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West Indian, Central American, and European Miocene and Pliocene M0llusks 1

West Indian, Central American, and European Miocene and Pliocene M0llusks 1

BULLETIN OF THE GEOLOGICAL SOCIETY OF AMERICA VOL. 35. PP. 867-886 DECEMBER 30. 1924 PROCEEDINGS OF THE PALEONTOLOGICAL SOCIETY

WEST INDIAN, CENTRAL AMERICAN, AND EUROPEAN AND M0LLUSKS 1

BY WENDELL P. WOODRING

(Read before, the Paleontological Society December 27, 1923)

CONTENTS Page Introduction...... : ...... 867 Composition of West Indian and Central American Miocene and Pliocene faunas...... 869 Composition of south European Miocene and Pliocene faunas...... 871 Piedmont basin...... 871 ...... 875 Comparison of West Indian and Central American faunas with European faunas...... 877 Miocene...... 877 ...... 877 ...... 879 Helvetian...... 881 ...... 884 Pliocene...... 885

I ntroduction The Miocene and Pliocene faunas of the West Indies and Central America should be compared only with European faunas that lived under essentially similar conditions. These American faunas, with the excep­ tion of a supplementary Mexican fauna, are almost purely tropical, ac­ cording to the distribution of their living genera. The Miocene and Pliocene faunas of the Mediterranean region and the Miocene faunas of Aquitaine, which most closely resemble these American faunas, include

1TMs paper is one of the composing a “Symposium on the correlation of the formations of southeastern North America, Central America, and the West Indies with the Tertiary formations of Europe.” Published by permission of the Director, U. S. Geological Survey, and of the Presi­ dent of the Carnegie Institution of Washington. Manuscript received by the Secretary of the Society September 2, 1924.

- L V I — B u l l . G e o l . S o c . A m ., V o l . 35, 1923 (867)

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the following temperate genera:2 Mytilus (MyUlus), Astarte, Cardium (C erastoderma), Macoma (Macoma), Solen (Sole'n), Ensis, Saxicava, and Aporrhais. None of these genera are in any of the West Indian Tertiary or living shallow-water faunas. All except Aporrhais are com­ mon in the Tertiary and living faunas of the Atlantic coast of the United States. Aporrhais has not been found in the Tertiary faunas of the

F ig u r e 1.—Graph showing approximate Composition of West Indian and Central American Miocene and Pliocene Faunas

Atlantic States, but is living in temperate waters on both sides of the Atlantic. Although these European faunas are not strictly tropical, they resemble the West Indian faunas in many features. So far as tempera­ ture is concerned, they are comparable in the same sense that the West Indian faunas are comparable to a subtropical fauna on the American mainland.

2 In this paper genus is used instead of continually repeating genus, subgenus, and section.

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All the West Indian and European faunas considered in this paper are essentially shallow-water faunas. Slight recognizable differences in depth are disregarded. The presence of some genera, such as Tindaria, Bathyarca, and Seguenzia, which are persistent deep-water dwellers in the present seas, is puzzling. Collections of predominantly shallow-water mollusks including these genera represent different bathymetric zones, or during Tertiary time they lived in shallower water. Bottom conditions are disregarded because not enough is known about them for the Tertiary faunas. Faunas of 500 or 600 species, like the Miocene West Indian faunas, or of 1,000 or more species, like those from the Miocene and Pliocene of the Mediterranean region, probably are not homogeneous. It seems unreasonable to believe that so many species could live on the same kind of bottom. Large collections from even one locality may include specimens washed from different kinds of bottoms. Superspecific group—genera, subgenera, and sections—for convenience called genera, are used in comparing the West Indian and south Euro­ pean faunas. The first appearance of a genus in the two regions is a significant feature, provided there is a possibility of free migration and a suitable habitat. Even when unrestricted migration is possible, the dispersal of a genu3 depends on the length of the swimming larval . Many genera of marine mollusks have no swimming larval stage and their distribution is essentially provincial.. This feature has been dis­ cussed by Doctor Vaughan in the introductory paper of this symposium.

C o m p o s i t i o n o f W e s t I n d i a n a n d C e n t r a l A m e r i c a n M i o c e n e

a n d P l i o c e n e F a u n a s The percentage composition of the West Indian and Central American Miocene and Pliocene faunas, based on the genera, is graphically repre­ sented in figure 1. The relative time spacing in the graph is arbitrary. Several families of gastropods—Turritidæ (=Pleurotomidæ), Epito- miidæ (=Scalidæ), Melanellidæ, and Pyramidellidæ—and the scapho- pods were not used in preparing the graphs, as their nomenclature is too chaotic. The graph is only a crude approximation of the actual compo­ sition of the faunas. Most of them have not been adequately studied and not all are comparable in size or in living conditions. Despite these unfavorable features, the graph clearly shows the principal elements of the faunas. It shows that they are predominantly West Indian. Even in the oldest fauna, 84.6 per cent of the genera are how living in West Indian waters. The percentage of living West Indian genera steadily; but not uniformly, increases as the decreases.

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The relatively large percentage of genera now living in the Pacific or the Indo-Pacific region is a striking feature of the Miocene faunas. These genera are as follows: Nucula (Acila) Peristernia Placunanomia Solenosteira Julia Metula Cardium (Lophocardium) Northia Clementia Columbella (Meta) Cooperella Strombina Mactra (Harvella) Oymia Jouannetia Malea Terebra (Terebra) Pustularia Cancellaria (Aphera) Cirsochilus Cancellaria (Narona) The percentage of these exotic genera decreases rather uniformly, but there is an apparent increase in the fauna of the Gatun formation of the Canal Zone and Costa Rica. If this actually is an increase, its signifi­ cance is not known. The small mollusks of the Gatun formation have so far been neglected. It may confidently be predicted that when they are studied the percentage of exotic genera will decrease, as most of the exotic genera are large. The percentage of extinct genera also decreases rather uniformly, except for the fauna of the Bowden formation of Jamaica, which has been more exhaustively studied than any other large fauna shown in the graph. The extinct genera, exclusive of new extinct genera in the Bow­ den formation, are as follows Spirulirostra Strombinella Barbatia (Obliquarca) Metulella Noetia (Sheldonella) Orthaulax Cuspidaria (Bowdenia) Solariorbis Neseromya Ampullina Alveinus Ampullinopsis Corbula (Bothrocorbula) Genera that are not living in West Indian waters, though not confined to the Pacific (other exotic genera of figure 1), are a minor feature. They include Cardita (Cardita), Cardium (Cardium), Halia, and Euthria. These exotic genera are common fossils in the Mediterranean Miocene and Pliocene deposits. According to available records, the following living American genera have not been discovered in West Indian waters:

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Venericardia (Pleuromeris) Alectrion (Tritia) [called Nassa Montacuta (Uzita) by some Europeans] Sportella Sycotypus Amauropsis

Only one of these genera—Amauropsis—is not recorded from the Gulf of Mexico or the east coast of Florida, and it alone detracts from the tropical aspect of the Tertiary faunas. The percentage of exotic and extinct genera is significant in determin­ ing the age of a West Indian Miocene or Pliocene fauna. This standard can be used only for large faunas. If faunas that lived under similar conditions were completely studied they probably would show smoother curves in a graph like figure 1. Under these favorable, but perhaps unattainable, conditions it might be possible to determine the age of a fauna by simply plotting the percentage composition.

C om po sitio n of S ou th E uropean M io cen e and P lio cen e F aunas

PIEDMONT BASIN The faunas of the richly fossiliferous beds in the classic Piedmont basin of are used as representative of the Mediterranean. Miocene and Pliocene. The type localities of the Tortonian, Plaisancian, and Astian stages are in the Piedmont basin. The' mollusks from these de­ posits have been described in the monumental series of monographs begun by Bellardi and completed by Sacco.3 The marine deposits carry­ ing large shallow-water faunas are of Helvetian, Tortonian, Plaisancian, and Astian age. The Aquitanian deposits of the Venetian basin (Schio beds)4 are used to supplement the Miocene faunas of the Piedmont basin. Apparently there are no large shallow-water Burdigalian faunas in these two Italian basins comparable to the West Indian faunas. Figure 2 is a graphic representation of the percentage composition of the faunas from the Piedmont basin. This graph shows remarkably smooth curves, although the data on which it is based are less reliable than in figure 1, as I naturally do not know the European faunas so well as the West Indian. The curves between the Astian stage and the living fauna would be more irregular if the faunas of the Sicilian, Milazzian, Tyrrhenian, and Monasterian-stages were plotted.

3 L. Bellardi and F. Sacco : I molluschi del terreni terziarii del Piemonte e della Liguria, 30 parts. Torino, 1872-1904. Bellardi is the author of the first five parts and Sacco of all the others. 4 P. Oppenheim : Ueber die Ueberkippung von S. Orso, das Tertiär des Tretto und Fauna wie Stellung der Schioschichten. Deutsch. Geol. Gesell. Zeltschr., vol. 55, 1903. pp. 98-235, pis. 8-10.

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The most striking feature shown by figure 2, as compared to figure 1, is the large percentage of exotic genera. These Miocene and Pliocene faunas are very different in some features from the living Mediterranean fauna, as has been known ever since the Mediterranean Tertiary faunas

F ig u r e 2.— Graph showing approximate Composition of Miocene and Pliocene Faunas of Piedmont Basin, Italy

were first studied. The has had a complicated geo­ logic history and its living fauna is due chiefly to invasions of a tem­ perate fauna from the Atlantic in Quaternary time. Some genera of the displaced Tertiary fauna are now living along the west African coast; others are living in the West Indian region; others are confined to the

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Pacific and Indo-Pacific regions. The percentage of exotic genera de­ creases steadily, but not uniformly, as the age decreases. The graph shows that the Mediterranean Miocene and Pliocene faunas have a relatively large percentage of genera that are not living now in any European or west African waters, but are living in the West Indian region. These genera, some of which are living also in the Pacific or Indo-Pacific region, are as follows: Leda (Jupiteria) Phos. Neilo Xancus (called Turbinella by most Neilo (Neilonella) Europeans) Tindaria Coralliophila Amusium Ancillaria Plicatula Lyrla Ouspidaria (Cardiomya) Columbella (Conidea) Deuteromya (= Dimya) Sconsia Modiolus (Brachydontes) Morum (Onlscidia) Phacoides (Cardiolucina) [included Ficus (called Ficula or Pyrula by in Phacoides (Oavilucina) by Amer­ most Europeans) icans] Terebra (Fusoterebra) Phacoides (Pleurolucina) [Phacoides Conus (several groups) (Here) miobarbieri Sacco probably Erato is a Pleurolucina] Rissoina (Rissolina) Protocardia (Nemocardium) Rissoina (Zebinella) Rocellaria (Spengleria) [= Gastro- Vanikoro chsena Spengleria)] Xenophora (Tugurium) M artesia Neritina (Puperita) Engina Modulus Distorsio (called Persona by most Pleurotomaria Europeans) Zeidora Bursa (called Apollon by most Euro­ Subemarginula peans) Rimula Gutturnium (called Ranularia by most Sabatia Europeans)

A smaller percentage of genera are now living in the West Indies and along the west African coast, but not in European waters. Some of these genera are living also in the Pacific or Indo-Pacific region. They are as follows: Isognomon (usually called Perna or d iv ella Pedalion) Latirus Crassatellites (called Crassatella by Cassis most Europeans) Conus (probably several groups) Phacoides (Linga) [included in Pha­ Strombus coides (Here) by Americans] Cancellarla (Trigonostoma) Codakia (Codakia) Seila

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Metis (called Oapsa by most Euro- Amalthea- peans) Cheilea (called Mitrularia by most Murex (Pteronotus) Europeans) Melongena Nerita Oliva Acteocina As most of the genera in the two preceding lists are common West Indian Miocene fossils, the faunas from these two regions are similar. So far as these numerous genera are concerned, the Mediterranean Mio­ cene and Pliocene faunas are more similar to the living West Indian fauna than to the living Mediterranean. The following genera are living on the west African coast. Some of them range northward to Spain and the Azores; others are living also in the Pacific or Indo-Pacific region: Vitularia Tympanotomus Pusionella Mesalia Halia Protoma Cancellaria (Solatia) Oxystele H arpa The Mediterranean faunas also resemble the West Indian faunas in having genera that are now confined to the Pacific or Indo-Pacific region. The percentage of these exotic genera is higher than in the West Indian faunas that seem to be of the same age. The Pacific and Indo-Pacific genera are as follows: Septifer Rostellaria Discors Rimella Isocardia (Miocardia) Pustularia Cardilia Erato (Eratopsis) Jouannetia Cancellaria (Scalptia) Clavellitlies (called Clavella by most Cancellaria (Merica) Europeans) Cancellaria (Ventrilia) Cominella Cancellaria (Ovilia) Volema (called PugiUna by most Eu- Cancellaria (Aphera) ropeans) Cancellaria (Massyla) Latrunculus Terebralia Iopas Telescopium Acanthina Stossichia Tudicla (Zaria) Latirus (Dolicholatirus) Delphinula Mitra (Mitra) Cirsochilus Mitra (Thala) Scutus Malea Scurria Comparison with the list on page 870 show that only 5 of these genera—Jouannetia, Malea, Pustularia, Cancellaria (Aphera), and Cir­ sochilus—also occur in the West Indian Miocene deposits.

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The percentage of extinct genera is greater than in the West Indian faunas. This may be partly due to the more refined separation of super­ specific groups by Sacco. The percentage for the West Indian faunas will increase, however, when they are as completely studied as the Mediterranean faunas. AQUITAINE BASIN The type localities of the Aquitanian and Burdigalian stages are in Aquitaine, in southwestern France. This basin also contains deposits of Helvetian and Tortonian age. The molluscan fauna of these 3 deposits is being monographed 1.a s § by Cossmann and Peyrot.6 As Í 1 1 their description of the gastro­ I' pods is not yet complete, only the pelecypods have been used in making comparisons with the West Indian faunas. The composition of the Mio­ cene faunas of Aquitaine is graphically shown in figure 3. z Although figure 2 is based on

actual increase, as it is only a F i g d b e 3.— Graph showing approximate Gorn- thirdwmu as lars-elarge as tnethe otners other* anaand BaHn> Franceof the (hased Miocene on FaunasPelecypoas) of Aquitaine therefore is hardly comparable. According to the graphs, the Helvetian fauna of Aquitaine has a smaller percentage of exotic genera than the Helvetian fauna of the Piedmont basin. Some of this difference probably would be eliminated if the graphs were based on similar data. The exotic genera in the faunas of Aquitaine are listed in the follow-

5 M. Cossmann and A. Peyrot: Conehologie ngoginique de l’Aquitaine, vol. 1, 718 pp., 28 pis., 3 maps, 135 text figs., Bordeaux, 1909-1912 ; vol. 2, 496 pp., 26 pis., 25 text figs., 1912-1914; vol. 3, 709 pp., 17 pis., 70 text figs., 1917-1919; vol. 4, pt. 1, 321 pp., 7 pis., 1922.

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ing lists. Genera living in the West Indies but not in European waters are as follows: Tindaria Erycina (Erycina) Neilo Aligena Pleurodon (called Nuculina by Coss­ Phacoides (Cardiolucina) [included mann and Peyrot) in Phacoides (Cavilucina) by Amer­ Noetia (Included in Anadara by Coss­ icans] mann and Peyrot) Protocardia (Nemocardium) Amusium Siliqua c Plicatula Basterotia Deuteromya ( = Dimya) M artesia Carditopsis a Rocellaria (Spengleria) [= Gastro- Miltha b chsena (Spengleria)] Dlplodonta (Felaniella) a Bahamas and Bermuda. 6 Brazil. o Atlantic coast of the United States.

The following genera are living in West Indian and west African waters, but not in European waters. Some of these genera, like some in the preceding list, are living also in the Pacific or Indo-Pacific region: Isognomon (usually called Perna or Phacoides (Linga) [included in Pha- Pedalion) coides (Here) by Americans] Crassatellites (called Crassatella by Tivela most Europeans) Metis (called Capsa by most Euro­ Codakia (Codakia) peans) Petricolaria Genera living along the west African coast, but not in the Mediter­ ranean Sea or in the West Indies, are as follows: Ungulina Jouannetia (Triomphalis) Tugonia The following genera are confined to the Pacific and Indo-Pacific; regions: Trisidos (called Paralelipipedium by Trapezium most Europeans) Circe Julia Sunetta (Solanderina) Septifer Phylloda Galeomma (Thyreopsis) Jouannetia Discors Only 5 (Neilo, Pleurodon, Aligena, Rocellaria (Spengleria), and Petricolaria) of the 25 genera in the first two of the preceding lists have not been discovered in Miocene deposits of the West Indies. Julia and

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Jouannetia, two of the Indo-Pacifie genera, occur in the West Indian Miocene. The preceding European faunas include the type faunas of the Aqui- tanian, Burdigalian, Tortonian, Plaisance, and Astian stages. In intro­ ducing the term Pontilevian for the littoral Helvetian deposits of Touraine, Dollfus and Dautzenberg6 reject Helvetian because as orig­ inally used by Mayer it included all the Miocene. Although this objec­ tion is based on valid grounds, the term Helvetian has been restricted and has a definite age significance.

Com parison of W est I n d ia n and Central A m erica n F aunas w it h E uropean F aunas

MIOCENE Aquitanian,—No large fauna of supposed Aquitanian age has been dis­ covered in the West Indies or Central America. Collections in.the United States National Museum from the upper part of the of the Canal Zone, which is supposed to be Aquitanian, have not been described. Vaughan7 has published an incomplete list of mol­ lusks from these deposits, and -Brown and Pilsbry8 described 5 species. These published lists give a wholly inadequate idea of the Culebra fauna. The Emperador limestone of the Canal Zone also is considered of Aquitanian age. It has a relatively meager molluscan fauna, like other coralliferous limestones, consisting principally of Pectens. Brown and Pilsbry9 have described most of the Emperador mollusks. Vaughan10 has listed a few additional undetermined species. Deposits of supposed Aquitanian age so far examined in the West In­ dies are coralliferous limestones. Their molluscan fauna is in no way comparable to the Aquitanian fauna of Aquitaine, which contains 219 species of pelecypods. The largest fauna from any of the West Indian deposits, obtained from the Anguilla formation of the Island of Anguilla,

8 G. F. Dollfus and P. Dautzenberg : Conchyliologie du Miocène moyen du bassin de la Loire. Soc. géol. France Mém. Paléontologie, no. 27, 1902, pp. 7-8. 7 T. W. Vaughan : The biologie character and geologic correlation of the sedimentary formations of Panama in their relation to the geologic history of Central America and the West Indies. U. S. Nat. Mus. Bull. 103, 1919, pp. 552-553. 8 A. P. Brown and H. A. Pilsbry : Fauna of the Gatun formation, Isthmus of Panama. II : Acad. Nat. Sci. Philadelphia Proc., vol. 64, 1913, pp. 500-519, pis. 22v26, 5 text figs. The fossils from the upper part of the Culebra formation are listed on page 503 as “species observed in the lignitic layers near Tower N, Las Cascadas.” 9 Idem. The mollusks from the Emperador limestone are listed on page 503 as “spe­ cies observed in the Pecten bed at Tower N, Las Cascadas.” 10 Loc. cit., p. 558.

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has been described by Cooke.11 The Anguilla fauna consists of 40 species. The earliest appearance of genera is the only basis of comparison with the Aquitanian fauna. Both faunas contain the earliest Metis (called Capsa by most Europeans) known in European and American Tertiary deposits. The Aquitanian deposits of Aquitaine contain the earliest European Phacoides (Linga) [included in Phacoides (Here) by Amer­ icans], which also occurs in the Anguilla formation. In America this group is well developed in the middle (Claiborne group), prob­ ably indicating its origin in American waters. Both Metis and Phacoides (Linga) belong to the group of genera now living in West Indian and west African waters, but not in the European seas. The Anguilla fauna bears no resemblance to the Aquitanian fauna of the Piedmont basin.12 It is more similar to the fauna of the Schio beds of the Venetian basin described by Oppenheim,13 probably because the Schio beds include some coralliferous limestones. So far as the mollusks are concerned, the correlation of the deposits of supposed Aquitanian age in the West Indies and Central America with the Aquitanian of Europe rests on their stratigraphie position rather than on any actual similarity. The Aquitanian fauna of Europe abruptly entered the enlarged Ter­ tiary basins and is totally different from the faunas.14 The work of Cossmann and Peyrot shows that more than 20 genera of pelecy- pods appear for the first time in the Aquitanian. Nine of these genera occur also in American Tertiary deposits. The following list shows their earliest appearance in the American Tertiary :

Aquitanian genera. Earliest appearance in American Tertiary. E rv ilia ...... Thomonde and Baitoa formations, Chipola marl (Burdigalian). Metis ( = C apsa)...... Anguilla formation (Aquitanian). Psammobia (Gobraeus) [= P. (Psam- mocola)] ...... Wilcox group (Lower Eocene). (Gurabo formation (Helvetian). Petncola (Rupellaria)...... j Torktown formation (Pontian). Tivela ...... Cercado formation (Burdigalian).

11 C. W. Cooke : Tertiary mollusks from the Leeward Islands and Cuba. Carnegie Inst. Washington Pub. 291, 1919, pp. 105-152, 16 pis. 12 The mollusks of these deposits are listed by F. Sacco : Les étages et les faunes du bassin tertiaire du Piémont. Soc. géol. France Bull., 4th ser., vol. 5, 1905, pp. 893-916, pis. 30-31 (maps). See page 900. 13 P. Oppenheim : Ueber die TIeberkippung von S. Orso, das Tertiär des Tretto und Fauna wie Stellung der Schloschichten. Deutsch, geol. Gesell. Zeitschr., vol. 55, 1903, pp. 98-235, pis. 8-10. 14 G. F. Dollfus : Essai sur l’étage Aquitanien. Service géol. France Bull., vol. 1!), no. 124, 1909, 116 pp., 6 pis.

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Cardium (Cardium)...... Claiborne group (Middle Eocene). Cardium (Cerastoderma)...... Claiborne group (Middle Eocene). Rochefortia ...... Duplin formation (Pontian). Phacoides (Linga) [= P. (Here) in part] ...... Claiborne group (Middle Eocene). The West Indian and Central American deposits that probably are Aquitanian in age have heretofore been referred to the Upper Oligocene. The difference between the faunas of probable Aquitanian age in the West Indies and Central America and of the Oligocene of the same region is not so striking as the difference between the European faunas of the same age. If the American deposits are of the same age as the Aqui­ tanian, there is no reason for continuing to call them Upper Oligocene. Burdigalian.—Deposits in the West Indies and Mexico that seem to be of Burdigalian age contain the earliest large Miocene faunas com­ parable in size to those of the Aquitanian basin. Two groups of faunas, for convenience called Upper and Lower Burdigalian, are recognized. The so-called Lower Burdigalian includes the .Baitoa formation of the Dominican Republic and the Thomonde formation of the Republic of Haiti. The known Baitoa fauna is small, consisting of 60 species, of which a preliminary list has been published.15 The Thomonde forma­ tion has recently yielded more than 350 mollusks. A preliminary list of these fossils has recently been published.16 It is the largest entirely undescribed Miocene fauna from the West Indies. The Baitoa and Thomonde formations, which are of approximately the same age, are the equivalent of the Chipola marl of Florida, but their fauna has a more tropical aspect. Their relation to the Miocene faunas of Aquitaine is not very clear. They are more similar to the Aquitanian and Burdiga­ lian than to the Helvetian and Tortonian. In America the last Alveinus appears in the Baitoa and Thomonde formations, in the Chipola marl, and in the later deposits near Santa Rosa, Mexico. The last European Alveinus (A. girondica (Benoist), called Lutetia by Cossmann and Peyrot),17 occurs in the Aquitaine basin, but is recorded from deposits of Aquitanian, Burdigalian, Helvetian, and Tortonian age. Ervttia is a conspicuous genus in the Baitoa, Thomonde, and Chipola faunas, where it first appears in American deposits. In Aquitaine, however, the earliest species comes from the Aquitanian stage.

15 W. P. Woodring and W. C. Mansfield: A geological reconnaissance of the Domin­ ican Republic. Dominican Rep. Geol. Survey Mem., vol. 1, 1921, pp. 113-114. 16 W. P. Woodring and W. C. Mansfield: Geology of the Republic of Haiti. Republic Haiti Geol. Survey, 1924, pp. 179-190, 194-197. ” See S. D. Harrfs: The genera Lutetia and Alveinus. Paleontographica Americana, vol. 1, no. 2, 1920, 12 pp., pi. 17, 8 tex t figs.

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The Cercado formation of the Dominican Republic and the deposits near Santa Rosa, Isthmus of Tehuantepec, Mexico, are called Upper Burdigalian. The Cercado formation carries about 500 mollusks, many of which have been described by Maury.18 Preliminary lists of unde­ scribed additional species collected by the U. S. Geological Survey ex­ pedition have been published.19 Bose20 described a few species from the beds near Santa Rosa. Mr. Wade, of the Transcontinental Petroleum Company, recently obtained a collection of about 300 species of mollusks from these beds. These mollusks are being described by Mr. Collins, of Johns Hopkins University. The relations of these faunas to the Burdigalian fauna of Aquitaine are not very striking. The Cercado formation contains an undescribed species of the curious Indo-Pacific genus Julia. Julia floridana Dali, from the Chipola marl of Florida (Burdigalian), and a new species from the Bowden formation of Jamaica (Helvetian) are the only other Amer­ ican species. The only other fossil species are J. girondica (Benoist), from the Aquitanian of Aquitaine; J. douvillei Cossmann and Peyrot, from the Helvetian (?) of Aquitaine; and J. lecointrece Dollfus and Dautzenberg, from the Helvetian of Touraine. In Aquitaine the earliest Myrtcea first appears in the Burdigalian, but in America there are Eocene and Oligocene species. The earliest American Tivela comes from the Cercado formation, but in Europe the genus first appears in the Aqui­ tanian. The Burdigalian of Aquitaine carries the earliest Macoma, but in America the earliest species comes from the Tampa formation of Florida (Aquitanian)-. The Santa Rosa fauna is less tropical than the Cercado fauna. It contains Alveinus, like the earlier Baitoa and Tho­ monde faunas. The Santa Rosa beds have furnished the only American Spirulirostra, recently described by Berry.21 The type species of this genus comes from the Helvetian of the Piedmont basin. Other species have been described from the Oligocene of Galicia and the Miocene of northern Germany and Australia. The Santa Rosa fauna contains the exotic genus Euthria, which is common in the Miocene and Pliocene de­ posits of the Mediterranean region and is now living in the Mediterra­

18 C. J. Maury: Santo Domingo type sections and fossils. Bull. Am. , vol. 5, 1917, pp. 165-459, pis. 27-68, 1 text fig. The Cercado mollusks are those de­ scribed from Bluff 2 and Bluff 3, Rio Mao; Zone G, Rio Gurabo; Zones H and I, Rio Cana. 19 W. P. Woodring and W. C. Mansfield: A geological reconnaissance of the Domin­ ican Republic. Dominican Rep. Geol. Survey Mem., vol. 1, 1921, pp. 116-130. 20 E. BSse: Sobre algunas faunas terciarias de México. Inst. geol. México Bol. 22, 1906, pp. 18, 23-27. 21E. W. Berry: An American Spirulirostra. Am. Jour. Sel., 5th ser., vol. 3, 1922, pp. 327-334, 5 text figs.

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nean Sea. Some of the Santa Rosa genera indicate that the fauna is of Helvetian age rather than Burdigalian. Although many genera that occur in the deposits of probable Bur­ digalian age in the West Indies and Mexico are recorded in the Bur­ digalian deposits of Aquitaine, the development of the faunas in the two regions is essentially provincial. The correlation of the American de­ posits with the European rests chiefly on their stratigraphic position. Helvetian.—Widely distributed deposits in the West Indies and Cen­ tral America carry a large fauna remarkably similar to the Helvetian of the Mediterranean region. The Gurabo formation of the Dominican Republic has about 400 mollusks, many of which have been described by Maury.22 Preliminary lists of additional species have been published in the report on .the geology of the Dominican Republic.23 The Bowden formation of Jamaica has a fauna of fully 600 species. Guppy,24 Guppy and Dali,25 Dali,26 and Pilsbry27 have described some of these mollusks. I am now completing a revision of a monograph of the Bowden fauna.

22 C. J. Maury: Santo Domingo type sections and fossils. Bull. Am. Paleontology, vol. 5, 1917, pp. 165-459, pis. 27-68, 1 text fig. The Gurabo mollusks are those de­ scribed from Bluff 1, Rio Mao; Zones A-F, Rio Gurabo; Area patrlcia beds, Rio Cana. Virtually all the mollusks described by Sowerby and Guppy are from the Gurabo formation. G. B. Sowerby: Descriptions of new species of fossil shells found by 3. S. Heniker [sic], Esq. Geol. Soc. London Quart. Jour., vol. 6, 1849, pp. 44-53, 2 pis. R. J. L. Guppy: On the Miocene fossils of Haiti. Geol. Soc. London Quart. Jour., vol. 32, 1876, pp. 516-532, 2 pis. Gabb described many unfigured Miocene species from the Dominican Republic, but as his collection was obtained from almost all the Miocene beds of the country, it has no stratigraphic value. W. M. Gabb: On the topography and geology of Santo Domingo. Am. Philos. Soc. Trans., new series, vol. 15, 1873, pp. 49-260, 2 maps. Pilsbry has recently published a fully illustrated revision of the Gabb collection. H. A. Pilsbry: Revision of W. M. Gabb’s Tertiary Mollusca of Santo Domingo. Acad. Nat. Sci. Phila­ delphia Proc., vol. 73, 1922, pp. 305-435, pis. 16-47, 48 text figs. 23 W. P. Woodring and W. C. Mansfield: A geological reconnaissance of the Domin­ ican Republic. Dominican Rep. Geol. Survey Mem., vol. 1, 1921, pp. 137-151. 24 R. J. L. Guppy: On the Tertiary Mollusca of Jamaica. Geol. Soc. London Quart. Jour., vol. 22, 1866, pp. 281-295, pis. 16-18. On the relations of the Tertiary forma­ tions of the West Indies: Geol. Soc. London Quart. Jour., vol. 22, 1866, pp. 570-590, pi. 26. Notes on West Indian geology, with remarks on the existence of an Atlantis in the early Tertiary period, and descriptions of some new fossils from the Caribbean re­ gion : Geol. Mag., Dec. 1, vol. 4, 1867, pp. 49-501, 6 text figs. On some new Tertiary fossils from Jamaica: Sci. Assoc. Trinidad Proc., vol. 2, no. 2, 1873, pp. 72-88, pis. 1-2. On the West Indian Tertiary fossils: Geol. Mag., Dec. 2, vol. 1, 1874, pp. 404-411, 433- 446, pis. 16-18. Supplement to the paper on West Indian Tertiary fossils: Geol. Mag., Dec. 2, vol. 2, 1875, pp. 41-42. On the Recent and Tertiary species of Leda and Nucula found in the West Indies, with notices of West Indian shells: Sci. Assoc. Trinidad Proc., pt. 12 [vol. 2], 1882, pp. 168-180, pi. 7. 25 R. J. L. Guppy and W. H. D ali: Descriptions of Tertiary fossils from the Antillean region. U. S. Nat. Mus. Proc., vol. 19, 1896, pp. 303-331, pis. 27-30. 28 W. H. Dali: Contributions to the Tertiary fauna of Florida. Wagner Free Inst. Sci. Philadelphia Trans., vol. 3, 1890-1903, 6 pts. Many pelecypods from the Bowden formation are described in parts 4-6. * 27 H. A. Pilsbry : Scapbopoda of the Jamaican Oligocene and Costa Rican Pliocene. Acad. Nat. Sci. Philadelphia Proc., vol. 63, 1911, pp. 165-169, 5 text figs.

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Mollusks from the Gatun formation of the Canal Zone have been de­ scribed by Dali,28 Toula,29 Brown and Pilsbry,30 and Cossmann.31 Very large collections in the United States National Museum have never been examined. Olsson32 has recently described more than 300 species from the Gatun formation of Costa Rica. In the same report he also describes some species from the Canal Zone that have not been discovered in Costa Rica. The total described fauna of the Gatun formation in Panama and Costa Rica consists of about 400 species. According to Olsson,33 the de­ posits in Costa Rica include higher beds than those in the Canal Zone, and some of them may be Upper Miocene. Many collections in the United States National Museum from Costa Rica have not been examined. The resemblance of these West Indian and Central American faunas to the Helvetian of the Piedmont basin is very striking. Perhaps this resemblance would lose some of its force if comparable Aquitanian and Burdigalian faunas were known from the Mediterranean region. The resemblance is particularly close in some families; for ex­ ample, the Ledidae, Malletiidac, Arcidas, Lucinidse, Ranellidse, Septidae ( = Tritonidse), Olividse, Mitridse, Marginellidse, Columbellidae, Cassidse, Terebridse, and Conidse. Leda (Lembulus), two groups of Yoldia, and Neilo are the only groups of Ledidae, Malletidse, and Arcidse in the Pied­ mont Helvetian deposits that are not also known in the West Indian Helvetian beds. The Bowden formation contains all of the seven groups of Area recorded by Sacco in the Helvetian of the Piedmont basin. Of the 12 groups of Lucinidas in the Italian Helvetian, only two—Lucina (Lorpinus) and Loripes—do not occur in the Bowden formation. The following is a list of common Helvetian genera in both regions that are not now living in the Mediterranean Sea: Leda (Jupiteria) Oliva Tindaria Olivella Isognomon (usually called Perna or Ancillaria Pedalion) Latirus

, 28 W. H. Dali : Contributions to the Tertiary fauna of Florida. Wagner Free Inst. Sci. Philadelphia Trans., vol. 3 (6 pts.), 1890-1903. Some peleeypods from the Gatun formation are described in parts 4-6. 29 F. Toula : Ein jungtertiäre Fauna von Gatun am Panama-Kanal. K.-k. geol. Keichs- anstalt Jahrb., vol. 58, 1909, pp. 173-670, pis. 25-28, 15 text figs. ; idem, vol. 61, 1911, pp. 487-530, pis. 30, 81. 30 A. P. Brown and II. A. Pilsbry : Fauna of the Gatun formation, Isthmus of Pan­ ama. Acad. Nat. Sci. Philadelphia Proc., vol. 63, 1911, pp. 336-373, pis. 22-29, 3 text figs. ; idem, vol. 64, 1913, pp. 500-519, pis. 22-26, 5 text figs. 81M. Cossmann : Étude comparative de fossiles miocéniques recueillis a la Martinique et à l’Isthme de Panama. Jour. Conchyl., vol. 61, 1913, pp. 1-64, pis. 1-5. æ A. A. Olsson : The Miocene of northern Costa Rica. Bull. Am. Paleontology, vol. 9, 1922, pp. 179-460, pis. 4-35. “ Idem, p. 20.

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Pllcatula Lyria Phacoides (Cardiolucina) [included Cassis in P. (Cavilucina) by Americans] Sconsia Phacoides (Linga) [included in P. Morum (Oniscidia) (Here) by Americans] Malea Codakia (Codakia) Ficus (called Ficula or Pyrula by Phacoides (Pleurolucina) [called P. most Europeans) (Here) by Sacco] Strombus Metis (called Capsa -by most Euro- Terebra (Fusoterebra) peáns) Conus (several groups) Murex (Pteronotus) E rato Melongena Cancellarla (Trigonostoma) Distorsio (called Persona by most Eu- Cancellarla (Aphera) ropeans) Rissoina (Rissolina) Bursa (called Apollon by most Euro- Rissoina (Zebinella) peans) Rissoina (Zebina) Phos Acteocina The following additional non-Mediterranean genera recorded by Sacco from the Tortonian, but not from the Helvetian, probably occur in Hel­ vetian deposits of other parts of the Mediterranean region. All these genera are common West Indian Miocene fossils: Crassatellites (called Crassatella by Amalthea most Europeans) Cheilea (called Mitrularia by most Protocardia (Nemocardium) Europeans) Xancus (called Turbinella by most Europeans) Barbatia (Obliquarca) and Myrtcea (Myrteopsis) are two extinct groups that occur in the Helvetian deposits of Italy and the West Indies. The Columbellid genus described by Sacco34 as Thiarinella seems to be the extinct genus Metulella, one of the most characteristic West Indian Helvetian genera. Cardiums of the subgenus Cardium occur in the Helvetian deposits of both regions, but are no longer living in the West Indies. Olsson36 has recently described a Halia from the Gatun formation of Costa Eica. This genus occurs in the Helvetian deposits of Italy and is now living only on the west African coast and northward to Spain. The similarity to the Helvetian fauna of Aquitaine is not so striking. Noetia is one of the exotic genera in the Tertiary deposits of Aquitaine, where it is represented by Noetia oleeni (Mayer)36 [called Area (Ana-

35 F. Sacco: I molluschi dei terreni terziarii del PiemOnte e della Liguria, pt. 6, 1890, p. 56, pi. 2, fig. 74. 35 A. A. Olsson: Bull. Am. Paleontology, vol. 9, 1922, pp. 251-252, pi. 7, fig. 7. 36 P. G. Sheldon : Atlantic slope Areas. Paleontographica Americana, vol. 1, no. 1, 1916, p. 69. LVII—B u l l . G e o l . Soc. A m ., V o l . 35, 1923

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dam) by Cossmann and Peyrot]. This species is confined to the Aqui- tanian and Burdigalian of Aquitaine, but also occurs in the Helvetian of Touraine [called Area, (Anadara) by Dollfus and Dautzenberg]. It is the only known European species of the genus. In the American Ter­ tiary the earliest species is in the Gatun formation (Helvetian). Other less spectacular genera now living in both the Mediterranean Sea and the West Indies are represented in the Helvetian deposits of both regions. Despite the resemblance shown by many genera, the West Indian Hel­ vetian faunas are clearly West Indian and the Mediterranean Helvetian faunas have an unmistakable European aspect. The following is a list of common West Indian Helvetian genera unknown in the European deposits: Yoldia (Orthoyoldia) Clementia Cunearca Cyclinella Placunanomia Chione Mytilopsis Tellina (Scissula) Verticordia (Trigonulina) Macoma (Cymatoica) Venericardia (Pleuromeris) Corbula (Bothrocorbula) Echinochama Cancellarla (Cancellarla) Ijucina (Lucina) Solenosteira Phacoides (Callucina) Metula Phacoides (Parvilucina) Strombina Phacoides (Bellucina) Crepitacella Phacoides (Parvilucina) Gaza (Callogaza) Cardium (Trigoniocardia) A large number of typical European genera that are common fossils in the Mediterranean Helvetian deposits have never been discovered in American Tertiary deposits. These genera are particularly abundant in the families Cardiidae, Veneridse, Tellinidae, Rissoidse, and Trochidae. Tortonian.—The Cerros de Sal formation of the Dominican Republic is the only West Indian formation of supposed Tortonian age having a relatively large molluscan fauna. A preliminary list of 75 species has been published.37 It is an almost pure West Indian fauna, with the ex­ ception of the two Pacific genera, Strombina and Solenosteira. The Cerros de Sal fauna affords no adequate basis of comparison with the Tortonian fauna of the Piedmont basin. This European Tortonian fauna resembles the preceding Helvetian fauna in many features, and therefore is like the West Indian Helvetian faunas; but the similarity between the Mediterranean and West Indian faunas reaches its climax

31 W. P. Woodring and W. C. Mansfield : A geological reconnaissance of the Dominican Republic. Dominican Rep. Geol. Survey Mem., vol. 1, 1921, pp. 163-164.

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in the Helvetian and thereafter declines, except for a slight increase in the Astian, as is graphically shown in figure 4. In this graph, as in figure 3, the Tortonian fauna of Aquitaine fails to agree with the other faunas. PLIOCENE The Pliocene faunas from the West Indies and Central America are very meager. Gabb38 described a so-called Pliocene fauna from Port

F ig u r e 4.— Graph showing Percentage of West Indian Miocene Genera in Miocene and Pliocene Faunas of Piedmont and Aquitaine Basins Solid line represents Piedmont basin; broken line represents Aquitaine basin.

Limon, Costa Bica. It has been known for some time that this is a mixed fauna, as deposits of both Miocene and Pliocene age crop out at Port

38 W. M. Gabb : Descriptions of new species of fossils from the Pliocene clay beds be­ tween Limon and Moen, Costa Rica, together with notes on previously known species from there and elsewhere in the Caribbean area. Acad. Nat. Sci. Philadelphia Jour., series'2, yol. 8, 1881, pp. 349-380, pis. 45-47.

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Limon.39 Olsson40 has recently described these deposits of Miocene and Pliocene age. The Toro limestone of the Panama Canal Zone may be Pliocene, but its fauna is negligible. Deposits of Pliocene age in the Eepublic of Haiti have recently been described, but their known mol- luscan fauna consists of only 17 species. I t is a pure West Indian fauna. If a large Pliocene fauna were discovered in the West Indies, it prob­ ably would contain a small percentage of exotic and extinct genera. The Pliocene Caloosahatchee formation of Florida carries a large subtropical fauna, including the Pacific genus Solenosteira and the extinct genera Corbula (Bothrocorbula) and Perplicaria. There are no Pliocene faunas in the West Indies and Central America comparable to the Pliocene faunas of the Piedmont basin. The resem­ blance of the Italian Helvetian and Tortonian faunas to the Helvetian faunas of the West Indies and Central America continues as a prominent but diminished feature of the Plaisancian and Astian faunas (see figure 4). This resemblance continues even in the early Quaternary Sicilian stage. Some of the living West Indian species are known by names given by Seguenza to fossils from Sicily. Refined work is gradually eliminat­ ing these names from the living West Indian fauna, but the confusion of fossil and living species in the two regions gives an indication of the similarity of some elements of the faunas.

*> T. W. Vaughan : TJ. S. Nat. Mus. Bull. 103, 1919, p. 221. 40 A. A. Olsson: Bull. Am. Paleontology, vol. 9, 1922, pp. 183-184.

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