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(:) FROM THE NEOGE~I<~ OF NORTHWESTERN ECUADOR

1 WILLIAM D. PITT i AND LOIS J. PITT l ABSTRACT East Pacific, as well as the Caribbean Recent and fossil faunas. NassarlUs mas­ The nassarid fauna from the Neogene tus Olsson, 1964, was descnbed froT!l the formations of northwestern Ecuador con­ Picaderos Formation, and occurs in both sists of five , none of which are the Angostura and Onzole formations. known from either the Recent or fossil fau­ Other species include: N. thielei Olsson, nas of the Caribbean or Pacific. There is 1964, from the Esmeraldas beds, with one species with affinities to the western affinities to an Indo-Pacific species: N. Pacific Recent fauna, one with affinities to hylus Olsson, 1964, known only from the­ Recent species in both the Caribbean and the Esmeraldas beds; and one species N. eastern Pacific faunas, the remaining repetiti Olsson, 1964, known on:y from •rp three species are known only from the Onzole FormatiOn. Ecuadorian Neogene, one of which, The Angostura Forme t!on of Late Nassarius mcmahini, n. sp., is confined to Miocene age (planktic for tminifpral zom• the Angostura Formation. N. 16) is the oldest formation di~cus~ed .n this paper. The name was first publi~hEd INTRODUCTION by Stainforth (1948, pp. 142, 143, 146 fol lowing an unpublished report of the It is generally accepted that there was International Ecuadorian Petroleum free passage of water between the Company ({lde Bristow and Hoff:!:e!:ter, Caribbean Sea and the Pacific Ocean, 1977' p. 32). through the Isthmus of Panama, for much The name "Esmeralda::: Formation \V<. of the Tertiary Period, allowing exchange proposed by Olsson ( 1942, p. 26Ul for tr1e of molluscan species. It has been suggest­ highly foraminiferal tuffaceous .,hale-, ed that the closure took place about 3.5 exposed along the coa,.:;t of E.'mt ra:da~ MA (Saito,1976). Jones and Hasson (1985) Province and the Esmeratdas Rivt r. accepted this date as the most likely time Bristow and Hoffstetter ( 1977, p. 141 rec when migration of species ended, with ognized the name Esmeraldas Formation final emergence of the Isthmus at about 3 but considered it to be part of the On.wle MA or later. Formation. Vokes (1988, p. 4) found .t tated that intended to be a monograph of the Family the name came from the village of N assariidae. Picaderos, which is near the center of th•' The five species discussed in this paper outcrop on the Rio Santiago. He h-,ted 16 are all endemic to the Neogene of north­ s pecies of mollusks from the PtcaderO'i western Ecuador. One species, N . mcmahi­ F ormation. Of these five occur at ni, n. sp., from the Angostura Formation Picaderos, four occur at Playa de Tign·, has affinities with species in the Recent three are not mentioned in the text, two on ly occur at Cueva de Angostura (Angostura Formation), one is from Playa de Tigre and Picaderos, and one is in tht' 1 Field Associate, Dept. of Geology, California text but not on the faunal list. Academy of Sciences, San Francisco, California, Olsson described the Picaderos For 94118 rna tion as occurring along the Rw 2 Home Address: 2444 38th Ave ., Sacramento, Santiago near the village of Picaderos: California, 95822 however, in the text he shows two

135 136 Tulane Studies in Geology and Paleontology Vol. 29

Picaderos species as occurring along the Class Cuvier, 1797 Rio Onzole and considerd some Picaderos Subclass species as being from the Borbon Milne-Edwards, 1848 Formation. Superorder Bristow and Hoffstetter (1977, p. 219, Cox, 1959 232) state that the name Picaderos Order NEOGASTROPODA Thiele, 1925 Formation is in disuse and consider it to Superfamily MURICOIDEA be equal to the Onzole Formation. The Rafinesque, 1815 Servicio N acional de Geologia y Minera Family Iredale, 1916 (Cuadro no. 83) also considers the Subfamily NASSARIINAE Picaderos Formation to be equal to the Onzole Formation; however, it places NASSARIUS Dumeril, 1806 Olsson's Playa de Tigre locality at the border of the Onzole and Angostura forma­ Nassarius DUMERIL, 1806, Zool. Anal., p. 166. tions and within the Angostura Formation. Type species: arcularia Linn., 1758, Olsson's collecting data is not precise by monotypy. enough to know exactly how close to Playa de Tigre he made his collections, thus it is Subgenus NASSARIUS s.s. difficult to know in which formation this NASSARIUS (NASSARIUS) REPETITI locality should be placed. Olsson, in the Olsson text, also listed some species as occurring Plate 1, figure 4 at Picaderos; Borbon Formation. It is pos­ Nassarius repetiti OLSSON, 1964, Neogene sible that some of his collecting data is in Moll. Northwest. Ecuador, p. 146, pl. 24, figs. 6, error and we may never know where some 6a. of his specimens were really collected. Description: "The shell is small, subovate, with an elevated sharp spire of about seven ABBREVIATIONS FOR whorls, including that of the small nucleus con­ REPOSITORY INSTITUTIONS sisting of one turn. The sculpture of the spire whorls consists of three rows of sharply beaded CASG - California Academy of Sciences, spiral cords which stand high above the adja­ San Francisco, California. cent sutural zones, which thus appear as if USNM- United States National Museum deeply excavated, the fine suture itself lying of Natural History, Washington D.C. away in the bottom as if hidden. The spirals on PRI - Paleontological Research Institution, the apertural side of the body whorl number Ithaca, New York. about seven with an additional eight or nine, much smaller ones, partly smooth lying along ACKNOWLEDGMENTS the basal sulcus and over the short beak in front. The .spiral cords are neatly beaded by We want to thank Elt:lily Vokes for mak­ small axials which faintly undulate the spiral ing Tulane University collections avail­ interspaces. On mature specimens, the sculp­ able and for help with the manuscript. We ture is obsolete on the back of the body whorl, also want to thank the following persons replaced there by a large smooth patch. The for making museum collections available: aperture is broadly subovate with a short longi­ Thomas Waller, United States National tudinal posterior canal at its junction with the Museum of Natural History; Gary parietal wall. The outer lip is thickened by a rib Rosenberg, Academy of Natural Sciences, on the back, and bears on its inner side four or Philadelphia; and Warren Allmon, more small denticles. There is a thin spread of Paleontological Research Institution. callus over the inner lip through which the Thanks also to Allan McMakin of Astoria, underlying spiral cords still show through; the Oregon, for the photography. parietal callus extends downward over the pil­ lar is a small shelflike feature [sic], bearing four small denticles a long it. The SYSTEMATIC PALEONTOLOGY notch is deep, its sides a little elevated, its fasci­ ole marked with small smooth spirals but has Phylum MOLLUSCA Cuvier, 1797 no bordering keel." (Olsson, 1964) No.4 N eogene Nassarius 137

Holotype: USNM 643999; height 14.6 mm, Pacific species N . iodes (Dall, l919 J, N. diameter 7.8 mm. moestus (Hinds, 1844), and !v. tiarula Type locality: Onzole Formation, Telembi, (Kiener, 1841), which occurs as far north Esmeraldas Prov., Ecuador. as San Francisco, Califorma 1 &'"> N. tegula Occurrence: Onzole Formation, Ecuador. (Reeve, 1853), see Skoglund, 1992, p. 85]. Nassarius iodes differs from N. mcmalani Discussion: Nassarius repetiti is similar in having three or four weak l1rations on toN. complanatus (Powys, 1835) from the the inner side of the outer lip and one Tropical East Pacific but differs in being much larger one posterior to the other:- at larger, more slender with the spire whorls about the center of the apertu.re The body slightly convex rather than straight sided. whorl of N. iodes has ten axml ribs, fcr tha'1 the NASSARIUS (NASSARIUS) MCMAKlNI rest, posterior to the other:;; at tht· rt·"ltPr of Pitt and Pitt, n. sp. the aperture. InN. moestus the body whorl Plate 1, figure 1 h as ten beaded, axial ribs with s-mooth Description: Shell small; five whorls, plus a interspaces, fading out on the last one protoconch of about two rounded, smooth third of the whorl. Nassarius tiurula di~'­ whorls, end broken. First spire whorl expanding fers in having five short lirations on the abruptly, axial sculpture appearing immediatly, inner side of the outer lip with one much but eroded on early whorls; remaining spire larger liration posterior to the re t, at tne whorls with eleven axial ribs crossed by about cen ter of the aperture. The PliO­ six spiral grooves (both specimens worn). Pleistocene and Recent western Atlant'c Suture distinct. Body whorl large with nine and Caribbean s pecies N. vibex (Say, strong, retractive axial ribs from suture to base, 1822), differs in having four liration-, on crossed by numerous, spiral, rounded grooves, the inner side of the outer lip and unt:c interspace of equal width. First axial posterior much larger liration posterior to the rel:>t, to aperture with signs of numerous spiral at the center of the aperture. The body bands; however, shell too worn to determine whorl has nin e axial ribs, cros..,eu by number or extent. Apertural callus pad heavy, twelve spiral bands, not fdding out on the 1 covering all of shell face; outer lip thickened. last one-third of the whorl, and the co - Anterior canal narrow, recurved. Outer lip with umellar callus is not well developed about six lirations with short secondary lira­ tions between. Subgenus PROFUNDINASSA Thiele. 1929 Etymology: This new species is named for Allan McMakin from Astoria, Oregon, who has Profundinassa THIELE, 1929, Handbuch der done most of the photography for this and past Systematische Weichtierkunde, pt. 1, p. 323. papers. Type species: Nassarius babylonims Watson, Holotype: CASG 67832.01; h eight 8.0 mm, 1882, by monotypy. diameter 5.1 mm. Paratype: PRI 44212; height 8. 7 mm, diame­ N ASSARIUS (PROFUNDINASSA) THIELE! ter 5.9 mm. Olsson Type locality: P-103, Angostura Formation, Plate 1, figures 5,6 sea cliffs about 300 meters west of Punta Verde, Nassarius (Profundinassa ) thielu OLSSO~, about 2 1/4 km east of the mouth of Rio Verde 1964, Neogene Moll. Northwest. Ecuador, p and about 22 km east of the bridge over Rio 145, pl. 24, fig. 7. Esmeraldas, Esmeraldas Prov. , Ecuador. Occurrence: Angostura Formation, Ecuador. Description: "Shell small, subovate, the spire twice the height of the aperture. The nucleus is Discussion: Nassarius mcmakini, n. sp., relatively large, turbinate, with two or three may be compared to the Tropical East smooth convex wh orls. Postnuclear whorls, 138 Tulane Studies in Geology and Paleontology Vol. 29

about four, are shouldered around the upper genera of Recent Mollusca, v. 1, p. 120. third, the angle of which carries a strong cord. Type species: Buccinum miga Bruguiere, The body whorl is large and forms about a half 1789, by subsequent designation, Cossmann, the whole shell, quadrate in shape, widest about 1901. the middle, the base strongly contracted. The NASSARIUS (UZITA) HYLUS sculpture is formed by about eight axial riblets Olsson which are noded by the shoulder cord and by Plate 1, figure 2 flat closeset spiral threads which cover the sur­ Nassarius (Uzita) hylus OLSSON, 1964, face between the shoulder angle and the beak. Neogene Moll. Northwest. Ecuador, p. 145, pl. The space above the shoulder angle is essential­ 37, figs. 10, 11 . ly smooth, except for small spirals. Aperture subovate, with a calloused shelf on the parietal Description: "Shell small, subovate, stout, the wall, the outer lip thin, smooth within. There is spire about twice the aperture and composed of a short, nassoid beak formed by the contraction about six whorls. The nuclear whorls are large, around the base. End of columellar pillar with a with a turbinate coil of two and one-half to sharp keel which margins the siphonal canal three rapidly enlarging smooth turas develop­ notch. Siphonal fasciole strong, bordered by a ing a small keel and a few axials near the end. nassoid keel." (Olsson, 1964) The sculpture consists of low, narrow, axial Holotype: USNM 644190; height 14.1 mm, rib lets ( 12 to 14) crossed by alternating spiral diameter 7.2 mm. threads producing a coarse cancellation. The Type locality: Esmeraldas beds, Punta Gorda, aperture is subovate, rimmed with a narrow, Esmeraldas Prov., Ecuador. nearly continuous, calloused peristome, finely Occurrence: Esmeraldas beds, Ecuador. denticulated on both lips. Beak short, encircled by a fasciolar fold ." (Olsson, 1964) Discussion: In comparing our specimens Holotype: USNM 644000; height 10.5 mm, of N. thielei to specimens from Olsson, diameter 5.8 mm. 1964, we find N. thielei to be a quite vari­ Type locality: Esmeraldas beds, Quebrada able species. Our specimens from Punta Camarones, Esmeraldas Prov., Ecuador. Gorda (fig. 6), mostly incomplete and Occurrence : Esmeraldas beds, Ecuador. worn, have the shoulder angle from almost turreted to moderately angled, the shoul­ Discussion: Nassarius exsarsus (Dall, der cord is almost non-existant in some 1908), a deep water species from the specimens. The axial ribs vary from verti­ Galapagos Islands, is similar in outline; cal in the figured specimen to protractive. however, the spire whorls are turreted, the The spiral threads are weak in our speci­ axial and spiral sculpture is closer togeth­ mens, although this appears to be due to er and more subdued. wear. The whorls on the figured specimen are straight sided, although on some of the NASSARIUS (UZITA) MASTUS fragmentary specimens they are more Olsson rounded. If we had more specimens this Plate 1, figure 3 could possibly prove to be an undescribed Nassarius (Uzita) mastus OLSSON, 1964, spec1es. Neogene Moll. Northwest. Ecuador, p. 146, pl. This common species, as noted by 38, fig. 7. Olsson, is similar toN. babylonica Watson, 1882, from the Philippine Islands but has Description: "The shell is small, stout, pupoid, a single cord at the shoulder rather than a its protoconch unusually large for its size, the double cord and the protoconch whorls are nuclear stage followed by a single, sculptured not carinate. Nassarius babylonica occurs turn (that of the body whorl). The large proto­ in the Recent western Pacific and Indian is composed of four, smooth convex Oceans. whorls, the first small, the others enlarging rapidly to form a turbinate coil. The nuclear Subgenus UZITA H. Adams stage is followed by the sculptured body whorl. and A. Adams, 1853 Assumption of the adult sculpture is gradual with the appearance of a few widely spaced Uzita H . ADAMS and A. ADAMS, 1853, The axial lines intersected by two or three spiral • 0. 4

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5

J>LXI E 1 Figc1res 1. assarius ( ras:wriusJ mcmakini Pitt and P1tt . n . :-; p CASG 67H:32.01 !holot. pe>l; hPight ' .0 mm. c!J ::lllwiPJ'5. 1 nlln t> ·, Locality: P -103, Ango.·lura Formntion 2. ossarws tL'zllaJ hvlus 01:-ison. 1~)(14 . . C'AS(~ 67 :3:3.0 1: he;ght 11.1 mm. dinnwt<•r G ~J mrn (;\ 1 .i1 Locality: P-100. Onzol> Fonn;Jtion, Ett~ador :3. assarius rUzitoJ mas/us Ol~son. 19f1J ..... C 'NM 64419 ( hololype 1; af'ter OJ....:-;on. pl.;).'), fig 7 '· heigl t ~ ~~ 1 Pll dwmeter 1. mm ! Xl~ . l l. Localil ·: Onzole Formntion. Ecuadt>r. ·L Nossarius r 'assanusJ repl'liti Olsson. 1 ~lf1 • ! L\ ·. 1 645:27 ( pnratyp '. al"tcr ( b .. on. pl. :21. fi ~. f) J; h!•Jgh • J.L~, mn diam •ter 6.4 mm r:>~Jrmn \ll J Locnhl.\: P-99. ( n7o]p Formatwn, E:cuador. 140 Tulane Studies in Geology and Paleontology Vol. 29 threads, the axials number about 12 and the Camarones, east of the village of Camarones, spirals finally to eight or nine, their points of about 10 km east of the mouth of Rio intersection forming small nodes on the final Esmeraldas, Esmeraldas Prov., Ecuador. portion of the whorl. The base is narrowly con­ P-103. Angostura Fm., sea cliffs about 300 me­ stricted a little above the short beaked anterior ters west of Punta Verde, about 2 114 km east canal, but there is no definite fasciole or a nas­ of the mouth of Ri6 Verde and about 22 km soid keel. The outer lip is thickened and a little east of the bridge over Ri6 Esmeraldas, reflected , marked with lirations within. Esmeraldas Prov., Ecuador. Columellar area is deeply excavated, smooth, the end of the pillar with a small ." (Olsson, LITERATURE CITED 1964) Holotype: USNM 644194, height 2.9 mm, BRISTOW, C. R. , and ROBERT HOFFSTET­ diameter 1.8 mm (not found ). TER, 1977, Ecuador in Lexique Stratigra­ Type locality: Onzole Formation, Picaderos, phique International, Second Edition, v. 5, Esmeraldas Prov., Ecuador. Amerique Latine, Fasc. 5A2, 410 p., 2 tables. Occurrence: Angostura and Onzole forma­ JONES, D. S., and P. F. HASSON, 1985, History tions, Ecuador. and development of the marine invertebrate faunas separated by the Central American Discussion: Olsson ( 1964, p. 146) noted Isthmus, in F. G. STEHLI and S. D. WEBB, that this species has a large protoconch eds., The Great American Biotic Interchange, and only one post-nuclear whorl, which p. 325-355, 6 text-figs, 3 tables. gives the appearance of an immature OLSSON, A. A. , 1942, Tertiary deposits of shell; however, this single whorl is sculp­ northwestern South America and Panama: tured and the outer lip is thickened and Eighth American Cong., Proc., v. 4, p. 231- lirate within as would be expected in an 287. adult shell. We have no explanation for OLSSON, A. A. , 1964, Neogene mollusks from this condition unless, as happens in some northwestern Ecuador. Paleontological Recent Tropical East Pacific species, after Research Institution, Ithaca, New York, 256 the eggs are laid, the adults leave the p., 38 pls. area. The juveniles could have been cov­ SAITO, TSUNEMASA, 1976, Geologic signifi­ ered by some natural condition and fos­ cance of coiling in the planktonic foraminifera silized with no adults present. Pulleniatina: Geology, v. 4, p. 305-309, 3 text­ We have not been able to locate the halo­ figs . type at USNM. There are two vials con­ STAINFORTH, R. M. , 1948, Applied micropale­ taining numerous specimens, one marked ontology in coastal Ecuador: Jour. Paleontol­ "Picaderos" and the other "various locali­ ogy, v. 22, no. 2, p. 113-151. ties." SKOGLUND, CAROL, 1992, Additions to the LOCALITY DATA Panamic Province Gastropoda (Mollusca) Literature: The Festivus, v. 24, Supplement, The following are Pitt fossil collecting 169 p. localities: VOKES, E. H. , 1988, (Mollusca: P-99. Esmeraldas beds, Punta Gorda (inter­ Gastropoda) of the Esmeraldas beds, north­ tidal) , about 8 km west of Las Palmas, western Ecuador: Tulane Stud. G eol. Esmeraldas Prov. , Ecuador. Paleont., v. 21, no. 1, p. 1-50, pis. 1-6, 1 table, P-100. Esmeraldas beds, east bank of Estero 15 text-figs.

February 26, 1997