THE (: FASCIOLARIIDAEl IN THE NEOGENE OF TROPICAL AMERICA

GEERAT J. VERMEI.J DEPARTMENT OF GEOLOGY AND CENTER FOR POP['LATIO.\' niOU>(r'r UNIVERS ITY OF CALIFORNIA AT DAVIS DAVIS. CALIFORNIA

ABSTRACT ABBREVIATIO. T~ ANSP -Academy of ~dtur .. l ~c1ew "', Phi­ The fasciolariid neogastropod genus ladelphia, Penn">) h arid Leucozonia Gray, 184 7, is represented in NMB - Naturhiston,rhe~ Mt...-.;et P1, Ba~t·l. the Neogene of tropical America by four Switzerland : L. ocellata (Gmelin, 1791) from USNM- United States Mt... eum et' ~c.tJrdl the Mare Formation (Late Pliocene) of History, Washington, D. C. Venezuela, not treated here; L. nassa PRI - Paleontological Rt>search Irt.,titJte, (Gmelin, 1791), of which L. caribbeana Ithaca, New York Weisbord, 1962, from the Playa Grande Formation (Pleistocene) of Venezuela is a SYSTE~ATICS synonym; L. rhomboidea (Gabb, 1873) from the Gurabo Formation (Early Family FASCIOLARIII>AE nray, L~:i3 Pliocene) of the Dominican Republic; and Subfamily PERISTERNI~. Af: L. striatula, n. sp., from the Cercado and Tryon, 1B80 lower Gurabo formations (Late Miocene to Genus LEUCOZO~IA Gray. ! 1..41 Early Pliocene) of the Dominican Republic. Leucozonia rhomboidea closely resembles Leucozonza GRAY, 1847, Zuo1 Jgical ::,oct tv or the Recent L. leucozonalis (Lamarck, London, Proceedings, v 15, o. h•. 1822) from the northwestern Caribbean Type species: IIGSW nmd '1, 1~'11 '1V and an unnamed Recent species from the monotypy. Brazilian island of Fernando de Noronha. Leucozonia striatula is very similar to the Remarks: Bullock ( 1971, dnd Lvons Recent L. brasiliana (d'Orbigny, 1841) (1991) have pointed out that Latlru'-' ~1b from the Atlantic coasts of Central and bulus (Linnaeus, 175kl, the Indo-Wes• South America. Pacific type species of \1o'1tro,·t, 1810, is very similar in the charac4-er of ·ts INTRODUCTION and shell to Leuro.zonia, and t'1< t Leucozonia may therrfore be a jumor "'ub­ Species of the fasciolariid neogastropod jecti ve synonym of Latirus. Woodnrr: genus Leucozonia are common inhabitants (1928) recognized that Latil·us i::s d hetero­ of low rocky intertidal and sublittoral com­ geneous group requiring thorou~,h re-v i­ munities in tropical America and the Cape s ion, especially among its Indo-We:t Verde Islands. Despite its ecological Pacific species. Lyons ( 1991 l refern d prominence, Leucozonia is poorly repre­ many fossil and Recent specie,; frmr tnt> sented in the fossil record. Only two fossil Americas to Latirus, but remarked thdt species, L. rhomboidea (Gabb, 1873) from none of these belong to Latints !n the the Neogene of the Dominican Republic strict sense, as typified by L. gihbulu~. I and L. caribbeana Weisbord, 1962, from have chosen to retain Leucozonia as a the Mare Formation (Late Pliocene) of valid genus, pending a thorough revision Venezuela have been described. A third of Latirus and of such related taxa ao species, the Recent L . ocellata (Gmelin, Ascolatirus Bellardi, 1884; Do/icholatiru~ 1791), has been recognized in the Mare Bellardi, 1884; Fractolatirus Iredale, L91ti; Formation (Gibson-Smith and Gibson­ Kuroda and Habe in Kuroda rt Smith, 1979), and will not be discussed al., 1971; Glaphyrina Finlay, 1927; fu.ther here. In this paper, I critically H emipolygona Rovereto, 1899 (Latirus assess the fossil species, and describe L. group 1 of Lyons, 1991); Hesperistania striatula, n. sp., from the Late Miocene and Gardner, 1944; Latirolagena Harris. 1897; Early Pliocene of the Dominican Republic. Latirulus Cossmann, 1889; Nrolatirus

129 130 Tulane Studies in Geology and Paleontology Vol. 29

Bellardi, 1884; Nodopelagia Hedley, 1915; folds, and a wider, more oblique, anterior Berry, 1958; . The holotype of L. Morch, 1852; Schumacher, 1817 caribbeana (PRI 26261) is an immature (Latirus group 2 of Lyons, 1991); Pseudo­ shell in which the inner side of the outer latirus Bellardi, 1884; Hutton, 1883; is smooth instead of being ornamented and Teralatirus Coomans, 1965. with granulated lirae. Like typical L. Leucozonia is characterized by a rela­ nassa from the insular Caribbean Recent tively short, broadly open anterior fauna, L. caribbeana has prominent shoul­ si phonal canal, by three or four basal col­ der knobs, three closely spaced spiral umellar folds, a well-marked parietal rib cords on the upper part of the last , at the adapical end of the inner lip, by and a well-developed labral tooth at the having the inner side of the outer lip end of a fourth primary cord, which is sep­ sculptured by granulated lirae, and usual­ arated from the three upper cords by a spi­ ly by having a labral tooth at the outer lip rally threaded sector. Recent L. nassa is situated at the end of a prominent spiral variable in the number and prominence of cord. Below this cord, the last whorl is con­ shoulder knobs and in the number of col­ stricted; that is, it has a concave lateral umellar folds. The shoulder always forms profile. At least two species, L. the widest part of the shell, and bears (Wood, 1828) from the tropical eastern seven to ten knobs. These knobs are Pacific and L. ocellata (Gmelin, 1791) from formed where the shoulder cord is crossed the tropical western Atlantic, differ from by strong axial ribs, whose lower end is typical Leucozonia by having the lirae on marked by the tooth-bearing cord. Recent the inner side of the outer lip smooth, by specimens have three to four (rarely five) having the outer lip planar instead of con­ columellar folds. I have seen Recent L. vex when viewed from the apertural side, nassa from Florida, the Bahamas, Saint and by lacking the labral tooth. The Maarten, Guadeloupe, Cura~ao, and the species discussed below all belong to the Brazilian island of Fernando de Noronha. more typical group of Leucozonia. Given the variation observed in Recent L. nassa, I see no basis for separating L. (Gmelin, 1791) caribbeana, and I therefore consider L. caribbeana as a junior subjective synonym L eu cozonia caribbeana WEISBORD, 1962, of L. nassa. Bulletins of American Paleontology, v. 42, no. 193, p. 361-362, pl. 32, figs . 9, 10. LEUCOZONIA STRIATULA, new species Figures 1, 2 Remarks: Weisbord (1962) named Leucozonia caribbeana from the Maiquetia Diagnosis: Leucozonia with well-developed Member of the Playa Grande Formation of axial ribs, fine spiral threads, a distinct labral Venezuela. Gibson-Smith and Gibson­ tooth, and a long anterior siphonal canal. Smith (1979) argued that this unit is con­ Description: Shell moderately large, maxi­ tinuous with,

ing from suture to suture in spire whorls, and umellar folds; well-de\elopE>r' pd~ l•Lll r bat from suture to tooth-bearing cord on last whorl; adapical end of inner lip i'1 Lug£> -;ptCI~"'f rt; axial ribs weak and irregular on last quarter elongate-ovate, ItS reigh. breddth fdtlO whorl of large specimens; outer lip sharp, crenu­ 2.7-2 .9; anterior siphonal canal hn~r, 23 Fl'r , f lated at edge, medially convex when viewed total apertural height; siphonc1 • ·"'!Ole low from apertural side; sharp labral tooth fo rmed rounded; absent.

at end of prominent spira l cord aL abapical end Holotype: USNM 490523 he1gH 3~ .. , r m of convexly rounded sector of last whorl; outer maximum diameter 22.7 mm, hPJght' t apPrt r, lip forms broad, shallow, concave sinus below plus canal 25.2 mm, brt>adth o• a Prtur e 9 1 labral tooth on abaxial side of siphonal canal; mm. inner side of outer lip with ten deeply recessed Para type I: NMB H 17R04 re1g•·• 48 mm lirae; inner lip adherent, with three basal col- maximum diameter 30.0 m"1

Figures 1, 2: Leucozonia striatula, new species. USNM 490523 (holotype), height 38.5 rnm; locality TU 1422. Figures 3, 4: L eucozonia rhomboidea (Gabb, 1873). USNM 490550, height 35.4 mm; locality TU 1354. 132 Tulane Studies in Geology and Paleontology Vol. 29

Paratype 2: ANSP 79634; height 44.0 mm, der and the suture. L. brasiliana was con­ maximum diameter 24.7 mm. sidered to be a synonym of L. nassa by Paratype 3: PRJ 44387; height 41.9 mm, max­ Marcus and Marcus (1962). Both L. brasil­ imum diameter 22.6 mm. iana and L. striatula differ from L. nassa, Type locality (holotype and paratypes 1 and however, in having about ten fine cords 2): locality TU 1422, Arroyo Bellaco, Cercado between the shoulder and the tooth-bear­ Formation, Dominican Republic. ing cord. In L. nassa, there are three large Other localities: Paratype 3, locality TU 1215, cords at and just below the shoulder, sepa­ lower Gurabo Formation, Rio Gurabo, bluffs rated from the tooth-bearing cord by fine above ford on Los Quemados-Sabaneta Road, threads. Most specimens of L. nassa , Dominican Republic. moreover, are less slender than L. brasil­ Other material examined: Three specimens, iana and L. striatula (diameter 57-69% as locality TU 1422; five specimens, locality TU compared to 52-61% of total shell height). 1215. Like L. rhomboidea (below), L. striatula Stratigraphic and geographic distribution: is found in reef facies (see Saunders et al., Cercado Formation (Late Miocene) and lower 1986). The living L. brasiliana occurs in Gurabo Formation (Early Pliocene), Dominican similar habitats. Republic. LEUCOZONIA RHOMBOIDEA (Gabb, 1873) Remarks: In several characters, Leuco­ Figures 3, 4 zonia striatula represents an intermediate condition between species of Polygona Lagena rhomboidea GABB, 1873, American Phi­ (Latirus group 2 of Lyons, 1991) and typi­ losophical Society, Transactions, new ser., v. cal Leucozonia. Like species of Polygona, 15, p. 218. the new species has a relatively long ante­ Leucozonia rhomboidea (Gabb). PILSBRY, 1922, rior siphonal canal and fine spiral sculp­ Proceedings of the Academy of Natural ture. In most species of Polygona, spiral Sciences of Philadelphia, v. 73, p. 345, pl. 21, sculpture is not as fine as in L. striatula, fig. 9. but the number of cords on the last whorl is low as in L. nassa, L. ocellata, the deep­ Description: Shell medium-sized, maximum water western Atlantic L. jacarusoi height 39.7 mm, fusiform; diameter 62-65% of Petuch, 1987, the eastern Atlantic (Cape total shell height; spire moderately low, last Verde Islands) L. triserialis (Lamarck, whorl comprising 61-65% of total shell height; 1822), or the eastern Pacific L. cerata protoconch consisting of one smooth whorl; (Wood, 1828), L. rudis (Reeve, 1846), and teleoconch consisting of six whorls separated by L. tuberculata (Broderip, 1833). Leuco­ adpressed, indistinct suture; last whorl without zonia striatula differs from tropical shoulder, evenly rounded above, weakly con­ American members of the genus Polygona stricted at base; spiral sculpture of penultimate by having a distinct labral tooth at the end whorl consisting of about sixteen fine threads; of a cord, which marks the abapical end of spiral sculpture of last whorl consisting of about the axial sculpture. ten weak basal cords, and of about twenty-three The species most similar to L. striatula very fine threads on upper part of whorl; axial is the Recent western Atlantic L. brasil­ sculpture consisting of about ten low, rounded iana (d'Orbigny, 1841), which is distrib­ ribs on spire whorls, absent on all or least half uted on the coasts of Central and South of last whorl; outer lip medially convex when America, south to southern Brazil. viewed from apertural side, with labral tooth at Leucozonia striatula differs from L. brasil­ end of uppermost spiral cord; inner side of outer iana by having a slightly lower spire (last lip with eleven lirae; inner lip adherent, with whorl comprising 65-68% instead of 63- three basal columellar folds; parietal rib situat­ 64% of total shell height), generally fewer ed at adapical end of inner lip in larger speci­ axial ribs (eight to eleven as compared to mens; aperture elongate-ovate, its height: ten to thirteen in L. brasiliana ), by having breadth ratio 2.5-2.7; anterior siphonal canal the widest part of the shell at the shoulder 27-31% of total apertural height; siphonal fasci­ instead of below the shoulder, and by hav­ ole rounded; umbilicus absent. ing finer spiral threads between the shoul- Material examined: ANSP 294 7 (lectotype), No.4 Neogene Leucozonia

Cibao region , Dominican Republic; ANSP 79165 NMB 16818), in the Early PliocPne pc.rt of (paralectotype), Cibao region, Dominican the Gurabo Formation. The assemb~age at Republic; NMB locality 16818, Canada de this locality represents a coral community Zamba, Dominican Republic; USNM 490550, of the kind in which L. leucozonalis carr Canada de Zamba, locality TU 1354, Dominican monly lives today (see Sa under-, et al., Republic (figs . 3, 4). 1986; Vermeij, 1973). Stratigraphic and geographic distribution: Two of five specimens of L rhomiJuideu Gurabo Formation (Early Pliocene), Canada de from TU 1354 bear eroded pits evidently Zamba, Dominican Republic. excavated by a hipponicid ga..,tr(Jpod. I shall report on these pit..., c..nd thE !l' mdker.., Remarks: Dall (1890, p. 106) thought in a separate paper. that Gabb's Lagena rhomboidea might be a young specimen belonging to the genus DISC'l'SSIO~ Mazzalina Conrad, 1860, but Pilsbry (1922, p. 345) correctly recognized it as a Its Late Miocene occurn nee mak L Leucozonia. The species closely resembles striatula the oldest known mE rnbE. r of thE. the Recent L. leucozonalis (Lamarck, genus Leucozonia. Furth E. r coli E. ctinr- tn 1822) from the northwestern Caribbean. older deposits will probablv reH,' C'VE n Both species reach the same maximum earlier representatives. The avc..ilablE t:vi height of about 40 mm. They have a dence shows that a :abr,tl tooth evoivE:d in rounded, finely spirally threaded last Leucozonia no lJ.ter tnan thE.' L tE whorl , and lack a distinct shoulder. Miocene. Many other vastroplla: ,.]..,o Leucozonia leucozonalis is generally some­ evolved a labral tooth during the l\-: on·ne what more slender (diameter 53-659f of in tropical America. These includE' 111em total shell height), lacks the axial sculp­ bers of the muricid :-.ubfc1miltes Muric nae, ture of the upper whorls of L . rhomboidea, Ocenebrinae, and Rdpaninae. Spt..ciE·. o~" and has a slightly longer canal (33-36% Pseudolividae and the olivid -,ubfamlly instead of 27-31 % of total apertural Ancillinae inherited the labral toot 1-t fl OP1 height). Paleogene ancestors. Odaly enot..gh, the An even more similar form is an large muricid fauna of the Miocent· anrl unnamed Recent species from Fernando de Pliocene of the Dommica1. Republ'c Noronha that previous authors have con­ (Vokes, 1989) is hightv unt..sual among sidered a variety of L. nassa (see Smith, Neogene tropical Amcr.can muncid as er1- 1890; Lopes and Alvarenga, 1955; blages in lacking species w1th a labral Matthews and Kempf, 1970; Leal, 1991 ). tooth. Pseudohvids and arcillmes are a\::o This Recent species differs from L. rhom­ absent from the Late Neogene I>ommic, n boidea by having a somewhat broader fauna. Did the evolutiOn of a labral tooth shell (diameter 68-7 4% instead of 62-65% in Leucozonia in the reef environments of of shell height), a relatively shorter the Dominican Republic fill an adaptive siphonal canal (17-21% instead of 27-31 (k gap left vacant by other group,.;? Fur~her of aperturallength), and in having ten low, work on the biogeography and phylogeny rounded, broad, obsolete axial ribs on the of gastropods with a labral tooth may sht•d last whorl instead of having the last whorl further light on this interesting possibiltty. partly or entirely devoid of axial ribs. The two previously known specimens of ACKNOWLEDGMENTE-: L. rhomboidea (ANSP 2947 and 79165) are immature shells with a height of about 10 I thank Emily H. Vokes for acce.:;s to mm. One of these CANSP 294 7, the lecto­ specimens, Janice Cooper and Mdry type) is the only specimen with the proto­ Graziose for technical assistance, and ~~F preserved. New material shows that Grant EAR-94-05537 for fundmg thi::-. the species attains a height of at least 39.7 research. mm, a height comparable to that of other species of Leucozonia. LITERATURE CITED Leucozonia rhomboidea has been collect­ ed at a single locality, TU 1354 (= locality BULLOCK, R. C, 1974, ContributiOn to the sys- 134 Tulane Studies in Geology and Paleontology Vol. 29

tematics of West Indian Latirus (Gastropoda: MARCUS, E. , and E . MARCUS, 1962, On ): Nautilus, v. 88, p. 69-79. Leucozonia nassa: Boletim da Faculdade de DALL, W. H., 1890, Contributions to the Terti­ Filosofia, Cii'mcias e Letras da Universidade ary fauna of Florida, with especial reference de Sao Paulo, Zoologia, v. 24, p. 11-30. to the Miocene Silex-beds of Tampa and the MATTHEWS, H. R. , and M . KEMPF, 1970, Pliocene beds of the Caloosahatchee River. Moluscos marinhos do norte e nordeste do Part I. Pulmonate, opisthobranchiate and Brasil. II. Moluscos do arquipelago de orthodont gastropods: Transactions of the Fernando de Noronha (com algumas referen­ Wagner Free Institute of Science, cias ao Atol das Rocas): Arquivos de Ciencias Philadelphia, v. 3, p. 1-473. Marinhas, v. 10, p. 1-53. GABB, W. M. , 1873, On the topography and PILSBRY, H. A., 1922, Revision of W. M. Gabb's geology of Santo Domingo: American Tertiary of Santo Domingo: Philosophical Society, Transactions, new ser., Proceedings of the Academy of Natural v. 15, p. 49-259. Sciences of Philadelphia, v. 73 , p. 305-445. GIBSON-SMITH, J., and W. GIBSON-SMITH, SAUNDERS, J. B. , P. JUNG, and B. 1979, The genus Arcinella (Mollusca: BIJU-DUVAL, 1986, Neogene paleontology in ) in Venezuela and some associated the northern Dominican Republic. 1. Field faunas: GEOS, v. 24, p. 11-32 [Escuela de surveys, lithology, environment, and age: Geologia y Minas, Universidad Central de Bulletins of American Paleontology, v. 89, no. Venezuela, Caracas I. 323, p. 1-79. GONZALEZ DE JUANA, C., J . M. ITURRALDE SMITH, E. A., 1890, Mollusca, in H. N. Ridley, DE AROZENA, and J. PICARD CADILLAT, Notes on the zoology of Fernando Noronha: 1980, Geologia de Venezuela y sus cuencas Journal of the Linnean Society, Zoology, v. 20, petroliferas: Ediciones Foninves, Caracas, v. p. 483-503. 2, p. 415-1031. VERMEIJ, G. J ., 1973, West Indian molluscan LEAL, J. H. , 1991, Marine prosobranch gas­ communities in the rocky intertidal zone: a tropods from oce anic islands off Brazil: morphological approach: Bulletin of Marine species composition and biogeography: Science, v. 23, p. 351-386. Universal Book Services/Dr. W. Backhuys, VOKES, E. H. , 1989, Neogene paleontology in Oegstgeest, 418 p. the northern Dominican Republic. 8. The LOPES, H . S ., and M . ALVARENGA, 1955, family (Mollusca: Gastropoda): Contribui~ao ao conhecimento dos moluscos Bulletins of American Paleontology, v. 97, no. da Ilha Fernando de Noronha , Brasil. 332, p. 5-94. Boletim do Institute Oceanografico, WEISBORD, N. E. , 1962, Late Cenozoic gas­ Universidade de Sao Paulo, v. 6, p. 157-190. tropods from northern Venezuela: Bulletins of LYONS, W. G., 1991, Post-Miocene species of American Paleontology, v. 42, no. 193, p. 1- Latirus Montfort, 1810 (Mollusca: 672. Fasciolariidae) of southern Florida, with a WOODRING, W. P., 1928, Miocene mollusks review of regional marine biostratigraphy: from Bowden, Jamaica. Part II: Gastropods Bulletin of the Florida Museum of Natural and discussion of results: Carnegie Institute History, Biological Sciences, v. 35, p. 131-208. of Washington, Publication 385, p. 1-564.

February 26, 1997