FIELD MUSEUM OF NATURAL HISTORY JUL 2196 -BULLETINS ; OF AMERICAN PALEONTOLOGY (Founded 1895) Vol. 54 No. 243 MORE ON VARIATION IN THE GENUS LEPIDOCYCLINA (LARGER FORAMINIFERA) By W. Storrs COLE 1968 Paleontological Research Institution Ithaca, New York, 14850, U.S.A. PALEONTOLOGICAL RESEARCH INSTITUTION 1967-1968 PRESIDENT Shae NOM Oe Rete aI TUN, fe eam ay wrath (Ny Voie! KENNETH E. CASTER WVIGHSPRESIDENT Vo. fy sue tore eee oi Se oh POOP oir, WILLIAM B. HEROY SEGRE TARY «1 RRASURER? o25-discctes eset RY IT ye ha REBECCA S. HARRIS DIRECTOR) ene IS EDA ENE. Me uettao KATHERINE V. W. PALMER GOUINSEE 2). SM Pe Be DS GH. Sa PTS MUR LE a Ae OU fed ARMAND L. ADAMS REPRESENTATIVE “AAAS GOUNGIL, 28) neh Be hideccescevshett oocowe KENNETH E. CASTER Trustees KENNETH E. CAsTER (1966-1972) KATHERINE V, W. PALMER (Life) DoNALD W. FISHER (1961-1967) WILLIAM B. HERoy (1963-1968) REBECCA S. Harris (Life) AxeEL A. OLSSON (Life) DANIEL B. Sass (1965-1971) Hans G. KuGLer (1963-1969) W. Storrs CoLe (1964-1970) BULLETINS OF AMERICAN PALEONTOLOGY and PALAEONTOGRAPHICA AMERICANA KATHERINE V. W. PALMER, Editor Mrs. Fay Brices, Secretary Advisory Board KENNETH E. CASTER HANS KUGLER A. Myra KEEN JAY GLENN Marks AXEL A. OLSSON Complete titles and price list of separate available numbers may be had on application. For reprint, Vols. 1-23, Bulletins of American Paleontology see Kraus Reprint Corp., 16 East 46th St., New York, N. Y., 10017 U.S.A. For reprint, vol. I, Palaeontographica Americana see Johnson Reprint Corpora- tion, 111 Fifth Ave., New York, N. Y. 10003 U.S.A. Subscription may be entered at any time by volume or year, with average price of $16.00 per volume for Bulletins. Numbers of Palaeontographica Americana in- voiced per issue. Purchases in U.S.A. for professional purposes are deductible from income tax, For sale by Paleontological Research Institution 109 Dearborn Place Ithaca, New York 14850 U.S.A. BULLETINS OF AMERICAN PALEONTOLOGY (Founded 1895) Vol. 54 No. 243 MORE ON VARIATION IN THE GENUS LEPIDOCYCLINA (LARGER FORAMINIFERA) By W. STORRS COLE June 21, 1968 Paleontological Research Institution Ithaca, New York 14850, U.S.A. Library of Congress Catalog Card Number: GS 68-136 Printed in the United States of America CONTENTS Page PAUDS UA Gtr semen Orem 08 Uy AOC IN a OVI ed Bed Ses Peart Pexo noe ate Pee ee eS 295 TritKO CU CHIO 1g meer ee ree ate NP Pe ee et een oe ta Sederoassti cae a teeas sh ama Ree, 295 Localities of figured specimens ........... 296 Examples of variation in species of Lepidocycl ind .......ccccccccescessseesceseseseseessessesecees 297 Shape and size of the embryonic chambers of Lepidocyclina ........00.00.000.... 297 Sulbcemenam ote e2c Oayl7 70 ctis saute. eeee selene seeseetee aes cie aa ssaudeeece st -Pandteaenescee 299 Multilocularsemibtyonic: Chambers issccrc:..052---<2-cersceee-aeecevie vances cesecseeeeeeteeaces-aee2- 301 History of specific names which have been applied to L. van ghani ............000005. 303 Lepidocyclina tournoueri Vaughan not Lemoine and R. Douvillé, 1904, ASV MONA Ola nels ae USI <..52hfacde cc eeetee cece sage ene seessunoesbteaueencoseati< 306 Type localities of certain American larger Foraminifera ..............c:cccceeceeeee < 307 Stratioraphicsimp lations isieac..f tcaccee se aes ee Ues can aiecauetiee cas ox bade esse owes seet bac B15 IREREREMCESCIE CC ete set cree ee PD Neda cee Wee Senne wea Wewean doe weed sacee Se atunrenseeeuelits 314 At eS arte see ae eenatt etre tepiese oot aas ce trre me ee cea-cuaseee rset cae nec cto noes vast eDeeaaa sven ee savent cesta seas 319 Mia Gl Koes eee cts AOR ce Ee See Ets eg reac ta cdso ira Wage roas tae sac ctactatae stds Cnbaccetocbaareceetes 326 MORE ON VARIATION IN THE GENUS LEPIDOCYCLINA (LARGER FORAMINIFERA) W. STorRS COLE Cornell University, Ithaca, New York ABSTRACT Although this article is primarily a study of the variation which may occur in Lepidocyclina vaughani Cushman (191924), additional data are presented to demon- strate the development of multilocular embryonic chambers in Lepidocyclina, and the subgeneric classification of this genus is revised. Lepidocyclina is subdivided into the subgenera Polylepidina and Lepidocyclina s. s. as Eulepidina is considered to be a synonym of Lepidocyclina s. s. Synonyms of L. vaughai are: Lepidocyclina tournouert Vaughan, not Lemoine and R. Douvillé, L. dartons Vaughan, L. tempani Vaughan and Cole, and L. crassimargo Vaughan. The type localities of certain Oligocene species of larger Foraminifera are reviewed and the associations of species at these localities are analyzed to demonstrate that these faunal associations are widespread in the Caribbean region. INTRODUCTION When Vaughan and Cole started their collaboration in the study of larger Foraminifera in 1931, generic definitions and specific descriptions were often based on insufficient data. New names, both generic and specific, were being erected at an astounding rate, in part because older names were not understood, and in part because variation was neither appreciated or understood. Vaughan (19334, p. 6) was the first American to appreciate the variation which may occur in species of Lepidocyclina. He wrote ‘The amount of variation in many species of orbitoids is bewildering.” How- ever, Vaughan (19334, p. 7) had also the philosophy expressed in his own words “... it 1s not always possible to study the variation of a species, but sometimes new species are, in my opinion, justifiably based on a single specimen or on a few specimens.” Vaughan (19334, p. 7) stated “The evaluation of the characters presented by meager material 1s dependent on the judgment, which ts a function of the experience of the investigator.’’ Galloway (1933, p. 469) had the same concept when he included in his definition of species the statement “A group of individuals having substantially the same struc- ture... which are identified by a specialist in that group under the same binomial name’ (italics mine). These concepts are valid, but clearly point to the fact that there must be a constant review and revision of generic and specific classification as data accumulate. This discourse is a continuation of earlier ones (Cole, 19576; 19584; 1960; 1961a,b; 1962; 1963b,c) in which the subgeneric classification and selected species of the genus Lepdocyclina were analyzed. There has been some resistance to studies of this kind. Hanzawa (1962, p. 130) wrote among other comments “ . . . Cole (1960, p. 101) 296 BULLETIN 243 postulated that Nephrolepidina is synonymous with Ewlepidina, despite their having been distinguished as two subgenera of the genus Lepidocyclina by various authors for half a century.’ Eames and Clarke (1965) retained the subgeneric name Pliolepidina H. Douvillé, 1917, for specimens of Lepidocyclina with multilocular embryonic chambers although Cole (1960; 1962; 1963b,c) and Sachs (1964) had presented conclusive evidence that multilocular embryonic chambers result from biologic ‘‘accidents’’ either in reproduction or developmental stages of individuals. However, on the other side Grimsdale (1959, p. 15) wrote ‘“The reality of this tendency among paleontologists—to over-estimate the taxono- mic significance of variable characteristics and so to erect superfluous species—is clear from various studies by Vaughan (1933a), Vaughan and Cole (1941), and Cole (1944, 1952, 1953, 19575) wherein some 70 described species have beeen reduced by synonymization to about 15 accepted.” Since my initial study (Cole, 1952, p. 29) of Lepidocyclina vaughani Cushman from the Panama Canal Zone, over 100 additional thin sections of this species have been prepared. Additional thin sections were made of supposedly valid species which had internal structures similar to those of Lepidocyclina vaughani. As a consequence of this study several sup- posedly valid species are assigned to the synonymy of L. vawghani, and an additional revision of the subgeneric classification of Lepidocyclina is suggested. The specimens on which this article is based will be transferred eventually from the Cole collection at Cornell University to the U.S. National Museum. The localities from which the specimens illustrated were obtained follow. LOCALITIES OF THE FIGURED SPECIMENS Antigua Loc. 1—Half Moon Bay (References: Vaughan, 1923, PP, 2552 1924 ph. 33, Hie. 9; 19334, pi 32). 2—Hodge Hill (Reference: Vaughan, 19334, p, 27). Cuba 3—USGS loc, 7552, just south of El Jique, about 6 miles above the mouth of the Yateras River, altitude about 250 feet (References: Vaughan, 19330, p. 354; 1933a, p. 22). LEPIDOCYCLINA: COLE 297 Florida 4—Exposure of white to cream-colored, soft limestone in a large sinkhole near Duncan Church, 7 miles southwest of Chipley (References: (Cole; 19347 ps 21; Veron, 1942.%p; 58; Cooke, 1945, p. 104). France 5—Le Sauset, Bouches du Rhone (Reference: Cole, 19614, p. 138). Mexico 6—Arbol Grande, near Tampico, Tamaulipas, Vaughan’s loc. M 12 V (Reference: Vaughan, 1924, pl. 33, figs. 6, 7; 19334, p. 26; Cole, 1961a, p. 377). 7—Between kilometer posts 17-18 on the Aguila Petroleum Com- pany’s narrow-gauge railroad between Potrero and Tanhuijo, Sate of Vera Cruz (sta. SiC; M-S 1; W. S:-Cole, collector (References: “Cole «and: Gillespie, 19305 Cole, 19614,"p. 377; 19635, p. 12). Panama 8—Quarry of Panama Cement Company, 200 feet north of the Transisthmian Highway and 0.9 mile in direct line northwest of highway bridge across Rio Gatuncillo; collectors, J. R. Shultz and W. P. Woodring, 1947 (References: Cole, 1952, p. 7; Woodring, 1957, p. 118). 9—Transisthmian Highway, 2 miles in direct line north-north-west of Rio Chagres bridge; collector W. P. Woodring, 1947 (References: Cole, 1952, p. 7; Woodring, 1957, p. 118). 10—Peninsular north of Barbacoas Island, in field 0.8 mile northeast of Lighthouse 13; collectors, S$. M. Jones and W. P. Woodring, 1947 (References: ‘Coles 1952, ps6; Woodring; 1957p; 116). Trinidad 11—Taparo Point (Reference: Vaughan and Cole, 1941, p. 17). 12—Pefial-Rock Road at mile 14, block from a mudflow (Kugler sta. K409a) (Reference: Vaughan and Cole, 1941, p. 14). EXAMPLES OF VARIATION IN SPECIES OF LEPIDOCYCLINA SHAPE AND SIZE OF THE EMBRYONIC CHAMBERS IN LEPIDOCYCLINA The genus Lepidocyclina has been subdivided into a number of sub- genera largely on the shape and arrangement of the embryonic chambers. However, the validity and necessity for such subdivision has been ques- tioned, or the issue has been evaded. 298 BULLETIN 243 As early as 1926 Vaughan (p. 394) wrote concerning the American Oligocene species Lepidocyclina undosa Cushman “The embryonic cham- bers are. . . of the Nephrolepidine type. They grade into the Eulepidine type of chambers...” Gravell (1933, p. 31) observed concerning Venezuelan specimens assigned to this species “A number of sections of this species show the embryonic apparatus varying from the nephrolepidine to the eulepidine type, making the assignment of this species to either subgenus virtually optional.” That the embryonic chambers of this species do vary from nephrolepidine to eulepidine has been substantiated recently by Sachs (1964, pl. 1, figs. 3, 10). Caudri (1934, text fig. 21a-m) illustrated the embryonic chambers of a number of specimens of L, martini Schlumberger, an Indo-Pacific Miocene species, which vary from nephrolepidine to eulepidine and trybliolepidine. Caudri (1934, p. 119) wrote “ .. . all the smaller nephrolepidine or trybliolepidine Lepidocyclinae from the Dutch East Indies belong to one group...” Drooger and Socin (1959, p. 419) wrote concerning the embryonic chambers of the European Miocene species L. sournoueri Lemoine and R. Douvillé “It is of various nephrolepidine to ‘trybliolepidine’ types.” This observation is confirmed by their excellent illustrations (pl. 1, fig. 1; pl. 2, all figs.). Cole (1945, pl. 25, figs. A-C; Cole et al, 1960, pl. 12, figs. 2,3) figured specimens of L. martini and of L. japonica Yabe (Cole, 1963a, pl. 10, figs. 9, 13, 14, 18), other Indo-Pacific Miocene species, with nephro- lepidine embryonic chambers in some specimens and eulepidine in others. The variation in shape of the embryonic chambers in these two species with relatively small embryonic chambers is duplicated in specimens of L. radiata (Martin), an Indo-Pacific Miocene species, with relatively large embryonic chambers (Cole, 1960, pl. 1; 1963, pl. 42). The embryonic chambers of the American Eocene species L. chaperi Lemoine and R. Douvillé vary from nephrolepidine to eulepidine (Cole, 19634, pl. 8, fig. 3; pl. 9, figs. 1-3). Moreover, specimens of this species have been illustrated (Cole, 1952, pl. 10, figs. 1, 5, 7) which have Lepidocyclina s. s. embryonic chambers. The American Oligocene species L. van ghani Cushman (PI, 20, figs. 1-3, 5-7) has embryonic chambers which at one end of the series are similar to the lepidocycline 5. s. kind and at the other end to large eulepidine or trybliolepidine shape chambers. LEPIDOCYCLINA: COLE 299 The American species L. ywrnagunensis Cushman (PI. 24, figs. 5, 8) has been assigned to the subgenera Nephrolepidina (Vaughan, 1924, p. 798) and Lepidocyclina s. 5. (Vaughan, 1926, p. 392; 1933a, p. 21; Cole, 1934 ps 24591952:.p,. 22)... Cole, (1960, ip. 1363 1961b; p. 143), at the time the suggestion was made that the subgeneric name Nephrolepidina was superfluous, assigned L. yuwrnagunensis to the subgenus Eulepidina. The embryonic chambers (PI. 24, fig. 8) of certain specimens of L. yurnagunensis are similar to those of specimens placed in the subgenus Lepidocyclina s. 5., but other specimens (Pl. 24, fig. 5; Cole, 1952, pl. 20, fig. 11; Sachs and Gordon, 1962, pl. 2, figs. 3, 5, 8) are definitely of the nephrolepidine (= eulepidine) kind. Specimens (PI. 24, figs. 4, 6, 9) from the Oligocene of Trinidad identified by Vaughan and Cole (1941, p. 75) as L. tempani Vaughan and Cole (= L. vaughani Cushman) have embryonic chambers which are not only variable in size but in shape. Specimens from Guam identified as L. parva Oppenoorth [= L. sumatrensis (Brady) } (Cole 7 Cole and Bridge, 1953, pl. 9, figs. 10, 11, 18; pl. 10, figs. 11, 12) and from the Eniwetok drill holes (Cole, 1957c, pl. 242, figs. 3-20) have embryonic apparatuses which are lepidocycline s. 5. to nearly typical eulepidine shape. Several specimens of L. vaughani (Pl. 20, figs. 1-3, 6, 7) from one locality are illustrated to demonstrate that relatively large differences in size occur. These same size differences are shown by specimens (PI. 24, figs. 4, 6, 9) from another locality. However, specimens of L. wndosa (PI. 22, figs. 1, 6; Cole, 1934, pl. 4, figs. 4, 5) from one geographic locality may have embryonic apparatuses which are small compared with those of specimens of the same species from another geographic locality (PI. 22, fig. 7; Cole, 19634, pl. 8, figs. 1, 2). Another species of which sufficient preparations have been made to show the difference in size of the embryonic apparatus is L. pwstulosa (Cole, 1963, pls. 23-25). Other illustrations could be presented to demonstrate that the shape and size of the embryonic apparatuses in species of Lepdocyclina are ex- tremely variable. Any subdivision of this genus into subgenera must recognize this fact. SUBGENERA OF LEPIDOCYCLINA In the ‘“Treatise on Invertebrate Paleontology,’ Part C, Protista 2 (1964, p. 721) Cole recognized four subgenera of Lepidocyclina, namely, Lepidocyclina s. s., Eulepidina, Pliolepidina and Polylepidina. In 1963 300 BULLETIN 243 during the time this volume of the Treatise was in press Cole restudied Pliolepidina and published (19634, p. 18) a revised opinion on the status of this subgenus. The conclusion reached was that Pliolepidina H. Douvillé, 1917, was a synonym of Lepidocyclina s. 5. Gumbel, 1870. Un- fortunately, this concept could not be incorporated in the Treatise. Cole (19634, p. 35) wrote “If Pliolepidina is accepted as a synonym of Lepidocyclina 5. s., there will be three subgenera, Polylepidina, Lepido- cyclina s. s., and Eulepidina in the genus Lepidocyclina.” The specimens on which this reduction in the number of subgenera was based and the interpretations which were made have been published (Cole, 19630, and references cited) and will not be repeated. The major structural difference in the supposedly valid subgenera 1s between Polylepidina and the other two subgenera. Polylepidina has a single, but pronounced, coil of periembryonic chambers which partially surround the embryonic chambers (Cole, 19634, pl. 1, fig. 3). In Lepido- cyclina s. 5. and Eulepidina two principal periembryonic chambers are developed, one on each side of the dividing wall between the embryonic chambers (for Lepidocyclina, see: Cole, 19630, pl. 1, figs. 1, 2, 4, 5; for Eulepidina, see: Cole, 19630, pl. 8, fig. 3; pl. 9, fig. 3; Pl. 21, fig. 7; PL 22, fig. 4). This structural arrangement is sufficient to distinguish Poly- lepidina from the other two subgenera. If Lepidocyclina s. s. and Eulepidina are to be recognized as valid subgenera, some criterion other than the development of the periembryonic chambers must be found. Thus, the shape and arrangement of the embryonic chambers have been used. Lepidocyclina s. s. was defined on the occurrence of two equal, or subequal embryonic chambers, whereas Eulepidina had an initial chamber which was either partially embraced or completely enclosed by the second chamber. A specimen (Pl. 20, fig. 1) of L. vawghani has embryonic chambers of the kind recognized as Lepidocyclina s. 5. This illustration should be compared with the one of L. pwstulosa illustrated as figure 2, plate 3 (Cole, 19636). This same relationship is demonstrated if the illustration (PI. 20, fig. 1) of L. vawghani is compared to certain equatorial sections of L. chaperi (Cole, 1952, pl. 11, figs. 4, 6). Many other species, if sufficient thin sections are available, show the same variation in shape of the embry- onic chambers, as do L. vaughani and L, chaperi, for example, L. suma- trensis (see: Cole, 1957c, pl. 242, figs. 3-20). In some species of Lepidocyclina the embryonic chambers run the LEPIDOCYCLINA: COLE 301 gamut from lepidocycline s. s. to eulepidine of which L. vaughani is an example. Other species have embryonic chambers which vary from lepidocycline s. 5. to nephrolepidine (for example, L. svmatrensis). Still other species have embryonic chambers which are more or less consistently of one kind (L. mantelli and L, canelle7). From the data available I am forced to the conclusion that there is no possibility of separating species into the two subgenera, Lepidocyclina s. S. and Exlepidina, and, therefore the proposal is made that Ewlepidina is a junior synonym of Lepidocyclina s. 5. Thus, the genus Lepidocyclina would be subdivided into Lepidocyclina (Polylepidina) and Lepidocyclina (Lepidocyclina) . The formerly used sugbeneric names are meaningful as descriptive terms similar to the manner in which gonuatitic, ceratitic, and ammonitic are used at present. Kummel (1961, p. 289) wrote concerning these am- monoid names ““. . . the terms have no taxonomic or stratigraphic signif- cance, but they are excellent descriptions of the basic suture patterns.” The fact has not been overlooked that the American middle Eocene species Lepidocyclina ariana Cole and Ponton has subequal embryonic chambers and that the first American species in which certain specimens have the initial chamber enclosed by the second chamber is the upper Eocene L. chaperz Lemoine and R, Douvillé. From these data alone it would appear that two subgenera could be recognized as there would appear to be evolution with time from lepido- cycline s. s. embryonic chambers to eulepidine. However, once the variability of the shape of the embryonic chambers from lepidocycline to eulepidine in several species has been demonstrated, the actual shape of the embryonic chambers loses its significance as an evolutionary structure and must be relegated to specific importance only. The basic structural change in the embryonic apparatus is the develop- ment of two principal periembryonic chambers. Polylepidina with one major periembryonic chamber has an entirely different arrangement of the periembryonic chambers than the other lepidocyclines have. This is the only evolutionary advance. The change in shape of the embryonic chambers is not constant varying from individual to individual in many species and can not be defined. MULTILOCULAR EMBRYONIC CHAMBERS The occurrence of two distinct sets of normal, bilocular embryonic chambers in specimens of Lepidocyclina has been established (Zuffardi 302 BULLETIN 243 Comerci, 1929, pl. 9, fig. 22; Rutten and Vermunt, 1932, pl. 3, fie. 25 Cole, 19634, pl. 4, fig. 6; this article Pl. 21, fig. 6). Specimens have been illustrated which have a single set of trilocular embryonic chambers (Pro- vale, 1909, pl. 3, fig. 18; Cole, 1962, pl. 4, figs. 4, 5; 1963, pl. 14, fig. 6; this article PI, 21, fig. 5) as well as specimens which have “twinned” trilocular embryonic chambers (Cole, 1962, pl. 4, fig. 1s 19630, pl. 44, fe1)i, Specimens in which two sets of embryonic chambers occur must have had two nuclei at the time the embryonic chambers developed although the ephebic development in these specimens is similar to that of mononucle- ate specimens with a single set of bilocular embryonic chambers, Certain specimens (Cole, 1962, pl. 5, fig. 2; 1963c, pl. 45, fig. 1) which obviously developed under the control of a single nucleus have deformed, bilocular embryonic chambers. Sachs, (1964) in a carefully reasoned and well-illustrated study of Lepidocyclina undosa with multi- locular embryonic chambers, demonstrated that specimens “. . . with mono- nucleate embryonts in which the initial chamber 1s so large with respect to the total size of the embryont that it has resulted in the protoplasm of the second chamber being concentrated into a series of small, interconnected lobes resembling chamberlets about the equatorial plane of the initial chamber.” The illustrations which have been cited demonstrate that a mono- nucleate specimen may develop multilocular embryonic chambers. How- ever, this kind of multilocular embryonic chamber can develop under the control of two or more nuclei. The specimen (PI. 21, fig. 6) has two distinct and separate sets of bilocular embryonic chambers which are not fused. Another specimen (PI. 21, fig. 4) from this same sample has two distinct sets of embryonic chambers in which fusion has occurred so that there is interconnection between the second embryonic chambers. A third specimen (PI. 21, fig. 5) has trilocular embryonic chambers, This speci- men has large periembryonic chambers on the periphery of the large chambers at each end and smaller periembryonic chambers at each side of the dividing walls of the central chamber. In addition each of these divid- ing walls has a stolon near the center. In the specimen (PI. 21, fig. 6) the two nuclei either were situated within a single mass of protoplasm so that two sets of bilocular embryonic chambers could develop, or two masses of protoplasm, each with a nucleus, were in close proximity to each other. However, there was sufficient LEPIDOCYCLINA: COLE 303 separation of the nuclei so that two sets of normal bilocular embryonic chambers developed. In the specimen (Pl. 21, fig. 4) the spacing of the nucle was such that the development of the walls of the second embryonic chambers at the zone of juxtaposition was inhibited, thus the two sets of embryonic chambers were interconnected producing in effect a trilocular embryonic apparatus. The specimen (PI, 21, fig. 5) seemingly developed from two closely associated nuclei in which the central chamber of this set represents the combined initial chamber developed by the two nuclei. The two large chambers on either end of the trilocular set represent the second embryonic chambers, each of which developed under the control of a separate nucleus. Proof has beeen given that specimens may develop multilocular embry- onic chambers either under the control of one or two nuclei. Specimens in which three sets of embryonic chambers are found must develop under the control of three nuclei. The specimen (Cole, 1962, pl. 7, fig. 5) has two sets of bilocular embryonic chambers in associaticn with a large multilocular chamber. A minimum of three nuclei are required to form this association. Sufficient proof has been presented by Cole (1960; 1962, p. 33; 1963, p. 14; 1963c) and Sachs (1964) to show conclusively that multiloc- ular embryonic chambers are only one possible variant in the develop- ment of embryonic apparatuses, and, thus, have no value on a subgeneric or higher taxonomic level. Eames and Clarke (1965), however, argued that Pliolepidina H. Douvillé, 1917, is a recognizable genus “. . . possessing a characteristic multilocular nucleus [and] is generically distinct from Lepidocyclina (Lepidocyclina) Gimbel, 1870..." Moreover, they reject the concept of Vaughan and Cole (1941, p. 65; Cole, 19636) that L. trinitatis is a synonym of L. pustulosa and that “Pliolepidina’ tobleri represents speci- mens of L. pustulosa which developed multilocular embryonic chambers. The evdence available refutes the postulates made by Eames and Clarke. HISTORY OF THE SPECIFIC NAMES WHICH HAVE BEEN APPLIED TO L. VAUGHANI Six specific names have been used or applied to American specimens, all of which are referred to Lepidocyclina (Lepidocyclina) vaughani Cush- man (1919). These names are: L. tournoueri of H. Douvillé, Vaughan, and Cole, not L. tournoueri Lemoine and R. Douvillé (1904) (a European 304 BULLETIN 243 species), L. verbeeki of Barker, nor L. verbeeki Newton and Holland (1899) (an Indo-Pacific species), L. tem pani Vaughan and Cole (1933), L. dartoni Vaughan (1933), L. crassimargo Vaughan (1933), and L. lehneri van de Geyn and van der Vlerk (1935). H. Douvillé (1924, p. 47) identified specimens from Erin Point, Trinidad, as Nephrolepidina tournoueri Lemoine and R. Douvillé (1904, p. 19), a European species. Vaughan (1924, p. 798) assigned specimens from Arbol Grande, near Tampico, Mexico, to this species stating it occurred in the Oligocene of Mexico and Trinidad. In 1933 Vaughan (a, p. 25) gave a complete description of the Mexican specimens, but he did not mention the specimens from Trinidad. However, Vaughan and Cole (1941, p. 75) definitely assigned the specimens from Trinidad which H. Douvillé (1924, p. 47) had referred to Nephrolepidina tournoueri to the synonomy of L. fempanii Vaughan and Cole (47 Vaughan, 1933a, p. 26). Vaughan and Cole (1941, p. 75) stated “Wan de Geyn and Van der Vlerk (1935, p. 271, figs. 14, 15) have named a species Lepidocyclina (Nephrolepidina) lehneri. Both the description and the figures are inadequate, but it is possible that this species is a synonym of L. tempanit.” Thus, two species were recognized by 1941, if L. lehneri is accepted as a synonym of L. fempanii, L. tournoueri based on specimens from a single Mexican locality, and L. tempanii reported from Antigua and Trini- dad. Vaughan and Cole (#7 Vaughan, 19334, p. 27) wrote “The main difference in the two species is readily seen in a comparison of the equatorial section. L. tempanii has much more elongate equatorial cham- bers than L. fowrnoueri.” Cole (1952, p. 29) demonstrated that the elongation of the equatorial chambers in Mexican specimens which Vaughan (1924, p. 798; 1933a, p. 25) had referred to L. tournoueri was equal to that of specimens of L. fempant. Equatorial chambers of Mexican specimens and of topotypes of L. tempani are illustrated on Plate 21, figures 1-3. As this difference in the degreee of elongation of the equatorial chambers was the only criterion by which these species could be separated, Cole (1952, p. 29) concluded that the Mexican specimens represented the same species as those from Antigua and Trinidad which had been named L. tempanii. In later studies Cole (19614, p. 388, 391; 1964, p. 141) used the specific name L. tournouer? Lemoine and R, Douvillé (synonym: L. tempanii) for speci- mens of this kind, LEPIDOCYCLINA: COLE 305 Species associated with the Mexican specimens of L. tournoueri re- ported by Vaughan (19332) are a ‘‘dwarf variety of L. caneller” (p. 15) and Lepidocyclina parvula Cushman var. crassicostata Vaughan and Cole (p- 17) (= microspheric form of L. canelles—see: Cole, 1961, p. 384). In addition specimens of Camerina panamensis (Cushman) {reported as Camerina dia (Cole and Ponton) } occur at this Mexican locality (Cole, 19614, pl. 29, figs. 6,9,10). Specimens from San Pedro, Peru, originally identified by Barker (1932, p. 278, 279, pl. 16, figs. 2, 3, 5) as Lepidocyclina (Nephrolepr- dina) verbeeki Newton and Holland, an Indo-Pacific species, were con- sidered by Cole (1952, p. 30) to be typical L. vanghani. These specimens were associated with Mvogypsina panamensis (Cushman). In 1933 (4, p. 36) Vaughan described a stellate species of Lepido- cyclina as L. (Nephrolepidina) dartoni from Cuba. Cole (1952, p. 27) identified specimens from a single locality in the Panama Canal Zone as L. dartoni. In 1961 Cole (4, p. 389) wrote “It should be recognized that in Lepidocyclina the stellate pattern is produced only in certain individuals, probably under the influence of ecological conditions, and that this pattern is not genetically produced. Therefore, it does not have value as a specific character.” The specimens of L. dartoni (Cole, 1952, pl. 19, figs. 1-8), except for their stellate pattern, are identical with other nonstellate specimens (Cole, 1952, pl. 19, figs. 8-12) which were assigned to L. tournoueri. Therefore, L. dartoni was placed in the synonomy of L. towrnoueri (Cole, 1961a, p. 388). Additional specimens of L. dartoni from Panama (Pl. 22, figs. 2, 3, 5) are illustrated for comparison. Although by 1961 Cole had suggested that specimens referred to L. tournouert, L. tempani, L. dartoni and L. lehneri represented only one species which he assigned to L. tournoueri, he considered that L. vawghani was a distinct and readily recognizable species. However, as more exten- sive faunal studies were completed in the Caribbean area, it became ap- parent that L. vaughani (reported from Antigua, Costa Rica, Carriacou, and Panama) never was found in association with L. tournoueri (Mexico, Antigua, Trinidad, and Panama) except for a doubtful occurrence of L. tournouert with L, vaughani at a single locality on Carriacou (Cole, 19582, pl. 28, fig. 8). 306 BULLETIN 243 This seemed surprising inasmuch as these two species of Lepidocyclina by associated species of other larger Foraminifera and general stratigraphic position should occur together. Thus, a question was raised whether there were in actuality two species, L. tournoueri (= L. tempanit) and L. vang- hani, or whether these specimens represented only one species. An attempt will be made to demonstrate that all of these specimens should be com- bined under one specific name, L. va ghani. LEPIDOCYCLINA TOURNOUERI VAUGHAN, NOT LEMOINE AND R. DOUVILLE, 1904, A SYNONYM OF L. VAUGHANI CUSHMAN. Specimens assigned to Lepidocyclina vanghani from Panama (Pl=19; fig. 3) and from Antigua (PI, 23, fig. 9) have a small, distinct umbo bordered by a wide flange which expands noticeably at the periphery. The lateral chambers are open, large, with slightly curved or straight roofs and floors, arranged in regular tiers with thin, but distinct pillars. The equa- torial chambers (PI. 19, figs. 1, 6; PI. 20, fig. 5; Vaughan, 19334, pl. 16, figs. 3, 4; Cole, 1952, pl. 21, fig. 5) are markedly rhomboid. Specimens assigned to “L. fournouer?’ (PI. 23, fig. 2; Vaughan, 19334, pl. 13, fig. 2) and “L. tempani’”’ (Pl. 23, figs. 1, 5, 11; Vaughan 1933a, pl. 13, fig. 3; Vaughan and Cole, 1941, pl. 39, fig. 6) are not only similar, but also resemble the umbonate part of specimens of L. vawghani inasmuch as the lateral chambers and pillars have the same shape and arrangement. However, in equatorial section specimens identified as “L. tournouer?” (PI. 21, figs. 2, 3; Vaughan, 19334, pl. 13, fig. 2) and “L. tempanu’ (pl. 1, fig. 5; pl. 3, fig. 1; Vaughan, 1933, pl. 13, figs. 4-6; Vaughan and Cole, 1941, pl. 39, figs. 5, 8) have elongate hexagonal equatorial chambers, at least, in the peripheral zone. Cole (1952, pl. 20, figs. 1-3; pl. 21, figs. 7, 11, 12) published several vertical sections of specimens of L. vawghani from Panama in which the flange either did not develop or had been removed by erosion, These illustrations should be compared with the specimens illustrated as figures 1, 2, 5, 11, Plate 23. There are no differences. Inasmuch as the embryonic apparatuses in all the specimens under discussion are the same, the only apparent difference is the shape of the equatorial chambers, rhomboid in L. vavghani and elongate hexagonal in “L. tournoueri” and “L. tempani.” LEPIDOCYCLINA: COLE 307 Examination of numerous equatorial sections of L. vawghani from one locality (loc. 9) in Panama demonstrated that certain specimens (PI. 19, figs. 1, 4) develop elongate, hexagonal equatorial chambers which are similar to those of “L. tempanz’”’ (PI. 19, fig. 5) and “L. tournoueri’ (PI. 210 higs:.2, 3). Moreover, many specimens of “L. tempanii’’ (Pl. 20, fig. 4; Plies figs. 6, 9) and “L. tournouer?’ (Pl. 24, fig. 7) have rhomboid equatorial chambers similar to those of L. vaughani in the zone adjacent to the embryonic apparatus. Cole (1957, p. 106) suggested that ‘The shape of any equatorial chamber must be governed by the mass of protoplasm from the surface of which the chamber walls form... A slight radial elongation of the mass will produce diamond-shape walls, and slightly greater radial expan- sion will result in hexagonal shapes... The availability of food may be a factor in certain situations, as a smaller supply of food may result in longer extensions of the protoplasm.” The fact once established that many of these specimens have equatorial chambers which vary from rhomboid to elongate hexagonal shape destroys the single criterion by which separate species can be recongized. Thus, specimens referred variously to “L. towrnouert’, “L. tempani’’ and other supposedly valid species are in reality L. van ghani Cushman, European specimens (PI. 23, fie. 63 Coley 1961), pll 16, tig. <1; Drooger and Socin, 1959, pl. 1, figs. 1-4; pl. 2, all figs.) of L. tournoueri Lemoine and R. Douvillé, 1904, are similar to the American specimens which have been assigned to this European species. Eventually, it may be possible to demonstrate that the American specimens assigned at present to L. vaughani represent the same species as the European specimens, If so, L. vanghani Cushman, 19192, will become a synonym of L. tournouers. TYPE LOCALITIES OF CERTAIN AMERICAN LARGER FORAMINIFERA Table 1 gives the type localities of 12 species of American larger Foraminifera and the species which have been reported in association with the type at each of these localities. Data on the type localities are sum- marized under each species. American specimens which Vaughan (1924, p. 798; 19334, p. 25) referred to the European species, L. tournoueri, are not listed in this section as they are discussed elsewhere in this article. poulejot o1e SoureuU o4L aa,o d 78bo peproryL aoa) 199L = 18hSL rqny Jeaenllcs: ab 16989 — 1c989 ensyuy Ob d d L “ATWO saty[RIO] addy aJeUsIsap 0} a7TqQv} sty} UT ‘PSOPUN * oO wAUOUAS v ST Sps7s * ue ‘7urgsnva * Oo surduouds o1e 7u0p,IPp ° ue 17uUYdiulal “JT wx z TJ I iP FURY Tj j 2 Tepe *(9x9} 908) woURIG PUag pasrauqns Jo AWUIDIA dy} UT SaTTTPIOT YW: “Aypero] addy woIZ poytodar sardads Joyyo = d tadd4} = J d Ae d dx 1ST09 11cO9 9 voURTg eUod auozZ [rury vuvurg "}XO} UI pasn sraquinu AWTRIOT SHSN1 a]OD puv uvysnea 1¥004qG1;"q SnadxpI0.114§ (uewysny) srruaupurd vursdtsony uruysny srruaunsvusnd uvwysny rropun uewysny suvgsuvad aJOD pure uvysnea, urdu] x x. uvwysny rynased UPUIYSN) SPITE ». ULYSNVA LOUD x x a]IAnog "Y pure ‘way sayjauvs vurphropideT uvWYsNy) rappur PUulsatso4saja Hy (uvurysny) sesuaurvurd rulsauPry “SOTPIPLIOT Asay qe sad4q ay} YIM paqeDossy satads JoyjO pur vioyrurUeIO, JoSIvy uvoAUTY UIEIIDd Jo saIro0] adA[— Tf >TquIL, LEPIDOCYCLINA: COLE 309 Camerina panamensis (Cushman).—The type of Camerina pana- mensis is Nummulites panamensis Cushman (19194, p. 98) from USGS loc. 6025, Panama Canal Zone, a locality in the middle member of the Caimito Formation of the Gatun Lake Area (Woodring, 1957, p. 117). Topotypes have been described and figured by Vaughan and Cole (1941, p- 46, pl. 10, figs. 13, 14; pl. 11, figs. 1-4) and Cole (1952, p. 10, figs. 1-4; 1958, p. 272; 1964, p. 141, pl. 14, figs. 2, 6, 9, 10, 14). The types of Heterosteginoides panamensis Cushman (19194, p. 97, pl. 43, figs. 3-8, vot figs. 1, 2) (= Miogypsina panamensis) are also from USGS loc. 6025. Vaughan and Cole (1932, p. 510) described from this locality Lepidocyclina (Lepidocyclina) pancanalis, a species which Cole (1952, p. 18) considered to be a synonym of L. cavellez Lemoine and R. Douvillé, Cole (1952, p. 7) identified from locality 55 (= USGS loc. 6025) Camerina panamensis (Cushman) (as Operculinoides panamensis) , Hetero- stegina antillea Cushman (as H. israelskyi Gravell and Hanna and H. panamensis Gravell), Lepidocyclina (Lepidocyclina) canellei Lemoine and R. Douvillé, L. (L.) canellei (as L. (Lepidocyclina) parvula Cushman), L. (L.) yurnagunensis Cushman, and Miogypsina panamensis (Cushman). Heterostegina antillea Cashman.—tThe type was described (Cushman, 1919b, p. 49) from USGS loc, 6869, Long Island, Antigua. The type description was brief and the illustrations were inadequate with the exception of one transverse section (Cushman, 1919, pl. 5, fig. 1). Vaughan and Cole (1941, pl. 16, figs. 1, 2) published illustrations of two oriented median sections of topotypes. H. antillea is associated at its type locality with abundant specimens of Lepidocyclina undosa Cushman (19196, p. 65), the types of which came from this locality. Lepidocyclina canellei Lemoine and R. Douvillé.—This species, one of the earliest described American species (Lemoine and R. Douvillé, 1904, p. 20), was based on specimens from Pena Blanca, Panama Canal Zone, a locality submerged by Gatun Lake. Associated species at the type locality have not been reported. Woodring (1957, p. 29) stated “Foraminiferal soft limestone, such as that at locality 54 / doubtless corresponds to the foraminiferal marl of publications issued before the flooding of Gatun Lake.” Earlier, Cole (1952, p. 7) wrote “The lithology at locality 53 suggests descriptions of the marl at the submerged locality at Pena Blanca...” 310 BULLETIN 243 Cole (1952, p. 7) identified three species at locality 53 in association with typical specimens of L. canellei. They are: L. ywrnagunensis Cush- man (as L. ywrnagunensis morganopsis Vaughan), L, vanghant Cushman, and Miogypsina panamensis (Cushman) [as M. (Miogypsina) antillea (Cushman) }. At locality 54 f Cole (1957a, p. 314) reported Heterostegina antillea Cushman, H. israelskyi Gravell and Hanna (= H. antillea Cush- man), Mrogypsina (Miogypsina) antillea (Cushman) [=M. panamensis (Cushman) }, and Miogypsina (Miolepidocyclina) panamensis (Cushman) in association with L. canellez, Lepidocyclina dartoni Vaughan.—In 1933 (a, p. 36) Vaughan des- cribed a stellate species of Lepidocyclina as L. (Nephrolepidina) dartont from the “". . . northern slope of La Piedra, northeast of Jamaica, north- east of Guantanamo, Cuba, USGS loc. 7664." From this locality Vaughan (19334, p. 354) reported L. yarnagunensis Cashman, L. (Eulepidina) sp. cf. L. dilatata (Micht.), a European species, and L. (Nephrolepidina?) crassimar go Vaughan (1933a, p. 33), a new species. The illustrations given by Vaughan (1933a, pl. 26, fig. 4; pl. 27, figs. 5, 6; pl. 28, figs. 1, 14) demonstrate that the specimens which he compared with L. dilatata are without question L. widosa, The specimens identified as L. crassimargo are Lepidocyclina vanghani Cushman. Lepidocyclina gigas Cushman.—Large microspheric specimens from USGS loc. 6862 were named L. gigas by Cushman (1919), p. 64). Vaughan (1924, p. 799) was the first to suggest that L. gigas“... is a microspheric form which appears to belong to the same species as L. wndosa Cushman.” In 1933 (a, p. 41, pl. 22, figs. 1-4) Vaughan illustrated the internal structure of this form to supplement the external views given by Cushman (1919%, pl. i, figs. 3-5). USGS loc. 6862 is also the type locality of L. parvula Cushman (19194, p. 58) and L. tempanii Vaughan and Cole (in Vaughan, 1933a, pu 26). Lepidocyclina parvila Cashman.—The types of this species were obtained from USGS loc. 6862 “. . . lower bed at Hodges Bluff, An- tigua... (Cushman, 1919b, p. 58). The types of L. gigas Cushman (1919, p. 64) and L. tempanii Vaughan and Cole (7 Vaughan, 1933a, p. 26) were obtained from this locality. Lepidocyclina tempanii Vaughan and Cole.—The type description of L. (Nephrolepidina) tempanii Vaughan and Cole (in Vaughan, 1933, LEPIDOCYCLINA: COLE 311 p. 26) was based on specimens from “. . . near top of Hodge Hill, Anti- gua, collected by W. R. Forrest . . . [and] USGS loc. 6862, Hodge Point, collected by T. W. Vaughan” (Vaughan, 19334, p. 48). As Cushman (19196, p. 58, 64) based the descriptions of L. parvula and L. gigas on specimens obtained from USGS loc. 6862, this locality is designated as the type for L. tempaniz. Lepidocyclina undosa Cushman.—The types of L. wndosa Cushman (19194, p. 65) are external views of two sellaeform specimens associated with numerous other specimens of the same kind, and a few specimens of Heterostegina antillea Cashman on a slab of foraminiferal limestone from USGS loc. 6869, Long Island, Antigua. Vaughan (1924, p. 798, pl. 34, figs. 5, 6) illustrated the embryonic chambers of two topotypes, and Vaughan and Cole (1941, pl. 41, fig. 1) figured a vertical section of a topotype. Vaughan (1926, p. 394) was the first to emphasize that the embryonic chambers of this species vary from nephrolepidine to eulepidine in shape. Lepidocyclina vaughani Cashman.—The type illustrations of Lepido- cyclina vaughani Cushman (19194, p. 93) are external views of two speci- mens (pl. 37, fig. 4; pl. 38) from USGS loc. 6021 (= 6673), Panama Canal Zone. These specimens are weathered, therefore the equatorial chambers are exposed in the marginal zone of the test. Other specimens (Cushman, 1919a, pl. 37, figs. 1, 2, 3, 5) from USGS loc. 6255, identified by Cushman as L. vau ghanz, later were named L, (Lepidocyclina) miraflorensis Vaughan (1923, p. 253). Cole (19614, p. 373) placed L. miraflorensis in the synonomy of L, canelle: Lemoine and R. Douvillé. Vaughan (1923, p. 254) recorded “Lepidocyclina canellei .. . occurs with L, van ghani at the type locality...” Cole (1952, p. 6, 7) studied larger Foraminifera collected by S, M. Jones and W. P. Woodring, 1947, from locality 43 “One-quarter mile southwest of USGS locality 6021. . ., the type locality of Lepidocyclina (Nephrolepidina) vaughant, but pre- sumably a little higher stratigraphically in the middle member.” At this locality L. vaughani was associated with Heferostegina antillea Cushman and Lepidocyclina canellei Lemoine and R. Douvillé. Cushman (19194, p. 90, 93) stated that the types of L. vanghani (USGS loc, 6021 = 6673) came from the Emperador Limestone. Wood- ting (1957, p. 116) assigned this locality to the middle member of the 312 BULLETIN 243 Caimito Formation. The specimens erroneously assigned to L. vanghani by Cushman (1919z, pl. 37, figs. 1, 2, 3, 5) from USGS loc. 6255 were from the La Boca Formation (Woodring, 1957, p. 125). Lepidocyclina yurnagunensis Cushman.—Specimens from USGS loc. 7548, 2 miles south of Yurnaguna, Cuba, were named L. canellez Lemoine and R. Douvillé, var. ywrnagunensis Cushman (1919b, p. 57). Waughan (1924, p. 798), without discussion, listed this variety as L. (Nephrolepr- dina) yurnagunensis. In 1926 Vaughan (p. 393) stated “L. yurnagunensis seems to me to be specifically distinct from L. canellei... In L. caneller the embryonic chambers are strikingly similar in size, and the equatorial chambers are hexagonal.” So far this species is the only one recorded from this locality. Miogypsina panamensis (Cushman).—In the type description of Heterosteginoides panamensis Cushman (19194, p. 97) specimens from two localities were figured. The types of M. panamensis (Cushman, 19192, pl. 43, figs. 3-8) were from USGS loc. 6025, Panama Canal Zone, a locality in the middle member of the Caimito Formation of the Gatun Lake area (Woodring, 1957, p. 117). The other specimens (Cushman, 1919a, pl. 43, figs. 1, 2) were from USGS loc. 6011, Panama Canal Zone, from the Culebra Formation (Woodring, 1957, p. 122), and were referred to M. cushmani by Vaughan (1924, p. 813), a species whose type locality is USGS loc. 6012d, Panama Canal Zone. USGS Loc. 6025 is the type locality for Camerina panamensis (Cush- man) (19194, p. 98) and for Lepidocyclina pancanalis Vaughan and Cole (1932, p. 510), a species which Cole (1952, p. 18) placed in the synon- omy of L. canellei Lemoine and R. Douvillé. Heterostegina antillea Cushman and Lepidocyclina yurnagunensis Cushman also occur at this locality (see under Camerina panamensis, this article). Spiroclypeus bullbrooki Vaughan and Cole.—The types are from Kugler’s loc. K 482, Marac River, Trinidad, West Indies. This is the only known American species. At its type locality Vaughan and Cole (1941) recorded the following: Heterostegina antillea Cushman, Lepido- cyclina canellet Lemoine and R, Douvillé (as L. parvula Cushman), L. tempani Vaughan and Cole, L. wndosa Cushman (as L. gigas Cushman) L. yurnagunensis Cushman, and Mogypsina panamensis (Cushman) (as M. hawkinsi Hodson). LEPIDOCYCLINA: COLE 313 STRATIGRAPHIC IMPLICATIONS Specimens identified as Lepidocyclina vaughani have been reported from the Panama Canal Zone (Cushman, 19192, p. 93; Cole, 1952, p29: 1957a, p. 314), Antigua (Vaughan, 19332, p. 33), Carriacou West Indies (Cole, 19584, p. 221) and Costa Rica (Malavassi, 1961, p. 500). In addition, specimens identified by Barker (1932, p. 278) as L. (Nephro- lepidina) verbeeki Newton and Holland, an Indo-Pacific species, were considered by Cole (1952, p. 30) to be typical L. vaughani. Specimens assigned to L. tempanii were recorded from Antigua (Vaughan, 19334, p. 26) and Trinidad (Vaughan and Cole, 1941, p. 75). American specimens referred to the European species, L. tournoueri, were known from one Mexican locality (Vaughan, 193324, p. 25). Vaughan and Cole (1941) identified 18 species associated with L. tempanii at 11 localities in Trinidad. Some 17 species have been identified as occurring with L. vawghani at the various localities from which more or less complete faunal identifications have been made. However, these species can be reduced to eight supposedly valid species if known synonyms are evaluated. Table 2 gives the species associated with L. vaughani and L. tempanii in several geographic areas. Table 2.—-Number of localities in the Caribbean area in which the various species are associated. Panama Trinidad | Carriacou,; Mexico Number of localities analyzed Bohio Caimito 2 7 11 4 1 Archaias angulatus (Fichtel and Moll) 1 — — — — Camerina panamensis (Cushman) —- -- 4 2 1 Heterostegina antillea Cashman 1 3 9 3 — Lepidocyclina cancellet Lem. and 2 5 9 2 1 R. Douvillé *tempanii Vaughan and Cole — — 11 —_ 1 vaughant Cushman fe. 7 — 4 — undosa Cushman 1 oa 7 -- — yurnagunensis Cushman 1 3 5 — — Miogypsina panamensts (Cushman) 2 2 y + — =e ze 2 ae _ A86 pps 3.7 plsalon). -..\. 5 Sek des pognoeeoeee Antillean Cretaceous Rudists, Canal Zone Foraminifera, Stromatoporoidea. (Nos. 165-176)... 447 pps \53, Dis.) ch 3 iebgns ass saetekeseavcsea shane Venezuela geology, Oligocene Lepidocyclina, Miocene ostra- cods, and Mississippian of Kentucky, turritellid from Vene- zuela, larger forams, new mollusks, geology of Carriacou, Pennsylvanian plants. (Nos;*177-183). ~~ 448) pp.,.36 pls. -...2..ccaetetscternecrnenees ae osacens Panama Caribbean mollusks, Venezuelan Tertiary formations and forams, Trinidad Cretaceous forams, American-Eur- opean species, Puerto Rico forams. 12.00 12.00 14.00 14.00 14.00 18.00 16.00 16.00 16.00 16.00 16.00 16.00 16.00 16.00 16.00 16.00 Bn ie 4 i ¥? a 7 — Y ; 7 a . “ait : Scan Under Sar ot c. 4A PP OBS F479