Mollusca:Gastropoda) in the New World Emily H
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THE GENUS HARPA (MOLLUSCA:GASTROPODA) IN THE NEW WORLD EMILY H. VOKES TULANE UNIVERSITY I. ABSTRACT the eastern Atlantic (H. doris Roding - The name Harpa americana has been H. rosea Lamarck) and one in the eastern applied to every fossil Harpa specimen Pacific (H. crenata Swainson). Two fossil found in the New World. A second exam species come from the Miocene of Europe, ple of true H. americana from the Gurabo and of course, the two Caribbean and east Formation, Dominican Republic, shows ern Pacific forms mentioned above as the that none should be so referred and the New World fossil record.* form that occurs in the Agueguexquite The only described species of Harpa in the Caribbean is that one originally refer Formation of Mexico is here named H. is red by Gabb (1873, p. 214) to the West Afri thmica, n. sp. The examples taken from can "H. rosea" and subsequently renamed the Esmeraldas Formation of Ecuador are H. americana by Pilsbry (1922, p. 337). better referred to the living West Coast Known only to come from the Dominican species H. crenata. Republic, with neither exact locality nor stratigraphic level being certain, and II. INTRODUCTION based upon a single incomplete shell, H. The gastropod genus Harpa is a good ex americana for some time remained the ample of the group Woodring termed only name consi,dered for any example of "paciphiles," that is, present in the Ter Harpa in the Neogene of the Caribbean. tiary of the Caribbean but now extinct Thus, when Perrilliat Montoya (1960) there while still living on the eastern monographed the "Middle Miocene" fauna Pacific coast (see Woodring, 1966, p. 428, of the Agueguexquite Formation of table 1). Until recently the entire known Mexico, the species of Harpa, which oc occurrence of the genus in the fossil record curs in some numbers in the formation, of the New World consisted of only two was referred to Harpa americana. In 1979, specimens: the types of H. myrmia Olsson, Gibson-Smith and Gibson-Smith cited •he 1931, from the Oligocene of P eru; and H. Harpa they found in the Lower Mwc< ne americana Pilsbry, 1922, from the Cantaure Formation of Venezut la as "Miocene" of Santo Domingo. "Harpa cf. americana. ·• In 1981, Pit attr There are also in the New World several buted a specimen from the Pliocene beds Eocene species assigned to Eocithara at Esmeraldas, Ecuador, to H americana, Fischer, 1883, once considered as a sub While this paper wa. in pre% the write learn- genus of Harpa but now recognized as an ed that David Dockery is . he procc ,_ f de extinct genus. Rehder (1973) has sum scribing a new species of HaJ"!!'l fron e 3vr.am marized all of the harpid species, both fos Marl (Vicksburgian -Lower Oligocene 01 '\tlts sil and Recent, of the entire w orld (not a sissippi (Miss. Dept. Ne· Res, Bur G1 ol Bull 124, in press). Although there are umb1 r ol formidable task, for there are fewer than similarities to Harpa <:: <:: the re<; mblanc. <;: to 50 species total) and conclude d that Eocithara are such that questions are raised as Eocithara, with 16 species-groups, is to whether the Mississippi form <:: uly to be re known from beds of Upper Paleocene to ferred to Harpa <:: s. I partie a the 1: te1 ril Middle Miocene age and geograp hically area is ;;trong'y ~ancell3.te :m1 thE. rJtoconc from Pakistan to Europe and in North seem;, t, con• <::t of 01 lv two r under wh1 rl America from Mississippi to California. However th1 re <:: 3. t in cc h "' al 1~ n1 t net g1· nated md tha1 dt e~ vve1' y thP ni)S ol th~ 1- r VI· The genus Harpa s.s., which is d istin ou~ ..v ,orl Ifl vpic:::~.J Harpa f; sh101 Th1s guished by its larger size, multispiral coni Oligoce 1e 4 ecies <:: erns to be n tr ·n~ttwn b1 cal protoconch, and expanded parietal cal tween Eoc1 hara and Harpa bt..t t doe~ not ap lus, was considered by Rehder to include pear to be c >sclv re ated tc w ubsequent eleven living and five fossil species. Nine of specie:.. ol Harp:t tther m ti-e Ola World 01 1-}e the living forms are Indo-Pacific, as is one New 11 >r ,}: blv ·e• r <;en1,; a pare l1 1 develop fossil. Only one Recent species is found in ment in Ev• tthara tho t led to C! eud- nt 53 54 Tulane Studies in Geology and Paleontology Vol. 18 thereby extending not only the geologic This new specimen is not adult but that range but also the geographic range. proved to be an advantage, for it does still However, the Cantaure specimens were have preserved the protoconch (see pl. 1, soon removed from the synonymy of H. fig. 7b), which has four and one-half sharp americana by Gibson-Smith and Gibson ly conical whorls, and a noticeably cancel Smith (1982) and referred instead to H. late shell surface (see pl. 1, figs. 7a and 8). myrmia Olsson, which had the effect of This is in contrast to the Agueguexquite shortening the geologic range consider species, which has a protoconch of three ably, as the so-called "Middle Miocene" and one-half rounded whorls and a nearly Agueguexquite Formation is now known smooth adult shell. Therefore, the Mexi to be Middle Pliocene (Akers, 1972, p. 30) can form is not to be referred to H. and the Dominican beds from whence H. americana and it is described below as H. americana is assumed to have been col isthmica, n. sp. lected are now known to be latest Meanwhile, a second example of the Miocene-basal Pliocene in age (Saunders Ecuadorian aH. americana" was reco et al., 1980, p. 157). vered by Pitt (see pl. 1, fig. 5) and this So long as the sole example of H. shows the nature of the shoulder, missing americana from the Dominican Republic in the first specimen. This new shell has a was the incomplete type specimen we doubled row of shoulder spines, very like were not in a very good position to deter that in the living H. crenata and it is con mine which, if either, of the remaining re cluded that the Ecuadorian form is much ferences should continue to be included nearer to that species than it is to the with H. americana. But, after nine trips to Dominican or Mexican one. the Dominican Republic, over a period of Thus, in the Caribbean we now have seven years, in which a total of over 150 three species of Harpa: the youngest being localities were collected, at long last our H. isthmica, n. sp., most like the Recent perseverance was rewarded on the tenth West African H. doris; H. americana, from trip with a second small example of H. the Dominican Mio-Pliocene, probably an americana. cestral to the Recent West Coast H. ere- PLATE 1 Figures 1- 3. Harpa isthmica Vokes, n. sp. 1. (X 1'/2) USNM 377402 (holotype); height 32.8 mm, diameter 22.0 mm. Locality: TU 1046. Agueguexquite Fm., Veracruz, Mexico. 2. (2a- X 3; 2b- X 10; 2c- x 2) USNM 377403 (paratype A); height 21.3 mm, diameter 12.4 mm. Locality: TU 638. Agueguexquite Fm. , Veracruz, Mexico. 3. ( x 1 V2) USNM 377404 (para type B); height of fragment 36.5 mm, diameter of frag ment 30.4 mm. Locality: TU 1046. Agueguexquite Fm., Veracruz, Mexico. 4. Harpa doris Roding (X 1) Tulane Collection; height 55.8 mm, diameter 33.2 mm. Locality: Ivory Coast, West Africa. 5, 6. Harpa crenata Swainson 5. ( x 1'/,) CAS 60735; height 39.0 mm, diameter (incomplete) 19.8 mm. Locality: Quebrada Camerones ( = TU 1397). Esmeraldas Formation Prov. Es- meraldas, Ecuador. ' 6. ( x 1) Tulane Collection; height 55.0 mm, diameter 35.5 mm. Locality: Mazatl8.n, Sinaloa, Mexico (fishermen). 7, 8. Harpa americana Pilsbry 7. (7a- X 2; 7b- X 10) USNM 377397; height 23.0 mm, diameter 14.3 mm. Locality: TU 1444. Gurabo Formation, Dominican Republic. 8. ( x 1 V2) ANSP 4061 (holotype); height 33.3 mm, diameter 19.4 mm. Locality: Unknown, Dominican Republic. No.2 Harpa in New World 55 2a 1b 2c 2b 3 7a 7b PLATE 1 56 Tulane Studies in Geology and Paleontology Vol. 18 nata and its mid-Pliocene relative in Formation of the Paraguana Peninsula, Ecuador; and in the early Miocene of Ven Venezuela, may or may not be referable to ezue Ia, a species which may or may not be H. myrmia, described from the Lower the same as H. myrmia, from the Oligocene Chira Formation of north Oligocene of Peru. The latter question can ernmost Peru. Both forms have extremely not be resolved unless more material of the heavy axial ribs, which cross over the su Oligocene form is recovered, for the ture, forming a series of lamellar flanges unique type specimen is woefully incom across the subsutural ramp. The surface plete. But, on the basis of the material pre texture of the shell is cancellate between sently available, there is no reason why the the smooth axial ribs. The parietal callus is Venezuelan form is not to be referred to H. somewhat marginate and suggests that this myrmia. Both are marked by having the line may be independently derived from suture crossed by extensions of the ribs, a the ancestral Eocithara line. In particular, feature that is not developed in any other Rehder (1973, p.