in length and in
W. G. Van Name—Embryology of Eustylochus. 27%
~I or
number, and may now be seen to be made up of microsomes, at least so far as their outer portions are concerned. ‘Toward the centers of the asters they appear homogeneous and their structure cannot be made out. They branch and connect with the cytoplasmic reticulum and thus indirectly with the periphery of the egg. The centrosomes (Figs. 5 and 36) have usually divided, so that there are now two in each centrosphere. This division, which may however be delayed till after the egg is laid, usually takes place in the direction of the axis of the spindle or nearly so. This fact I shall have occasion to speak of again, as it affects the process of formation of the sec- ond polar spindle. The two derived centrosomes remain quite near together, nevertheless the centrospheres usually become slightly elongated in the direction in which they separate. The centro- spheres are now bounded by a definite outline where the aster rays take their origin.
Fertilization.
Eustylochus, like other polyclads, is hermaphroditic, but the eggs of one individual are fertilized by the spermatozoa of another. The spermatozoa may be injected into any part of the body, and find their way to the eggs by their own activity, since they are able to penetrate through the tissues. This process of hypodermic impreg- nation is described by Lang (18) and other writers in other polyclads. The spermatozoa are introduced into a puncture made in the epi- dermis of the dorsal surface by the penis of another individual. They are mixed with a quantity of thick mucus so as to form a spermatophore of jelly-like consistency, and of fairly definite form (Fig. 31). This mass of mucus eand spermatozoa is not inserted entirely below the epidermis, but the greater portion of it remains projecting out of the animal as a white spherical mass, which may be 0°6™™ or even more in diameter. Sometimes as many as four or five of these spermatophores may be seen projecting from the same individual at various points on the dorsal surface, but more often toward the posterior than the anterior end.
Living spermatozoa are 35-40 » long, and very slender and thread- like. They taper at each end, but the anterior end is much more slender, exceedingly sharply pointed, and moves in snake-like curves or waves. These waves may travel in either direction, sometimes with considerable rapidity, but with little effect in moving the spermatozoon along in water, though no doubt efficient in penetrat- ing the body-tissues. No definite demarcation between the active
276 W. G. Van Name—Embryology of Eustylochus.
and slender end and the thicker and inactive part, probably the middle piece, is visible, nor can any tail be distinguished.
The spermatozoa are often present in the tissues in almost incredi- ble numbers, and some difficult questions in regard to the details of the process of fertilization present themselves. Although they can penetrate most of the body-tissues without difficulty, there must be ~ others that are more or less impermeable to them, as for example those of the male reproductive organs. Otherwise it would not be possible for an individual to store up the fully developed spermato- z0a, or to prevent fertilizing its own eggs.
I have not seen the spermatophores of Planocera nebulosa and therefore have not examined and measured the living spermatozoa. I often observed the animals apparently in the act of impregnation, — but on examining them found no spermatophore. Judging from preserved specimens of the animals and eggs, the spermatozoa of — Planocera do not differ from those of Hustylochus.
The entrance of the spermatozoon into the egg takes place about the time the process of forming the first polar spindle begins, which as I have said occurs either in the ovary itself or, usually at least, before the egg has got far from it. The possibility suggests itself that the entrance of the spermatozoon may be the stimulus which induces the disappearance of the germinal vesicle and the formation of the spindle.
Another question that arises is why the unripe eggs in the ovaries are not prematurely fertilized. Evidently it is not because the spermatozoa do not have access to the ovaries, although from the numbers of them generally to be seen just outside the ovary, the membrane enclosing the same mugt serve to keep them out to a con- siderable extent. The most plausible explanation is that the chemi- cal nature of the egg protoplasm is not such as to attract the sper- matozoon until it is of a certain degree of maturity.
It is exceedingly rare to find more than one spermatozoon in an g. If double fertilization is prevented by the formation of a vitelline membrane as soon as one spermatozoon has entered, the membrane must be extremely delicate, for I have been unable to distinguish such a structure. If present, it doubtless afterwards forms the inner layer of the tough shell which is deposited around each egg by the shell glands just before laying, and later becomes
or fo]
e
separated from the egg by a considerable space containing a clear fluid, for if this were not the case we should see it when the polar bodies separate, At that time, however, there is no interior mem-
doe
W. G. Van Name—Embryology of Eustylochus. 277
brane visible, but the polar bodies lie free in the space under the shell.
Eggs which lie in the uterus invariably contain a spermatozoon, which lies fully as far removed from the surface as at the end of the second maturation spindle (Figs. 5 and 36). It stains very deeply throughout its whole length and lies curved and twisted into various loops and bends. It is not apparent that it enters at any particular part of the egg’s surface or has at this stage any particular position in relation to the spindle, which it often approaches very closely, especially in Planocera nebulosa.
At this stage, especially in eggs of Hustylochus (it is not usually conspicuous in Planocera), the egg-cytoplasm, immediately about one end of the spermatozoon, has a more densely granular appear-
_ ance and stains more deeply (Figs. 5 and 6). In some preparations
this is exceedingly conspicuous (Fig. 6). I have not discovered anything of the nature of a centrosome in this and attribute it to the disintegration of some non-nuclear portion of the spermatozoon. By the time the egg is laid it has disappeared.
In poorly preserved specimens it is not uncommon to find spots and discolorations of various kinds and shapes near the spermatozoon. There is no constancy in their appearance and they do not occur in the best preparations. The egg can remain in the uterus in the conditions shown in Figs. 5 and 36 for an indefinite period. No further change appears to take place until it is laid.
Later Stages of the First Polar Spindle.
Lad
A section of a recently laid egg is shown in Fig. 7. The spindle as in other polyclad eggs has a central position, and measures between the centers of the asters three-eighths or more of the diameter of the egg. The two centrosomes in each aster often le so near together that unless the extraction of the stain is carried sufficiently far they cannot be distinguished as separate, though usually the centrosome, if but one appears, has an elongated form showing that it may be double. The centrosphere surrounding them appears homogeneous in structure, and is elongated in the direction of the axis of the spindle. It has gradually become bounded by a distinct and smooth outline, from which the rays, still rather few in number and about one-half the length of the spindle, have their origin. They branch and are lost in the spaces between the yolk globules, which are quite uniformly distributed in the egg, and come close to the spindle and centrospheres.
278 W. G. Van Name—Embryology of Eustylochus.
The spindle fibers, as in the earlier stages, are few in number and slender, but have become much more regular. The chromosomes generally have still the ring structure though they are now elongated and oval in outline.
At this time the asters of the spindle are alike in size and appear-
ance and it cannot be predicted which one is to pass into the polar ~
body. But soon one pole of the spindle approaches the surface of the egg.
This is accomplished by a movement of the whole spindle, which meanwhile shortens to about one-third of the diameter of the egg or even less. The aster of the outer pole becomes reduced in size, and some of the rays necessarily disappear altogether as the centrosphere approaches and finally comes in contact with the surface of the egg (Fig. 8). Finally the last of the rays at this pole and the centro- sphere itself fade away.
Division of the Chromatin.
The shape of the chromosomes in the equatorial plate is very variable, and at first sight it would appear difficult to derive some of the forms from the closed rings. Very similar if not absolutely identical forms are found in all the species of polyclads described by the writers already referred to. Van der Stricht has given the most thorough discussion of the relation of the different forms of chro- mosomes to the original elongated segment and to the ring derived from it. His conclusions are for the most part confirmed or at least supported by what I have observed in EZustylochus and Planocera.
There is, I think, little room for question that we have here a true case of heterotypical mitosis (Flemming, 2). No longitudinal split- ting can be seen in the elongated beaded segment which is the first stage of the chromosome. The first sign of its approaching division, the cleft or opening through it, does not appear until it has con- tracted into so nearly round or oval a form that from its shape we cannot do much more than guess which diameter represents the original long axis of the chromosome. If we are to ascertain whether this cleft really is a longitudinal cleavage, an “aequations- teilung” of the chromatin, we must find some other means to deter- mine it.
The way that the ring opens and divides proves that it consists of two elongated segments, each bent into a semi-circle or U and joined end to end. The junctions of these segments therefore come’ at opposite points on the ring. The spindle fibers attach themselves
W. G. Van Name—Embryology of Eustylochus. 279
to points midway between these junctions, and as they draw upon them the ring becomes a more and more elongated ellipse. As the points where the spindle fibers are attached move apart the line join- ing them becomes the long axis of the ellipse, and the points of union of the segments the extremities of the short axis. The ring then breaks at one of these points, and the tension of the spindle fibers then tends to straighten the whole chromosome (which now consists of two long segments joined at one end only) and to bring it into a line parallel to the spindle fibers. The junction of the seg- ments then comes at the middle point of the long slender rod thus produced, which is not necessarily a straight one. Eventually it breaks here also and the derived chromosomes, each having the form of a shorter rod, usually bent into a U form, pass to their respective
poles of the spindle. As the spindle fibers are not attached to the
ends of the segments but near their middle, the long rod before divi- sion is usually more or less curved or hooked at each end, the extreme ends not being so rapidly drawn toward the spindle poles as the points half way toward the middle where the spindle fibers are inserted. The segments derived from either this or the straighter form regularly have a U form on account of the tension being applied to the middle of their length instead of the end or ends. Another effect of the tensions thus applied, is to twist the chromosome as soon as the ring has broken at one point. This, together with the hooked end anda tendency which the segments have to become knobbed at their ends (including those that remain joined) often causes the chromosome as a whole to assume an appearance strik- ingly like an italic letter 7. The successive stages I have shown dia- grammatically in Fig. 41. Actual examples appear in Figs. 9, 10 and 36.
This process of division may be regarded as the regular one, and the other methods which occur as modifications of it. The most important modification is produced apparently by an increased amount of cohesion of the ends of the future segments when the chromosome still formsaclosedring. As before, itis drawn out into an ellipse with the junctions of the end of the segments at the ends of the short axis. Instead, however, of the ring opening at one of these points the segments continue to hold together, the ellipse becomes more and more elongated, and the sides eventually come together and appar- ently fuse, making instead of a long narrow rod consisting of the future segments united to each other by one end of each, a short thick rod in which each segment has the form of a loop the two
280 W. G. Van Name—Embryology of Hustylochus.
arms of which have come together and fused. As in the form first described, a transverse division of the rod at its middle point finally separates the derived segments. In the latter form, however, they are joined by both ends, in the former by but one end of each. This process is shown diagrammatically in Fig. 42. Actual examples in different stages appear in Figs. 9, 10 and 11.
Both these processes were recognized by Francotte but described with especial care and fullness by Van der Stricht (28). Klinckow- strom figured them but gave little explanation. We often—indeed
I might say generally—find different stages of both of these processes — going on in different chromosomes of the same spindle. Of each
there are many minor modifications, so that a great variety of forms results. Most of them are due to the tendency of the derived seg- ments to thicken at certain points, especially at the middle (where, as I shall describe later, they are probably destined to divide in the second polar spindle) and at the ends, where they form knobs. These thickenings often become conspicuous long before there is any sign of separation at these points, or even in the ring stage. The result, when it occurs early, is that the ring is thickened at four points, giving the appearance of a tetrad group connected by narrower commissures. Such figures are mentioned by Francotte., It is how- ever in these thickened places, and not between them, that the chromosome finally divides.
By far the most interesting modification is one which gives us some suggestion in regard to the distribution of the chromatin. The actual partition of the chromosome in both the methods above described is transverse to its long axis. The important question is whether this partition is transverse to the original long
axis of the beaded segments found in the early stages of spindle
formation, in’ other words, whether we have an equal or a reducing division of the chromatin. The modification which gives us a hint in regard to this important matter is shown in Fig. 43, examples of it also in Fig. 10. 5
This diagram (Fig. 43) shows not successive stages in the history of a chromosome as the other two diagrams, but different degrees of modification from the type shown in Fig. 42. As will be understood from the figures, in its extreme form it results in a cross-shaped chromosome. Such forms are described but not explained by Van der Stricht. The interesting point about this form is that it would hardly be possible to produce it unless the cleavage of the chromo- some which produces the ring is actually a longitudinal one. Here
‘i> asa
W. G. Van Name—Embryology of Eustylochus. 281
we see that the segments are not attached by their ends only, but also for a considerable distance (varying in different cases) along towards their middle points, showing what thejr original relations to each other are.
This supports Van der Stricht’s opinion that the first division is in reality a longitudinal or equal one, and ‘his suggestion that the same explanations would clear up the apparently transverse division described by Korschelt (15, 16) in Ophryotrocha.
Van der Stricht describes “secondary rings” formed by the re- fusion of the ends of segments already separated. It is not necessary to assume such an unlikely process for the explanation of any of the forms. Where we find the ends of the segments united, it is much more reasonable to suppose that they have not yet separated. I
_ cannot believe that the open loops figured by Van der Stricht are destined to form rings again. The only case when there is indica- tion of fusion is in the form of chromosome shown in Fig. 42, where the sides of the ellipse apparently do actually fuse, converting the ellipse into a solid rod.
The derived segments of chromatin which result from the form of division shown in Fig. 42 are short thick rods and are really the same as the loops resulting from the form shown in Fig. 41 except that the two sides of the loop have come together and fused.
The Centrosomes and Asters.
While the aster rays of the other pole of the spindle have, as already mentioned, become reduced, those of the inner pole have become longer, stouter and more numerous. The centrosphere of this pole has become proportionately larger, and its two centrosomes have moved a little farther apart. The centrosomes seem to vary
_ greatly in size. MacFarland (20) in describing the first polar spindle of Diaulula, says that the size of the central granule (“ centralkorn ”’), which undoubtedly corresponds to what I have in this description called the centrosome, is dependent, within certain limits, upon the
__ extent to which the stain is extracted. Kostanecki and Siedlecki (17) found much the same thing in the case of the cleavage centrosomes of __ ~Asearis, where Fiirst (5) was even led to believe that they are noth- _ ing more than artefacts, Though I think that it would be impossible _ to explain the distinctness, the definite form, and especially the division of the black staining center of the asters in such eggs as Diaulula and Eustylochus, on the supposition that the centrosome is
282 W. G. VanName—Embryology of Eustylochus.
only the portion left colored after a partial extraction of the stain, the iron-hematoxylin method may evidently mislead us as to the actual size of this organ. In the present case the deeply stained center of the aster may be reduced by continued extraction with the alum solution from a body apparently occupying the whole centrosphere, down to one so minute as to be scarcely visible with . the highest powers of the microscope, or it may be entirely dissolved away. ;
The enormous centrosomes, often unsurrounded by any indication of a centrosphere, which appear in some of the illustrations of Klinckowstrém and Francotte, are probably due to the whole centro- sphere being stained.
Figs. 12 and 18 are later stages of the spindle, showing the separation of the derived chromosomes and the formation of the polar body. The divergence of the centrosomes of the inner pole in preparation for the second polar spindle may already be noticed. — The rapidity with which they move apart, or the time when they start to do so, is not dependent upon the rate of separation of the chromosomes. This will readily be seen by comparing the progress made in these processes in the eggs shown in Figs. 12 and 13. Often as the centrosomes separate a constriction appears in the outline of the centrosphere between the centrosomes (Fig. 12). This appears to happen only when the latter move apart at an early stage. If their separation does not proceed far before the first polar body is separated the constriction does not appear. The centrosphere simply elongates and the new spindle forms as described below.
The segments resulting from the division of each chromosome remain connected by distinct and rather thick fibers even after they have become widely separated (Figs. 12 and 13). In the late phases they are generally closely grouped together. In the final stage of the spindle no true cell-plate is found, although such a structure is — often quite noticeable in the second polar spindle. This was also noted by Wheeler (30) in Mysostoma and by Coe (1) in Cerebratulus.
The Number of Chromosomes.
In Eustylochus ellipticus the number of chromosomes in the polar spindles is ten, and therefore twenty in the cleavage spindles, though they are then much more difficult to count. I have determined the number to be the same in Planocera nebulosa with almost as great a degree of certainty.
W. G. VanName—Embryology of Eustylochus. 283
Van der Stricht found nine in the polar spindle of Thysanozoon Brocchi, Klinckowstrém and Francotte six in the polar spindle of Prosthecereus vittatus and Francotte eight in the polar spindles of Leptoplana tremellaris, Oligocladus auritus, Cycloporus papillosus and Prosthiostomum siphunculus.
The Second Polar Spindle.
The centrosomes at the inner pole of the spindle continue to move apart and the centrosphere to become more elongated. Meanwhile the spindle fibers remaining in the egg degenerate and disappear, and the aster fibers also undergo degeneration, becoming fewer in number as well as shorter and less conspicuous. The extent to which this occurs appears to vary greatly in individual eggs but is never,
_as far as I have seen, carried to complete disappearance. The centrosphere of the inner pole of the first spindle soon begins to assume the form of a spindle (Fig. 14), even while the centrosomes are still quite near together. <A light area appears between the cen- trosomes in the elongating centrosphere. This approaches almost to each centrosome and is, as MacFarland (20) found in Diaulula, without definite outline toward the ends of the incipient spindle. The constriction of the centrosphere mentioned in describing the first polar spindle, has disappeared.
At a little later stage (Fig. 15) the centrosomes have moved far- ther apart and in the light central area a few fibers may be distin- guished which increase in number and length as the spindle lengthens. The aster rays also increase in number and length as the spindle grows, though they never attain the development found in the first polar spindle. The portion of the old centrosphere surrounding each centrosome assumes the outline and appearance of a complete cen- trosphere, while some of the aster rays, attaching themselves to the chromosomes draw them into the equator and become the mantle fibers. Thus with the exception of the chromosomes, and the outer portions of the aster rays as far as they are built up from the gran- ules of the cytoplasm, the entire mitotic figure appears to be derived from the centrosphere of the inner pole of the preceding spindle.
This seems enough to prove the incorrectness of calling the cen- trosphere the “centrosom,” as MacFarland (20) has done in describ- ing the maturation of the mollusk Diaulula. It is evident that we are not dealing with the centrosome only, but with much else in addition, and that such a use of the term is not in harmony with its definition aS a permanent and self-perpetuating organ of the cell. If we are
284 W. G. Van Name—Embryology of Eustylochus.
to apply the term to any visible object it must be the black staining “‘centralkorn.”
I have already stated that when the centrosome in each aster of the first polar spindle divides, the derived centrosomes usually sepa- rate in the direction of the axis of the spindle or nearly so (Figs. 5, 7,8, 12 and 13). This applies especially to Hustylochus,; in Plano- cera we find variations more frequent, and it is not rare to find them moving apart in a direction very oblique or even transverse to the spindle axis (Fig. 37), which is rather uncommon in Lustylochus, though not sufficiently so to be abnormal. The result of this is that the second polar spindle when formed has already a radial or nearly radial position, and no rotation of the spindle, or at least only a slight one, is necessary. The outer pole simply moves toward the surface of the egg, the spindle meanwhile lengthening, while the inner pole does not change its position to any great extent. Accord- ing to Lillie (19), a similar position of the spindle is common in Unio. As the outer pole moves toward the periphery of the egg it often pushes directly through the group of chromosomes, and thus some of the aster rays attach themselves to these and draw them into the equatorial part of the spindle. :
Van der Stricht, on the contrary, finds the centrosomes in Thysan- ozoon lying usually in a line perpendicular to the spindle axis, though ~ occasionally parallel or identical with it or in intermediate positions. Klinckowstrém in Prostheceraeus also finds the second polar spindle in a tangential position during its early stages.
The question of the relation of the aster rays of the second polar spindle to those of the inner pole of the first, is a most difficult one to answer. This is largely because of the very different appearances which eggs in about the same stage present.
MacFarland (20) describes the rays of the second polar spindle as arising anew, but before the outer portions of the rays of the old aster have entirely disappeared, so that we have at one time two sets of radiations present, the short new rays, and outside these, sur- rounding the whole spindle, the old aster rays whose central portions have disappeared.
Some of my preparations suggest such a process. Fig. 37, repre- senting an egg of Planocera nebulosa, shows one of these. The rays of the first polar spindle aster become weak as they approach the centrosphere, which looks as if it were developing new rays. Yet a careful examination of this specimen shows that the old rays may be traced clear to the centrosphere. Such a specimen as that shown in
W. G. Van Name—Embryology of EHustylochus. 285
Fig. 14 also suggests that new rays are being formed, the old ones having already disappeared.
Yet, on the other hand, there are many cases where we can scarcely doubt that the centrosphere simply lengthens and divides, each part carrying its rays with it, although the latter may become reduced in strength. In support of this, it is worth mentioning that although I have a very large amount of good material in this stage, particu- larly of Planocera nebulosa, | have never found a case where the rays had disappeared entirely nor one where I could distinguish an old and a new set of rays with certainty.
I think therefore that the conclusion is inevitable that the aster rays of the inner pole of the first polar spindle persist in part as rays of the second polar spindle asters, and my preparations further indi- cate that the extent to which they do this, and the extent to which new radiations are developed, varies greatly in individual eggs.
I do not find the rays at any period penetrating the centrosphere and reaching the centrosome as described by Van der Stricht, Fran- cotte and many who have studied the eggs of animals of other classes, though during the process of forming the spindle the cen- trospheres have a more or less indefinite outline. As the centro- sphere of the first polar spindle elongates, the part of it immediately surrounding the derived centrosomes becomes the centrosphere of the second polar spindle aster.
The second polar spindle (Fig. 16) is shorter and wider than the first, the aster rays are fewer and shorter and the chromosomes more or less widely separated in a lateral direction, which is in strong contrast to their crowded position in the late stages of the first polar spindle (Fig. 13).
The chromosomes of the second polar spindle are many times smaller than in the first and are less easily stained. Their form is usually irregular and they often divide while the spindle is in an early stage (Figs. 14-and 15).
The cross form described by Klinckowstrém, Francotte and Van der Stricht is also found occasionally in my preparations. It is explained by the latter as two short rods, the segments derived from the division of the chromosome lying across each other at right angles.
Although some specimens appear to support the theory that the division of the chromatin is here a transverse and reducing division, I cannot claim to have determined the relation of this plane of division of the chromatin to the previous one, as Van der Stricht
286 W. G. VanName—Embryology of Eustylochus.
appears to have done with some degree of certainty. Neither Klinckowstrém or Francotte give any evidence in support of this conclusion, upon which, nevertheless, they base their schemes of the distribution of the chromatin in the egg and the polar bodies.
Fig. 17 shows a very late stage of the second polar spindle, though most of the second polar body, which is here already constricted off, lies in another section. The chromosomes have swelled up into vesicles, in which a few threads of a network are beginning to appear, but as yet no nucleoli. The aster rays, centrosphere and
centrosome (it does not divide) have not yet begun to degenerate. —
This is the last stage in which I have found this centrosome.
The Female Pronucleus.
The vesicles formed from the chromosomes remaining in the egg
after the second polar body has been expelled, increase in size and
fuse together to form a single large one, the female pronucleus, which is at first irregularly lobed and elongated in a direction trans- verse to the main axis of the egg. ‘This fusion may occur while the vesicles are still small, or it may be delayed till shortly before the formation of the cleavage spindle. The following figures illustrate the process: Figs. 17, 19, 20 and 21.
The longer diameter of the pronucleus may exceed one-quarter of that of the egg. In the vesicles which form it, and in the pronucleus itself, there is a rather coarse network, which, however, does not stain deeply during these stages. At the junctions of the fibers of the network, and where they touch the nuclear membrane, are thick- enings due apparently to accumulations of the substance composing
the fibers. In addition to the network, there are in the fully formed _
pronucleus séveral nucleoli of unequal size. One of these usually originates in each vesicle, and as the latter become fused the nucleoli also increase in size and unite, so that in the completed pronucleus instead of ten nucleoli there are perhaps three or four quite large ones. Their staining power, which is at first considerable, meanwhile decreases so that they stain only with plasma stains. They are spherical and bounded by a distinct outline. From the way they fuse together they are probably of a viscous or semi-fluid nature. in the living egg.
A short time before the membrane of the pronucleus begins to disappear for the formation of the cleavage spindle the nucteoli vanish. They are sometimes vacuolated before the time of dis-
LR
a
W.-G: Van Name—Embryology of Eustylochus. 287
appearing ; in other preparations they become very pale and appear to dissolve away in the contents of the pronucleus.
These nucleoli seem to be waste matter, and resemble in every particular the nucleoli of the male pronucleus, and those of the nuclei of the cleavage stages.
While these changes have been going on, the female pronucleus has moved a little toward the center of the egg, keeping, however, on or near the line from the center to the animal pole. The aster of the second polar spindle, which remains in the egg, meanwhile degenerates, the centrosome and centrosphere disappear, and all that remains is a spherical mass of bluish stainy granules, apparently the microsomes of which the aster rays were made up (Figs. 19a, 20 and 21). This also moves to a more central position
in the egg. In the eggs of Planocera it is much less conspicuous
than in those of Hustylochus, where, however, it is much more noticeable in some preparations then in others. No yolk globules are found within the area occupied by this granular mass.
The Male Pronucleus.
I have given an account of the spermatozoon up to the time when the egg reaches the uterus. No change appears to take place in it until after the egg is laid. In a recently laid egg (Fig. 7) the dark colored area in the egg cytoplasm by which one end (probably the posterior, or middle piece) of the spermatozoon was surrounded, has disappeared and the spermatozoon has contracted into a shorter and thicker form, so that it is now spindle shaped. During the succeeding stages of the first polar spindle it contracts still more, becoming irregularly rounded or oval. It also loses its homogeneous appear- ance and becomes transformed into a vesicle in which a reticulum, and later one or more nucleoli similar to those in the female pro- nucleus, make their appearance (Figs. 18, 39 and 40).
It is generally still quite small at the end of the maturation period. In its final stages it appears to have precisely the same structure as the female pronucleus, but may generally be dis- tinguished by one or more of the following characteristics : its greater distance from the animal pole, more regular form, slightly smaller size, and smaller number of nucleoli. As a comparison of the above figures will readily show, its development does not necessarily keep pace with that of the female elements, but each develops independently of the other.
288 W. G. Van Name—Embryology of Eustylochus.
As has already been described, the spermatozoon quickly penetrates to a considerable distance from the surface of the egg and appar- ently remains practically stationary until maturation is complete. — When the first polar spindle has taken its final position and we can recognize the animal pole of the egg, we find that it more often lies in the hemisphere away from the animal pole and has evidently” entered at some point on the surface of that hemisphere. This state- ment however applies to Hustylochus only; in Planocera it is perhaps more frequent to find the sperm-nucleus in the upper hemisphere. In this species it usually lies closer to the center of the egg and con- sequently often very near the polar spindle aster (Figs. 38 and 40). This more central position is often noticeable even in uterine eggs (Fig. 36).
The movements which bring the pronuclei together seem to be chiefly movements directed toward the center of the egg, both on — the part of the male and the female. Just before the formation of the cleavage spindle we find them usually quite near together in the central part of the egg, the female nearer the animal pole.
The Polar Bodies.
During the pronuclear stages the polar bodies usually lie side by side, touching each other and the egg, showing that the point of separation of the second is nearly but probably not quite coincident with that of the first. There is generally considerable difference in the arrangement of the chromatin in the two polar bodies: In the first it remains in a compact group, which often appears to be com- posed of beaded threads. In the second, however, the rounded chromosomes are scattered irregularly in the cytoplasm of the polar body and are often widely separated. This difference is suggestive of, and doubtless largely due to, the form and arrangement of the chromo- somes in the later stages of the first and second polar spindles respectively. The first polar body rarely if ever divides.
The Sperm-centrosomes and their Asters.
These are shown in a number of my specimens of Planocera nebulosa, though the examples illustrated (Figs. 38, 39 and 40) are cases where they are visible with unusual clearness. I attribute my failure to find them in Hustylochus to the same causes which make them indistinguishable in many preparations of Planocera nebuldsa. The radiations are very few and short as well as very weak and
ee eile ed
A
W. @. Van Name—Embryology of Eustylochus. 289
developed during a brief stage only, and they are doubtless often obscured by the densely crowded and dark colored yolk spheres. The centrosomes and centrospheres moreover greatly resemble in size and staining qualities the yolk spheres with black centers which I have already alluded to.
In Planocera I have distinguished the sperm-centrosome with cer- tainty only in eggs in the late stages of the first polar spindle and in the second polar spindle stages. Earlier than this it is probably also visible, but until definite radiations develop I cannot certainly identify it. When the centrosome can be recognized it is still sin- gle (Fig. 38). It is surrounded by a distinct centrosphere and by short radiations which can be seen to be made up of microsomes even close up to the centrosphere. Subsequently the centrosome divides into two (Fig. 39) which separate, the centrosphere mean- while elongating (Fig. 40) and eventually dividing, forming two asters. Krom the position of the aster or asters, which is in nearly every case somewhere on the line which the spermatozoon has prob- ably travelled in penetrating to the position where we find it, they have separated from the spermatozoon at a considerably earlier stage. If this surmise is correct it may explain one specimen which for a time seemed very puzzling, in which the polar spindle lies between the spermatozoon and the sperm-aster. But if the sperm- centrosome separates from the nucleus before the polar spindle reaches its peripheral position, it is not unlikely that this would occasionally happen to move between the sperm-nucleus and _ its aster.
Van der Stricht found sperm-centrosomes and asters only in the eggs where the sperm-nucleus lay near the vegetative pole or near the center of the egg. Ido not find that the position of the sper- matozoon has anything to do with their presence or apparent absence. I have not succeeded in learning anything of the subse- quent history of these asters or centrosomes, nor demonstrated any connection between them and the cleavage centrosomes.
The First Cleavage Spindle.
For a lone time after the second polar body has separated the 8 , I granular remains of the inner aster of the polar spindle continue to be visible. It has already been described as an area free from yolk where the cytoplasmic microsomes, which stain blue with hama- toxylin, are particularly abundant and conspicuous (Figs. 19a and Trans. Conn. Acap., Vou. X. Avucusr, 1899,
19
290 W. G. Van Name—Embryology of Eustylochus.
20). These have fora time a suggestion of their former arrange- ment in radial lines (Fig. 19a).
This may have practically disappeared by the time the pronuclei have their final position, but there remains at least an area free from yolk the position of which (central to the female pronucleus) is
between the pronuclei when they approach each other, and in it the’
cleavage spindle appears.
Some of my specimens (Figs. 22 and 24 for example) plainly show that the spindle is at first short but afterwards increases in length. This indicates that the asters arise by the division of a single one. If we are ready to believe that the cleavage-centrosomes are derived from the spermatozoon, this would support Van der Stricht’s theory that the separation of the sperm-centrosome from the sperm-nucleus, and consequently its division into two, is often delayed until this stage.
On the other hand, my preparations of P/anocera indicate that the sperm-centrosome separates from the nucleus very early and that it divides during the maturation stages. I have not observed a case where the cleavage aster is still single. Fig. 22 shows the nearest approach to this condition that I have found. It also shows the first appearance of the cleavage centrosomes.
From their first appearance (Fig. 22) the asters are connected by a central spindle and the centrosomes are surrounded by centrospheres which become more definite in outline in the later stages. Beaded threads of chromatin, staining more deeply than the remainder of the network, appear within the pronuclei. The aster rays begin to penetrate the pronuclei, whose membranes begin to dissolve away in the vicinity of the centrosomes. The threads of chromatin break up into segments, the chromosomes of the cleavage spindle, which continue to shorten and thicken and finally assume the form of a rather short rod with smooth outline bent into the shape of a U or a V with the point rounded. Ten are formed from each pro- nucleus.
The spindle continues to grow, its aster rays and fibres increasing in number, If it did not at first lie transverse to the main axis of the egg, it rotates sufficiently to take such a direction, It usually lies almost in the center of the egg. The aster-fibres attach them- selyes to the chromosomes and draw them into the equator of the spindle, where they take a position with the free ends of the loops directed away from the axis of the spindle. They lie near the outer part or circumference of the equatorial plane of the spindle,
aaa 5
=
> 2 a
W. G. Van Name—Embryology of Lustylochus. 291
Through the center of this circle or ring of chromosomes a large number of central fibres pass directly from one aster to the other. Figs. 23, 24 and 25 show the successive stages in this process.
I have already described the way in which the nucleoli disappear. The portion of the reticulum of the pronuclei which is not to develop into the chromosomes disappears about the same time as the nuclear membranes.
In Hustylochus and Planocera the pronuclei ordinarily do not fuse, and their membranes may disappear while they are still quite far apart, so that the spindle contains for a time two well separated groups of chromosomes, derived from the egg and spermatozoon respectively (Fig. 24). At a later stage we can no longer dis- tinguish these groups. I have also found a few instances like the one mentioned by Francotte, where the chromosomes of one pro- nucleus developed and the membrane disappeared while the other pronucleus was still intact.
The minute structure of the centrospheres, aster rays and spindle fibres does not appear to differ from that already described in the polar spindles. In length the cleavage spindle does not exceed that which is often attained by the first polar spindle, but it is much thicker and the asters are generally larger.
When the chromosomes finally divide each splits lengthwise, form- ing two U-shaped pieces which are drawn toward the opposite poles of the spindle (Fig. 26). An indistinct cell-plate, consisting of a thickening of the central fibres at their middle point, is then gener- ally noticeable.
Meanwhile the centrosomes in each pole of the spindle have divided, the derived centrosomes usually separating in a direction transverse to the axis of the spindle (Fig. 26). They lie near the middle of the centrosphere, not toward the outside or part most removed from the equator of the spindle, and therefore much nearer to the chromosomes than occurs in Zhalassema as described by Griffin (12). I have not observed the small asters within the centro- sphere described by this author and by Coe (1). When the egg has divided, the chromosomes swell up into vesicles in which nucleoli and a reticulum appear, and the vesicles in each blastomere fuse into a resting nucleus, but the process so closely resembles the formation of the female pronucleus that its description and illustration would be superfluous,
292 W. G. Van Name—Embryology of Eustylochus.
The Second and Later Cleavages.
The conditions in the case of these planarians are particularly unfavorable for tracing the cleavage centrosomes through from one division to the next, for a stage intervenes when the aster rays of the preceding spindle have disappeared or practically so, and the new rays have not as yet developed. All that remains is an area of granular protoplasm lying against the nucleus, very similar in appearance to that found in the earlier stages already described, and there are generally black staining spots or granules present, making it impossible to recognize the centrosomes. By the time the new aster rays are visible these lie, if not on opposite sides of the nucleus, at least widely apart, and no central spindle can be seen connecting them. At an earlier stage such a spindle may exist, but the specimens upon which this opinion is based are too obscure to be of much value in proving it.
The chromosomes of the second cleavage spindle are formed in the same way as those of the first, except that they are all derived from the single nucleus instead of from the two pronuclei. A part of the nuclear reticulum begins to stain deeply, the nucleoli disappear and the membrane is dissolved as in the case of the pronuclei. In the subsequent phases the second cleavage spindle resembles the first.
As already mentioned, the second cleavage is somewhat unequal, and the third is still more so. Mitosis in the later stages seems to follow the same rules, but as the cells become smaller its observation presents more difficulties. In the late stages the centrosomes and centrospheres are larger and the spindle proportionately shorter and wider. This is already quite apparent in the third cleavage spindle shown in Fig. 29.
Later Development of the Embryo.
The development goes on both by the division of the small cells and by the budding off of more small cells from the large ones, but I have not attempted to work out the cell-lineage. Hustylochus isa most unfavorable animal for this kind of work because of the small differences in the size of the blastomeres, in which it is unlike some other polyclads, especially /iscocelis, described by Lang (18). The slowness of development and the opaqueness of the blastomeres add to the difticulty.
The result of the succeeding divisions is that a cap or envelope of small ectoderm cells extends further and further from the animal
W. G@. Van Name—Embryology of Eustylochus. 293
pole, and finally encloses the whole egg, the large cells of the lower hemisphere becoming enclosed within the ectoderm layer. In this manner a spherical embryo is formed which is really a modified gastrula.
The time occupied in developing up to this stage is variable. Eggs may occupy eight hours or more between the two and the thirty- two cell stage. In the course of a few hours more the cells become so numerous that it is difficult to count them. During the two or three days following the time when the ectoderm overgrows the egg the interior cells undergo a change, their outlines become less dis- tinct than those of the ectoderm cells, and nothing but the nuclei and yolk spheres can be distinguished in the living specimen. The consistency of the interior cells becomes less firm and they seem to be more or less free to move about.
As the development proceeds a number of large oval bodies of different sizes become conspicuous within the embryo. These are shown in the figures of the later stages (Figs. 32, 33 and 34). They were noticed by Girard, who considered them to be cavities. They are, however, solid bodies,—large cells from the lower hemisphere of the egg densely packed with yolk, which seems in some of them to fuse to a single large mass occupying most of the cell, like the fat in adipose tissue.
When four or five days old the embryo, still practically spherical, develops cilia, and begins to revolve within the egg-membrane. About the same time active contractile movements commence. As development proceeds it becomes somewhat elongated, and therefore is necessarily more or less bent or folded within the membrane. By this time the anterior and posterior ends may be readily distin- guished, whereas in the earlier stages this is a matter of great difficulty on account of the small difference in the size of the blastomeres. There is, however, no reason to doubt that this is already determined at a very early stage, as it is in polyclads like Discocelis, where the conditions for demonstrating it are not so un- fayorable. As the embryo lies in the egg the dorsal surface is the more convex; the ventral surface shows a transverse fold due to the bending of the embryo already mentioned. On the posterior ven- tral surface two longitudinal lobes appear, separated by a median groove or depression (Fig. 32).
About six days after the egg is laid a very dark brown pigment spot, or eye, becomes visible near the anterior end on the left side.
It is at first small and irregular in shape, though it afterwards in-
TRANS. Conn. AcaD., VOL. X. Aucust, 1899. 20
294 W. G. Van Name—Embryology of Eustylochus.
creases in size and becomes circular. It lies in or attached to the ectoderm but does not reach the surface. A day or so later another spot appears in a similar position on the right side. This also in- creases in size, but for many days that on the left side is conspieu- ously larger (Fig. 32). Occasionally an embryo presents an-abnor- mality in regard to the number and arrangement of the pigment spots, but so far as I have discovered, the earlier appearance and larger size of that on the left side is invariable. Except for this the embryo is bilaterally symmetrical, if we leave out of account the large yolk-masses which change their position as the animal turns and contracts.
Later the surface of each eye-spot becomes concave, and in the concavity a transparent vesicle is developed.
By the time the eyes have appeared, the ectoderm in the depres- sion between the ventral lobes, mentioned above, can be seen to be deeply bent inward into a funnel-shaped cavity extending upward and somewhat forward. At the anterior end of the embryo, almost between the eyes, a few stiff hairs grow out in a group and generally adhere so closely together that the group appears like a single coarse flagellum. Later a similar, but at first shorter, group appears at the posterior end. All these structures are developed by the time the embryo escapes from the egg-membrane. They can, however, be much more easily observed after this has occurred, which may be anywhere from ten days to two weeks after the egg is laid, or ocea- sionally longer. Toward the last the egg-membrane becomes very thin, and with the most careful handling many embryos will be pre- maturely released. For several days before the embryo escapes its movements are very active, and if the membrane is torn it swims rapidly away by means of the cilia. Contrary to what Girard says, I find that the embryo almost always swims with the anterior end directed forwards, but it keeps turning about its longer axis so that the ventral surface is often upward.
Fig. 32 shows an embryo which was drawn as it was endeavor- ing to escape through a small hole torn in the egg-membrane, and Fig. 33 one in about the same stage removed from the membrane and seen from one side. These were about eight days old. But little further change takes place up to the time of hatching. In the course of the first few days after hatching the embryos assume the curious form shown in Fig. 34, though there does not seem to be any considerable change in the internal structure during the interval. Fie. 85 shows a still later stage; the external form is similar to that
W. G. Van Name—Kimbryology of Eustylochus. 295
in Fig, 34; the posterior part of the body is, however, much more prominently developed than in the earlier stage, and now forms a distinct posterior median lobe. The large yolk-cells have by this time become divided up, and most of the yolk has evidently been absorbed, but some dark colored substance is left which may be a remnant of it.
The invagination between the posterior ventral lobes has also become wider and more conspicuous, but I have not been able to find at this stage any internal cavity with which it might communi- cate. I have never seen one of the embryos eat anything, neither will they ingest powdered carmine.
Their bulk has not changed appreciably from that of the egg, and so far they have been living on the yolk stored up within the cells. -When this is gone (as in the stage shown in Fig. 35) the embryo soon dies. I have never been able to raise them any farther. If they live longer they do not develop, but degenerate and lose their activity. I fourd no indications of the peculiar chrysalis stage which Girard described, and am convinced that he mistook some other organism present with the embryos for a stage of the latter. As might be expected, the development of Hustylochus resembles closely that of the related polyclad Stylochus pilidium, described by Goette (9, 10, 11) and Lang (18).
The later development of Planocera nebulosa does not differ much from that of Hustylochus ellipticus. In the early cleavage stages there is a little more difference in the size of the blastomeres. The embryos are of course a little larger and the development is some- what slower. I have kept them for nine or ten days before the eyes began to appear. The left eye develops first, as in Hustylochus.
In conclusion I wish to express my gratitude to Dr. Wesley hk. Coe of the Sheffield Biological Laboratory, not only for the material for which I am indebted to him, but for constant advice and assist- ance during the course of the work, and especially for the personal interest he has taken in it. My thanks are also due to Prof, A. E. Verrill for valuable advice in regard to the illustrations and to Prof. Sidney I. Smith, in whose laboratory the work was done.
Sheffield Biological Laboratory of Yale University, June, 1899,
296 W. G. Van Name—Embryology of EHustylochus.
LITERATURE.
1. Coz, W. R.—The maturation and fertilization of the egg of Cerebratulus. Zool. Jahrbiicher. Vol. 12, 1899. 2. Fiemminc, W.—Neue Beitriige zur Kenntniss der Zelle. Arch. f. mik. Anat,. Vol. 29, 1887. 3. Francorre, P.—Recherches sur la maturation, la fécondation et la segmenta- tion chez les Polyclades. Mém. cour. Acad. Roy. de Belgique, 1897. 4. Francorre, P.—Recherches sur la maturation, la fécondation et la segmenta- tion chez les Polyelades. Arch, de Zool. Exper, 3d series. Vol. vi. No, 2, 1898. . Fuerst, E.—Ueber Centrosomen bei Ascaris megalocephala. Arch, f. mik, Anat. Vol. 52, 1898. 6. GarpInER, E. G.—Early development of Polychoerus caudatus Mark. Journ. of Morph. Vol. xi, No. 1. r . Girarp, C.—On the development of Planocera elliptica. Proc. Boston Soc. Nat. Hist. Vol. iii, 1848-1851. 8. Grrarp, C.—Embryonic development of Planocera elliptica. Journ, Acad, Nat. Sci. Philadelphia. Second series. Vol. ii, 1854. 9. Gorrre, A.—Zur Entwickelungsgeschichte der Seeplanarien. Zool. Anzeiger,
ol
-~)
1878. 10. Gorrre, A.—Zur Entwickelungsgeschichte der Wiirmer. Zool. Anzeiger, 1881.
11. Gorrre, A.—Abhandlungen zur Entwickelungsgeschichte der Thiere, — Leipzig, 1882.
12. Grurrin, B. B.—History of the achromatic structures in the maturation and fertilization of Thalassema. Trans. N. Y. Acad. Sci. June, 1896.
13. Hatiez.—Contribution & histoire naturelle des Turbellariés. Lille, 1879.
qe wa aaa.
14. Kurnckowstroem, A. v. — Beitraege zur Kenntniss der Eireifung und Befruchtung bei Prostheceraeus vittatus, Arch. f. mik. Anat. Vol. 48, 1897.
15. Korscue rt, E.—Mittheilungen ueber Eireifung und Befruchtung. Verh. der Deutéchen Zool. Gesellschaft: 1895.
16. KorscueLt, E.—Ueber Kerntheilung, Eireifung und Befruchtung bei Ophryotrocha puerilis. Zeitschrift fiir wiss. Zool. Vol. 60, pt. 4. 1899.
17. Kosraneckt, K. and Srepteck1, M.—Ueber das Verhiltniss der Centrosomen zum Protoplasma. Arch, f. mik. Anat. Vol. 48, 1896.
18. Lane, A.—Fauna und Flora des Golfes von Neapel. Die Polycladen. 1884.
19, Linum, F. R.—Centrosome and sphere in the egg of Unio. Zool. Bull. Vol. 1, No. 6, 1898.
2), MacFarianp, F. M.—Celluliire Studien an Mollusken-Kiern. Zool. Jahr- biicher. Vol. 10, 1897.
21, Sevenka, E.—Ueber eine eigenthiimliche Art der Kernmetamorphose. Biol. Centralblatt, Vol. i, 1881-1882.
22. VAN DER Srriowt, O.—Contribution d l'étude de la sphére attractive, Arch, de Biolog. TT, xii, 1892,
Oe Ss eee eee ee ee ee
W. @. Van Name—Embryology of Eustylochus. 297
23. Van DER Srricut, O.—Origine des parties constituantes de la figure achro- matique de Thysanozoon Brocchi. Verhandl. der Anat. Gesellsch. 1894,
24. Van per Srricut, O.—Le premier amphiaster de l’ovule de Thysanozoon Brocchi. Une figure mitosique peut-elle rétrograder? Bibliographie ana- tomique. No. 1, 1896,
25. VAN per Stricut, O.—Anomalies lors de la formation de l’amphiaster de rebut, Bibliographie anatomique, No. 1, 1896.
26. Van preR Strricut, O.—La maturation et la fécondation de l’ceuf de Thy- sanozoon Brocchi. Associat. frangaise pour l’avancement des sciences, Congrés de Carthage, 1896.
27. VAN pER Srricut, O.—Les ovocentres et les spermocentres de Vovule de Thysanozoon Brocchi, Verhandl. d. Anat. Gesellsch. in Gent. 1897,
28. VAN DER Stricat, O.—La formation des deux globules polaires et l’appara- tion des spermocentres dans l’ceuf de Thysanozoon Brocchi, Archives de Biologie. Vol. xv, pt. 3, Oct., 1898.
29. VeRRILL, A. E.—Marine Planarians of New England. Trans. Connecticut Acad. Vol. viii, 1893.
30. WHEELER, W. M.—The maturation, fertilization and early cleavage of Mysos- toma glabrum Leuckart. Archives de Biologie. Vol. xv, 1897.
31. Witson, E. B. and Leamine, E.—An atlas of the fertilization and karyo- kinesis of the ovum. 1895.
32. Van DER Stricut, O.—Etude de plusiéres anomalies interessantes lors de la formation des globules polaires. Etude de la sphére attractive ovulaire & état pathologique, etc. Livre jubilaire dedié a Charles Van Bambeke, Brussels. 1899.
EXPLANATION OF PLATES.
With the exception of Figs. 30, 31, 41, 42 and 43, all the figures were drawn with the aid of a camera lucida.
In illustrating the development of the egg, I have avoided making up figures by combining into one drawing structures which appear in different sections in the preparation, preferring to reproduce more than one section where this is necessary.
The preparations from which the figures of sections were drawn were made with corrosiye-acetic solution and stained by Heidenhain’s iron-hematoxylin method.
To this Figs. 1 to 5 inclusive, also 11, 25, 26, 27 and 56 are exceptions. These were preserved with picro-acetic solution, which, as mentioned in the descriptive part, renders the yolk globules inconspicuous, but brings out the reticulum of the cytoplasm strongly.
Any other exceptions are mentioned in describing the individual figures.
All represent the eggs or embryos of Eustylochus ellipticus (Girard) Verrill, unless otherwise mentioned,
298 W. G. Van Nume—Embryology of Lustylochus.
PLATE XXXVI. Fig. 1.—Young ovarian egg, showing the large germinal vesicle, the nuclear reticulum and large nucleolus, Enlarged 860 diameters. Fig. 2.—Ripe egg in which the first polar spindle asters are visible, though they do not show the centrosomes, Preparation stained with Delafield’s hama-
toxylin. Most of one aster lies in another section, The nucleolus has dis-. _
appeared, and the beaded threads of chromatin haye become conspicuous, x 860.
Figs, 3a and 3b.—Sections of the same egg, which is in a later stage than fig. 2. Shows the polar spindle centrosomes and the rounded chromosomes, one of which in fig. 8a has already opened into a ring. x 860.
Fig. 4.—Slightly later stage. Somewhat over-stained, so that the centrosomes appear very large. Their elongated form indicates their approaching divi- sion. x 860.
Fig. 5.—Uterine egg showing polar spindle and spermatozoon. The centrosomes have already divided. The centrosphere has acquired a distinct outline. x 860.
Fig. 6.—Spermatozoon (from an egg in the stage of fig. 5), one end of which is surrounded by the dark area which is less distinctly seen in fig. 5. x 860.
Fig. 7.—Recently laid egg. The polar spindle is still in a central position, The spermatozoon has contracted into the shape of a spindle. x 860.
Fig. 8.—First polar spindle after it has reached the surface of the egg. This specimen is unusual, as all the chromosomes still have the ring form. In this stage some of the rings at least have usually broken. x 1240.
Norr.—Figs. 5, 7 and 8, as well as some that follow, illustrate the separation of the derived centrosomes, which in this species generally occurs in a longitu- dinal direction, parallel to the axis of the spindle or nearly so.
Figs. 9 and 10.—Chromosomes of eggs in the same stage as fig. 8. x 1240.
PLATE XX XVII.
Fig. 11.—First polar spindle, introduced to show the chromosomes. x 1240.
Fig. 12.—Later stage of the first polar spindle. The centrosomes of the outer pole of the spindle, which is somewhat over-stained, lie in an oblique instead of the usual axial position. The centrosomes of the inner pole have begun to separate. x 860.
Fig. 13.—Formation of the first polar body. x 860.
Figs. 14, 15, 16.—Three stages of the second polar spindle, showing the reduc- tion of the aster rays in strength and number, and the formation of the new spindle from the centrosphere. In the earliest of these (fig. 14), the chromo- somes have already split into two segments. The chromosomes are more widely separated from each other than in the first polar spindle. Figs. 14 and 15 x 1240. Fig. 16 x 860.
Puats XXX VIII.
Fig. 17.—Final stage of the second polar spindle, showing the cell-plate. Most of the second polar body is in the next section, The chromosomes have swelled up into vesicles. x 1240,
W. G. Van Name—Embryology of Eustylochus. 299
Fig. 18.—Sperm-nucleus from an egg in the stage of fig. 17. x 860. Figs. 19a@ and 19b.—Somewhat later stage than fig. 17. The polar spindle aster
has degenerated, its centrosome and centrosphere have vanished, and the rays become very indistinct. The vesicles are larger and each contains a nucleolus. The sperm-nucleus and several of the female vesicles appear in fig. 196, which is another section of the same egg. x 860.
Figs. 20 and 21.—Show the fusion of the vesicles into the female pronucleus, the
remains of the second polar spindle aster and the male pronucleus. The latter lies unusually far from the center of the egg in fig. 20. x 860.
Norre.—Figs. 19 and 20 show the difference in the condition of the chro-
matin in the first and second polar bodies. In fig. 19a the first polar body lies on the right, in fig. 20, on the left.
Fig. 22.—Very early stage of the first cleavage spindle, showing the first appear-
ance of the cleavage centrosomes. The female pronucleus, which is three- lobed, lies above; the male, below and to the right. Threads of chromatin, which will develop into the chromosomes of the cleavage spindle, are begin- ning to be noticeable. = 860.
PLATE XX XIX.
Fig. 23.—A rather early stage of the first cleavage spindle: The threads of
chromatin can be seen in the pronuclei, but their membranes are still com- plete or nearly so. The centrosomes are not stained. x 860.
Fig. 24.—First cleavage spindle, shortly after the disappearance of the pronu-
clear membranes, which have dissolved while the pronuclei were still some distance apart. Two groups of chromosomes are shown, derived from the male and female pronucleus respectively. The centrosomes are not stained and the centrospheres are poorly preserved, showing an outer ring not normally present. x 860.
. 25.—First cleavage spindle fully formed. x 860. . 26.—Late stage of the same showing the centrosomes of one pole only.
The single centrosome shown in each pole in fig. 25 has divided into two, which separate in a direction transverse to the axis of the spindle. The centrosphere is more differentiated than in the earlier cases, as it is in the later stages of the polar spindles also. x 860.
Figs. 27a and 27b —Early stage of the second cleavage spindles. One aster,
Fig.
lying in the next section, is shown in fig. 270. x 860.
PLATE XL.
28.—Second cleavage spindles. The U-form of the chromosomes can be seen in the right hand blastomere. The centrospheres exhibit the same defect described in fig. 24. x 860.
Fig. 29.—Two cells of an egg in the four cell stage with third cleavage spindles.
x 860.
Fig. 30.—Process of forming the first polar body. Outlines of the forms
assumed by the polar body in one instance in the course of ten minutes.
Fig. 31.—Outline of a spermatophore. The small part only is inserted through
the opening made in the epidermis.
300 W. G. Van Name— Embryology of Eustylochus.
Fig. 32.—Embryo about eight days old endeavoring to escape through an open-_ ing accidentally torn in the egg membrane. Ventral surface of embryo shown, x 285.
Fig. 38.—Embryo about eight days old, prematurely hatched by tearing the egg membrane. x about 285.
Fig. 34.—Embryo a few days after hatching. x about 285.
Fig. 35.—Embryo a week after hatching. x about 285.
Puate XLI.
Fig. 36.—Planocera nebulosa, Uterine egg showing a first polar spindle. This
is peculiar in that all the chromosomes have already assumed the elongated —
form, the rings having broken at one point. A part of the spermatozoon is seen to the right of the spindle. x 860.
Fig. 37.—Planocera nebulosa. Beginning of the second polar spindle. In this —
specimen the centrosomes are separating in a direction nearly transverse to the axis of the spindle. x 860.
Fig. 38.—Planocera nebulosa. Egg in a little earlier stage than the last, yet the 7
second polar spindle is further advanced. Near it is the sperm-nucleus, in
this case rather larger than usual at this stage, and in the upper right hand
part of the section the sperm-aster, whose centrosome appears very large owing to insufficient extraction of the stain with the alum solution. x 860.
Fig. 89.—Planocera nebulosa. Sperm-nucleus and aster from an egg in the second polar spindle stage. The centrosome has just divided, and the cen- trosphere has become elongated. x 1240.
Fig. 40.—Planocera nebulosa. Second polar spindle more advanced than in fig. 38. Most of the chromosomes lie in another section. Near the spindle the sperm-nucleus, and to the left, near the edge of the section, the sperm-aster is shown. The latter has proceeded somewhat farther in its division than the one shown in fig. 39. x 860. 7
Notr.—Figs. 36, 38 and 40 show the position of the spermatozoon and later the sperm-nucleus very near the center of the egg and the polar spindles, which is so common in this species as to be quite characteristic.
Fig. 41.—Planocera or Eustylochus. Diagram showing successive stages in the division of a chromosome of the first polar spindle,
Fig. 42.—The same, showing another method of division,
Fig. 43.—Diagram showing different degrees of modification (not different stages, as in the last two figures) of one of the forms of chromosome of the first polar spindle.
Fig. 44.—Kustylochus ellipticus. Entire egg in two cell stage. 285. :
Figs. 45 and 46.—Same in four and eight cell stages respectively, seen from the vegetative pole. x 285.
4
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