A NEW SPECIES OF (:) FROM THE MIO-PLIOCENE OF NORTHWESTERN ECUADOR WILLIAM D. PITT 1 ·~ and LOIS J. PITT

ABSTRACT Brazil; E. guildingi (Sowerby, 1844, p. Two species of Enaeta occur in the Re­ 151), Antilles to Brazil; and E. reevei (Dall, cent fauna of the Tropical East Pacific: E. 1907, p. 353), Cuba to Honduras. There are cumingii (Broderip, 1832), and E. barnesii two species in the Recent Tropical East (Gray, 1825). The fossil record from the Pacific: E. cumingii (Broderip, 1832, p. Tropical East Pacific is represented by E. 33), and E. barnesii (Gray, 1825, p. 511); barnesii from the Pleistocene of Panama, both occur from the Gulf of California to and a new species, E. propecumingii from Peru. the Miocene Angostura Formation and the Enaeta propecumingii, n. sp., from the Pliocene Esmeraldas beds, Onzole Forma­ Late Miocene Angostura Formation of tion, of northwestern Ecuador. The oc­ northwestern Ecuador, is the earliest currence of E. propecumingii in the Angos­ known fossil record of the Enaeta in tura Formation is the oldest known fossil the Tropical East Pacific. The genus record of the genus Enaeta in the Tropical s.s. is known from older formations in the East Pacific. eastern Pacific, as far north as the state of Washington (Weaver, 1942) but not in the INTRODUCTION Tropical East Pacific. The genus Enaeta Adams and Adams, In the geological record there are more 1853, which has sometimes been consi­ species of the genus Enaeta in the western dered a subgenus of Lyria (Keen, 1971, p. Atlantic than in the Tropical East Pacific 619; Hoerle and Vokes, 1978, p. 106), is a and it is probable that the genus originated New World genus (Woodring, 1964, p. 289; in the western Atlantic. Abbott, 1974, p. 245; Skoglund, 1992, p. 94) represented by five species in the western Abbreviations for Repository Institutions Atlantic fossil record, with E. isabellae CAS - California Academy of Sciences, (Maury, 1910, p. 17) from the late Lower San Francisco, California. Miocene Chipola Formation of Florida the LACM - Los Angeles County Museum of earliest recorded occurrence. Other spe­ Natural History, Los Angeles, Califor­ cies included are: E. trechmanni Jung ma. (1971, p. 201), from the Middle Miocene PRI - Paleontological Research Institu­ Grand Bay Formation, Carriacou, West In­ tion, Ithaca, New York. dies; E. perturbatrix (Maury, 1917, p. 76), TU - Tulane University, New Orleans, from the Mio-Pliocene Gurabo Formation, Louisiana. Dominican Republic; and two species from USNM - United States National Museum the Pliocene Gatun Formation, Panama: of Natural History, Washington, DC. E. economia (Woodring, 1964, p. 289) and E. olssoni Hoerle and Vokes (1978, p. 116). The eastern Pacific has two known fossil ACKNOWLEDGMENTS species: E. barnesii (Gray, 1825), a Recent We want to thank Dr. Emily Vokes of species that also occurs in the Pleistocene Tulane University for help with the manu­ beds of the Burica Peninsula, Panama, script and for making available specimens and E . propecumingii, n. sp. The genus from Tulane collections, mcluding para­ Enaeta is represented in the Recent west­ types, and Allan Me Makin, from Ham­ ern Atlantic by three species: E. cyllenifor­ mond, Oregon, for the photography. mis (Sowerby, 1844, p. 151), Florida to SYSTEMATIC PALEONTOLOGY 1Field Associate, Dept. of Geology, California Academy of Sciences, San Francisco, Califor­ Phylum MO~LUSCA Cuvier, 1797 nia, 94118 Class GASTROPODA Cuvier, 1797 2Home address: 2444 38th Ave. , Sacramento, Subclass California, 95822 Milne-Edwards, 1848 123 124 Tulane Studies in Geology and Paleontology Vol. 28

Superorder Paratype D: USNM 484522; height 19.5 mm, Cox, 1959 diameter 8.8 mm; locality TU 1399, Esmeraldas Order Thiele, 1925 beds, Ecuador. Superfamily MURICOIDEA Paratype E: PRI 33194; height 17.9 mm, diam­ eter 9.2mm; locality TU 1399, Esmeraldas beds, Rafinesque, 1815 Ecuador. Family VOLUTIDAE Rafinesque, 1815 Paratype F: LACMIP 12298; height 20.9 mm, Subfamily LYRIINAE diameter 10.7 mm; locality TU 1399, Esmeraldas Pilsbry and Olsson, 1954 beds, Ecuador. Genus ENAETA Occurrence: Angostura Formation and E s­ H. Adams and A. Adams, 1853 meraldas beds of the Onzole Formation, Ecuador. Enaeta H . Adams and A. Adams, 1853, The Genera of Recent , v. 1, p. 167. Discussion: Enaeta propecumingii, n. Type species: harpa Barnes, 1824, by sp. , is represented by eight specimens, subsequent designation, Cossmann, 1899; Re­ three from locality P-101 , three from locali­ cent, eastern Pacific. [Voluta harpa Barnes, ty TU 1399, one from locality TU 1397 and 1824, non Mawe, 1823 = Voluta barnesii Gray, one from locality P-103. The fossil speci­ 1825 (as barnsii)]. mens examined are most similar to the fos­ sil E. trechmanni Jung, 1971, and theRe­ E NAETA PROPECUMINGII, n. sp. cent E. cumingii (Broderip, 1832). Enaeta Pitt and Pitt trechmanni differs in having a somewhat Plate 1, figure 1 globose, but evenly tapering spire and 18 Description: Shell elongate, of medium size, axial ribs rather than the slender, evenly with seven whorls; protoconch of about three, tapering spire and approximately 12 axial smooth, rounded whorls with a blunt ending. ribs of E. propecnmingii. The body whorl Spire whorls (o rnamentation appearing im­ of E. trechmanni does have a row of nodes mediately) evenly tapering, bearing two rows of anterior to the suture, a narrow smooth in­ nodes; one row of low nodes anterior to suture; a narrow smooth interspace; and one row of terspace, and axial ribs extending from the larger, e longated nodes just posterior to suture. interspace to the base of the shell; thus, be­ Body whorl with a row of nodes anterior to the coming the closest species to E. prope­ suture; narrow, smooth interspace, separating cumingii in the fossil record. Enaeta prope­ major axial sculpture, which consists of 11 to 13 cumingii is also similar to the Recent E. smooth, rounded ribs, extending from in­ cumingii but the latter has only a single terspace to base of shell. Aperture elliptical, row of nodes about midway on the spire more than one-half of shell height; outer lip whorls, in contrast to two rows in E. prope­ thickened, with a small blunt tooth midway on cumingii. Enaeta cumingii also differs in inner side of outer lip. Columellar wall with three strong plications anteriorly, and three that the body whorl has a smooth in­ faint plications, evenly spaced posteriorly. Ante­ terspace anterior to the suture, a row of rior canal recurved. nodes and nine axial ribs extending to the Etymology: From the Latin, prope meaning base of the shell with a secondary row of near and cumingii from the Recent E. cumingii nodes at their mid-point. (Broderip, 1832). The Recent E. cumingii apparently Holotype: CASG 67695 .01 ; height 29 .6 mm, di­ arose in the Tropical East Pacific from E. ameter 14 .3 mm (Plate 1, fig. 1). propecumingii. Type locality: P-101, Esmeraldas beds, road cut (south side), immediately west of the village LOCALITY DATA of Camarones, about 20 km (by road) east of the bridge over Rio Esmeraldas, Esmeraldas Prov., The following are Pitt localities: Ecuador ( = TU 1399). P-101. Esmeraldas beds, road-cut (south side), Paratype A: CASG 67695.02; height 31.4 mm, immediately west of the village of Camaro­ diameter 14.9 mm; locality TU 1397 , Esmeraldas nes, about 20 km (by road) east of the bridge beds, Ecuador. over Rio Esmeraldas, Esmeraldas Prov. , Paratype B: CASG 67695.03; height 24.2 mm, Ecuador. diameter 13.5 mm; locality P-103, Angostura P-103. Angostura Fm., sea cliffs about 300 me­ Formation, Ecuador. ters west of Punta Verde, about 2 114 km east Paratype C: USNM 484521; height 26.0 mm, of the mouth of Rio Verde and about 22 km diameter 13.7 mm; locality P-101, Esmeraldas east of the bridge over Rio Esmeraldas, Es­ beds, Ecuador. meraldas Prov., Ecuador. No.4 New Enaeta From Ecuador 125

2 b

PLATE1 Figures 1. Enaeta propecumingii Pitt and Pitt, n. sp. CASG 67695.01 (holotype); height 29:6 mm, diameter 14.3 mm (X 1.8). Locality: P-101, Esmeraldas beds, Ecuador. 2. Enaeta cumingii (Broderip, 1832) CASIZ 099363; height 33.2 mm, diameter 17.0 mm (X 1.4). Locality: PR-100, San Carlos Bay, Guaymas, Sonora, Mexico; Recent.

PR-100. Recent, San Carlos Bay, Guaymas, ADAMS, H., and A. ADAMS 185:3 58 Tlw gen Sonora, Mexico. era of Recent Moll usca, arranged acc-ord ~ ~ The following are Tulane University to their organization. London, v. I. p. 'vt-xl, I 484; v. 2, p. 1-661; v. 3, pis. 1-1:>8 [da1 ing o" localities: parts, v. 2, p . 661]. 1397. Esmeraldas beds, Quebrada Camarones, BRODERIP, W.J., 1832, CharaC''E'r" of new •me• cut-bank on east side of canyon, which is at cies of Mollusca and Conchif'era, collected by east edge of village of Camarones, 20 km (by Mr. Hugh Cuming: Zoo!. Soc. Londor Proc. road) east of bridge over Rio Esmeraldas, or for 1832, pt. 2, p. 25-33. approximately 10 km east of mouth of Rio Es­ DALL, W.H., 1907 , A re\ 1ew of' the American meraldas, Esmeraldas Prov. , Ecuador. Volutidae: Smithsoma n Misc. Col!.. v. 48, pt 1399. Esmeraldas beds, road-cut on west side of 3, no. 1663, p. 341-3 73 village of Camarones, which is 20 km (by EMERSON, W.K., 1985, T wo new species of' road) east of bridge over Rio Esmeraldas, at Lyria from the western Atla ntic (Gastro­ Esmeraldas, Esmeraldas Prov., Ecuador. poda:Volutidae): Nauttlus , v. 99, no. 1, p . 28- 33, 15 text-figs. GRAY , J .E., 1825, Monograph on the Cyp­ LITERATURE CITED raeidae, a family of testaceous Mollusca: ABBOTT, R. T. , 1974, American , Sec­ Zoo!. Journ., v. 1, p. 489-517, errata Sheet. ond Edition. Van Nostrand Reinhold Co. , pl. 12. . New York, 663 p., 24 color plates, 6405 text­ HOERLE, S.E., and E.H. VOKES, 1978, A r·e­ figs. view of the volutid genera Lyria and Fal- 126 Tulane Studies in Geology and Paleontology Vol. 28

silyria (Mollusca: Gastropoda) in the Tertiary Panamic Province Gastropoda (Mollusca) lit­ of the western Atlantic: Tulane Stud. Geol. erature: The Festivus, v. 24, Supplement, Paleont. , v. 14, no. 3, p. 105-130, 5 pis. 169 p. JUNG, P. , 1971, Fossil mollusks from Carriacou, SOWERBY, G.B., Jr. , 1844, Descriptions of six West Indies: Bulls. Amer. Paleontology, v. new species of Voluta: Zoo!. Soc. London, 61, no. 269, p. 14 7-262, 21 pis. , 2 text figs. Proc., pt. 12, p. 149-152. KEEN, A.M., 1971, Sea Shells of tropical west WEAVER, C.E. 1942, Paleontology of the ma­ America, Marine mollusks from Baja Califor­ rine Tertiary formations of Oregon and nia to Peru, Second Edition. Stanford Uni­ Washington: Washington Univ. [Seattle] versity Press, Stanford, California, i-xiv + Pubis. Geology, v. 5, pts. 1-3, 789 p ., 104 pis. 1064 p., 22 color plates, ca. 4,000 figs., 6 WOODRING, W.P., 1964, Geology and paleon­ maps. tology of Canal Zone and adjoining parts of MAURY, C.J., 1910, New Oligocene shells from Panama. Description of Tertiary mollusks. Florida: Bulls. Amer. Paleontology, v. 4, no. (Gastropods:Columbellidae to Volutidae): 1, p. 119-164, pis. 18-26. U .S. Geol. Survey, Prof. Paper 306-C, p. 241- SKOGLUND, CAROL, 1992, Additions to the 297 , pis. 39-4 7. November 15, 1995