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smithsonian contributions to zoology • number 645

Types of Ampullariidae 

(Mollusca: Gastropoda) 

in the National Museum 

of Natural History, 

Smithsonian Institution, 

with Lectotype 

Designations

Robert H. Cowie, Kenneth A. Hayes, 

and Ellen E. Strong
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smithsonian contributions to zoology • number 645

Types of Ampullariidae 

(Mollusca: Gastropoda) 

in the National Museum 

of Natural History, 

Smithsonian Institution, 

with Lectotype 

Designations

Robert H. Cowie, Kenneth A. Hayes, 

and Ellen E. Strong

WASHINGTON, D.C.

2015
ABSTRACT

Cowie, Robert H., Kenneth A. Hayes, and Ellen E. Strong. Types of Ampullariidae (Mollusca: Gastropoda) in the National 

Museum of Natural History, Smithsonian Institution, with Lectotype Designations. Smithsonian Contributions to Zoology, 

number 645, iv + 13 pages, 1 figure, 2015.—The caenogastropod family Ampullariidae Gray, 1824, is a family of freshwater snails predominantly distributed in humid tropical and subtropical habitats in Africa, South and Central America, 

and Asia. The family includes the largest of all freshwater snails and frequently constitutes a major portion of the native 

freshwater malacofauna of these regions. Ampullariid taxonomy is confused primarily because most species were described 

on the basis of only shell morphology, which is highly variable within species yet relatively conservative among species 

within the family. The need for rigorous taxonomic treatment of the group is acute, and a starting point for such research 

is a comprehensive study of type material. With type or possible type material of 20 nominal species-group taxa of Ampullariidae, belonging to the genera Afropomus Pilsbry and Bequaert, 1927 (1 taxon, from Africa), Felipponea Dall, 1919 (2, 

South America), Forbesopomus Bequaert and Clench, 1937 (1, Asia), Lanistes Montfort, 1810 (2, Africa), Pila Röding, 

1798 (2, Asia), and Pomacea Perry, 1810 (11, South, Central, and North America), as well as 1 incertae sedis taxon in Pila

or Pomacea, the U.S. National Museum of Natural History, Smithsonian Institution, has significant holdings, for which 

this annotated catalogue is provided. To stabilize the nomenclature, lectotypes are designated for eight species-group taxa. 

No new synonyms are proposed, other than as a result of the provision of a replacement name for elongata Dall, 1921, 

non d’Orbigny, 1842, nec Rochebrune and Germain, 1904. Accounts are arranged alphabetically by original published 

species-group name. Information in each account includes original name and combination, citation to the original description, current taxonomic status, type material held by the U.S. National Museum (holotypes, paratypes, syntypes, lectotypes, 

paralectotypes, and one neotype) with catalogue number(s) and number of specimens in each catalogue lot, type locality, 

and other remarks or additional information as appropriate. All name-bearing types are illustrated.

Cover images: Apertural and abapertural views of ampullariid shells of (left to right) Paludina sinistrosa, Ampullaria pinei, 

A. superba, and A. (Felipponea) neritiniformis (from Figure 1J, H, K, F, respectively, herein).

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Library of Congress Cataloging-in-Publication Data

National Museum of Natural History (U.S.).

Types of Ampullariidae (Mollusca:Gastropoda) in the National Museum of Natural History, Smithsonian Institution, 

with lectotype designations / Robert H. Cowie, Kenneth A. Hayes, and Ellen E. Strong.

 pages cm. — (Smithsonian contributions to zoology ; number 645)

Includes bibliographical references and index.

1. Ampullariidae—Type specimens—Catalogs and collections—Washington (D.C.) 2. National Museum of Natural History (U.S.)—Catalogs. I. Cowie, Robert H., 1954– II. Hayes, Kenneth A. III. Strong, Ellen E. IV. Title.

QL430.5.A57N38 2015

594'.3209753—dc23

2015024160

ISSN: 1943-6696 (online); 0081-0282 (print)

Publication date (online): 25 September 2015

ZooBank registration: 15 September 2015 (LSID: urn:lsid:zoobank.org:pub:DB1AF5F1-0629-4839-BA1D-5848F6AB463E)

The paper used in this publication meets the minimum requirements of the American National Standard for Permanence 

of Paper for Printed Library Materials Z39.48–1992.
Contents

INTRODUCTION 1

Approach and Format of Accounts 2

Abbreviations and Acronyms 2

Acknowledgments 2

CATALOGUE OF TYPE MATERIAL OF AMPULLARIIDAE GRAY, 1824 3

atalanta Bequaert and Clench, 1937 3

aurostoma Lea, 1856 3

balanoidea Gould, 1850 4

canaliculata Lamarck, 1822 4

columellaris Gould, 1848 4

dalli Cowie, Hayes, and Strong, nomen novum 4

elongata Dall, 1921 4

gracilis Lea, 1856 6

meridaensis Pain, 1950 6

miamiensis Pilsbry, 1899 6

neavei Melvill and Standen, 1907 6

neritiniformis Dall, 1919 6

oligista Pilsbry and Olsson, 1953 6

palmeri Marshall, 1930 7

pinei Dall, 1898 7

sanjosensis Morrison, 1946 7

sinistrosa Lea, 1838 7

superba Marshall, 1926 7

tubaeformis Lea, 1856 8

turbinis Lea, 1856 8

zeteki Morrison, 1946 8

MATERIAL PREVIOUSLY CONSIDERED AS TYPES 9

zischkai Blume and Pain, 1952 9

REFERENCES 11

Types of Ampullariidae (Mollusca: 

Gastropoda) in the National Museum 

of Natural History, Smithsonian 

Institution, with Lectotype Designations

INTRODUCTION

The caenogastropod family Ampullariidae Gray, 1824 (junior synonym Pilidae Preston, 1915—Cowie, 1997; ICZN, 1999a; Bouchet and Rocroi, 2005), within the superfamily Ampullarioidea, is placed in the informal group Architaenioglossa, which also 

includes the Cyclophoroidea Gray, 1847, and Viviparoidea Gray, 1847 (Bouchet and Rocroi, 2005). The Campaniloidea Douvillé, 1904, is closely related, but the relationships 

among the four superfamilies remain unresolved (Ponder and Lindberg, 1997; McArthur and Harasewych, 2003; Strong, 2003; Simone, 2004; Colgan et al., 2007; Jørgensen 

et al., 2008; Ponder et al., 2008; Hayes et al., 2009).

Ampullariidae are freshwater snails predominantly distributed in humid tropical 

and subtropical habitats in Africa, South and Central America, and Asia (Hayes et al., 

2015). They include the largest of all freshwater snails (Pomacea maculata Perry, 1810, 

can attain greater than 165 mm in adult shell height; Hayes et al., 2012) and frequently 

constitute a major portion of the native freshwater mollusc faunas of these regions. Of 

14 Neotropical freshwater snail families, Ampullariidae constitute ~20 % of the Recent 

species (Strong et al., 2008).

Within the Ampullariidae, nine extant genera have been recognized in modern treatments (Cowie, 2015; Hayes et al., 2015). The great majority of the species are referred 

to just three genera: Lanistes Montfort, 1810 (Africa, including Madagascar), with 43 

recent species currently considered valid; Pila Röding, 1798 (Ampullaria Lamarck, 1799, 

and Ampullarius Montfort, 1810, are junior synonyms—Cowie 1997; ICZN 1999a) 

(Africa and Asia), with 29 species; and Pomacea Perry, 1810 (South, Central, and North 

America) with 97 species (Hayes et al., 2015). The remaining six genera contain only 

one or a few species: Afropomus Pilsbry and Bequaert, 1927, and Saulea Gray, 1868 

(1 species each) are African; Asolene d’Orbigny, 1838 (8 species), Felipponea Dall, 1919 

(3 species), and Marisa Gray, 1824 (2 species) are South American; and Forbesopomus

Bequaert and Clench, 1937 (1 species) is Asian (Hayes et al., 2015).

Taxonomy within the family is confused and complicated (e.g., Cowie et al., 2006), and 

delimitation of the genera and assignment of species to genera is in some cases uncertain 

1

 Pacific Biosciences Research Center, University of Hawaii, 3050 Maile Way, Gilmore 408, 

Honolulu, Hawaii, 96822, USA.

2

 Department of Biology, Howard University, 

415 College St. NW, Washington, D.C., 20059, 

USA.

3

 National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC 163, 

Washington, D.C., 20013, USA.

* Correspondence: kenneth.hayes@howard.edu 

Manuscript received 27 February 2015; accepted 14 May 2015.

Robert H. Cowie,1 Kenneth A. Hayes,2,3* and Ellen E. Strong3
2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

clarification, in square brackets. The type locality is as given in 

the original description, with original orthography within quotation marks, although when needed, changes for capitalization 

are in square brackets. Also in square brackets is additional type 

locality information when available.

Only the following type specimens are included in this catalogue: holotypes, syntypes, lectotypes, and one neotype, as well 

as paratypes and paralectotypes. Other specimens are only discussed as necessary for clarification, including non-type material of one species (Pomacea zischkai Blume and Pain, 1952). 

Catalogue numbers are those of lots in the mollusc collection 

in the USNM. The number of specimens in each lot is indicated. 

The current taxonomic status includes generic and subgeneric 

placement (the latter only if not the nominotypical subgenus), 

whether a valid taxon, and if not, the appropriate synonymy. 

Unless otherwise stated, generic placements and synonymies for 

New World taxa follow Cowie and Thiengo (2003) and for Old 

World taxa (Africa and Asia) follow Cowie (2015).

All interpretations follow the fourth edition of the International Code of Zoological Nomenclature (ICZN 1999b).

Abbreviations and Acronyms

ANSP Academy of Natural Sciences of Drexel University, Philadelphia, Pennsylvania, USA

Code International Code of Zoological Nomenclature

(ICZN, 1999b)

FMNH Field Museum, Chicago, Illinois, USA

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA

MMUE The Manchester Museum, University of Manchester, Manchester, UK

NMNH National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA

NMW National Museum of Wales, Cardiff, UK

n. syn. new synonymy

USNM U.S. National Museum (within the NMNH), 

Washington D.C., USA

ZSM Zoologische Staatssammlung München, Germany

Acknowledgments

We thank Neal Evenhuis for discussion of our interpretations of the Code and Norine Yeung for assistance with photography and help locating specimens in the collection. Funding 

was provided in part by National Science Foundation grant 

DEB-0949061.

or incorrect and in need of revision (Hayes et al., 2009, 2015). 

These problems are primarily because most species were described 

only on the basis of shell morphology, which is highly variable 

within species yet relatively conservative among species, such that 

the variable shell morphology of many species appears to overlap that of others. In the light of this confusion and the need for 

rigorous taxonomic research, it is crucial to study type material. 

Ascertaining the location of type material and its correct status is 

the necessary first step.

By far the largest museum collection of ampullariid type 

material is in the Natural History Museum, London. Other major 

collections, in roughly decreasing order of total numbers of lots 

of ampullariid types, include the Muséum national d’Histoire 

naturelle, Paris (its ampullariid types catalogued by Cowie and 

Héros, 2012); the Museum of Comparative Zoology, Harvard 

University; the Museum für Naturkunde, Berlin (its ampullariid 

types catalogued by Köhler and Glaubrecht, 2006); the Museo 

Nacional de Historia Natural, Santiago, Chile (its ampullariid 

type material dominated by the collections of Philippi and noted 

by Cowie, 2015, and by A. R. Kabat [Harvard University, Museum 

of Comparative Zoology] and E. V. Coan [Santa Barbara Museum 

of Natural History, Invertebrate Zoology], unpublished); the 

Academy of Natural Sciences of Drexel University, Philadelphia 

(its ampullariid types catalogued by Cowie et al., submitted); the 

National Museum of Wales, Cardiff; and the Forschungsinstitut 

und Naturmuseum Senckenberg, Frankfurt-am-Main. The ampullariid type material of the Zoologische Staatssammlung München 

is dominated by the South American collections of Spix, whose 

material was catalogued by Fechter (1983) and discussed briefly 

by Cowie et al. (2004). Many other museum collections, including 

the U.S. National Museum (USNM) within the National Museum 

of Natural History, Smithsonian Institution (NMNH), Washington, D.C., house additional type material (Cowie and Thiengo, 

2003; Cowie, 2015). The purpose of the present catalogue is 

to document the type material of Ampullariidae housed in the 

USNM, following Recommendation 72F.4 of the International 

Code of Zoological Nomenclature (ICZN, 1999b).

Approach and Format of Accounts

In this catalogue, taxa are arranged alphabetically by species-group name. The name and original combination are given 

following the name, verbatim as in the original description, and 

are followed by the bibliographic details of the original description. Bibliographic details of subsequent publications bearing directly on the original description are provided, as necessary for 
Catalogue of Type Material of 

Ampullariidae Gray, 1824

atalanta Bequaert and Clench, 1937

Forbesopomus atalanta Bequaert and Clench, 1937:54, pl. 2, figs. 1–7.

Current Taxonomic Status: Forbesopomus atalanta Bequaert and Clench, 1937; valid species (Cowie, 

2015:10).

Type Material: Paratype USNM 530146 (1 specimen).

Type Locality: “Lake Lanao, Misamis, Mindanao, Philippine Islands.”

Remarks: Only known from the type locality and the type series. Holotype, by original designation, figured, 

and with dimensions given (Bequaert and Clench, 1937:55, pl. 2, figs. 1–7), in the MCZ; additional 

paratypes in the MCZ and the NMW (Bequaert and Clench, 1937:55; Cowie, 2015:10). The gender 

of Pomus in Latin, and hence of Forbesopomus, is feminine.

aurostoma Lea, 1856

FIGURE 1A

Ampullaria aurostoma Lea, 1856:110.

Current taxonomic status: Pomacea aurostoma (Lea, 1856); valid species (Cowie and Thiengo, 2003:57).

Type Material: Lectotype USNM 106299 (1 specimen), here designated; paralectotypes USNM 1270616 

(formerly 106299X; 3 specimens); possible paralectotypes USNM 106273 (11 specimens).

Type Locality: “Carthagena” [Colombia].

Remarks: No holotype was designated and there are multiple type specimens, although the specimen 

(USNM 106299) figured subsequent to the original description by Lea (1866, pl. 22, fig. 4) is labeled “holotype”; based on the banding pattern of the shell, it is the one illustrated, and it bears the 

corresponding figure number inked on the body whorl. We therefore designate the shell in USNM 

106299 as the lectotype to stabilize the name. The paralectotypes (USNM 1270616) were not noted 

by Cowie and Thiengo (2003:57), who listed the possible paralectotypes in USNM 106273 (labeled 

“unconfirmed type”) as syntypes. A comment in the original handwritten ledger indicates that the 

latter lot comprises “fossil shells found 10 ft. below surface.”
4 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

dalli Cowie, Hayes, and Strong, 

nomen novum

FIGURE 1D

Felipponea dalli Cowie, Hayes, and Strong, nomen novum pro elongata Dall, 

1821 (see also elongata Dall).

Current taxonomic status: Felipponea dalli Cowie, Hayes, and Strong, 

2015; valid species (Cowie and Thiengo, 2003:49, as “elongata

Dall”).

Type Material: Lectotype USNM 333024 (1 specimen), here designated.

Type Locality: “Uruguay River, Dept. of Paysandú” (Dall, 1921:133, as 

“elongata”).

Remarks: The name elongata Dall is a junior primary homonym of elongata d’Orbigny and of elongata Rochebrune and Germain (Cowie 

and Thiengo, 2003:49; Cowie, 2015:35). There is no available and 

potentially valid synonym of elongata Dall, so a new substitute 

name, dalli Cowie, Hayes, and Strong, is here provided (Code, 

Article 60.3). The name honors William Healey Dall, American 

naturalist, malacologist, honorary curator at the NMNH, and the 

author of elongata Dall. No holotype of elongata Dall was designated, nor has it been determined for certain that Dall based his description on only a single specimen, although he provided dimensions of only one shell and a catalogue number (USNM 333024) 

of a lot, labeled “holotype,” that contains only one shell. On the 

basis of the label, Cowie and Thiengo (2003:49) listed this shell 

as the holotype; their listing did not constitute designation of the 

specimen as a lectotype. We here avoid assumption of a holotype 

(Code, Recommendation 73F) and formally designate the shell in 

USNM 333024 as the lectotype to stabilize the name. This specimen is automatically also designated as the lectotype of elongata

Dall (Code, Article 72.7).

elongata Dall, 1921

Ampullaria (Felipponea) elongata Dall, 1921:133.

Current nomenclatural and taxonomic status: Junior primary homonym of 

Ampullaria fasciata variété elongata d’Orbigny, 1842, and of Ampullaria kordofana variété elongata Rochebrune and Germain, 1904, replaced here by dalli Cowie, Hayes, and Strong (see dalli, above); objective synonym of dalli Cowie, Hayes, and Strong (Code, Article 72.7); 

Type Material: Lectotype USNM 333024 (1 specimen), here designated.

Type Locality: “Uruguay River, Dept. of Paysandú” (Dall, 1921:133).

Remarks: The name-bearing type (lectotype) of both elongata Dall and dalli

Cowie, Hayes, and Strong is the same specimen (Code, Article 72.7). 

Ampullaria elongata Bennett, 1831, is a nomen nudum, so unavailable, 

and therefore does not enter homonymy; it was noted by Cowie and 

Thiengo (2003:81) as not being an ampullariid.

balanoidea Gould, 1850

Ampullaria balanoidea Gould, 1850:196.

Current taxonomic status: Afropomus balanoidea (Gould, 1850); valid 

species (Cowie, 2015:10).

Type Material: Paralectotype USNM 611247 (1 specimen).

Type Locality: “Grand Cape Mount, Liberia.”

Remarks: No holotype was designated, nor has it been determined that 

Gould based his description on only a single specimen. The lectotype 

(MCZ 169054) was designated by Johnson (1964:45). The gender of 

Pomus in Latin, and hence of Afropomus, is feminine.

canaliculata Lamarck, 1822

FIGURE 1B

Ampullaria canaliculata Lamarck, 1822:178.

Current taxonomic status: Pomacea canaliculata (Lamarck, 1822); valid species (Cowie and Thiengo, 2003:58; Hayes et al., 2012:737).

Type Material: Neotype USNM 1185844 (1 specimen, in ethanol).

Type Locality: “Palermo, Buenos Aires, Argentina.”

Remarks: Hayes et al. (2012:741) considered the original type material to be 

lost and therefore designated a neotype. The original type locality was 

“les rivières de la Guadeloupe,” which may have been Lago Guadalupe (also referred to as Laguna Guadalupe, but now known as Laguna 

Setúbal) in Argentina and not the island of Guadeloupe, although this 

is conjecture (Pain, 1946:58; Hylton Scott, 1958:300; Thiengo et al., 

1993:68; Cazzaniga, 2002:74; Cowie and Thiengo, 2003:58). However, 

in designating a neotype the type locality was fixed as above.

columellaris Gould, 1848

FIGURE 1C

Ampullaria columellaris Gould, 1848:74.

Current taxonomic status: Pomacea columellaris (Gould, 1848); valid species 

(Cowie and Thiengo, 2003:60).

Type Material: Holotype USNM 5547, by monotypy.

Type Locality: “Province of Maynas, Peru.”

Remarks: No holotype was designated in the original publication. However, 

Gould (1852:128) implied that the original description was based on 

a single specimen (“the shell”), which he illustrated (Gould, 1852: 

pl. 9, figs. 147, 147a). The specimen in USNM 5547 is the shell illustrated, based on both the shell banding pattern and the configuration of 

repair scars, especially the prominent scar on the early part of the body 

whorl. We accept this specimen as the holotype by monotypy (Code, 

Article 73.1.2). The reference by Johnson (1964:57) to this specimen 

as the “figured holotype” was taken incorrectly by Cowie and Thiengo 

(2003:60) as fixation of a lectotype, citing Article 74.6 and Recommendation 73F of the Code.
NUMBER 645 • 5

FIGURE 1. Shells (apertural and abapertural views) of selected Ampullariidae species discussed herein. Dimensions given for each are shell 

height (length) in millimeters measured with digital calipers parallel to the columella. (A) Ampullaria aurostoma Lea, 1856, lectotype USNM 

106299, 28.9 mm; (B) A. canaliculata Lamarck, 1822, neotype USNM 1185844, 54.1 mm; (C) A. columellaris Gould, 1848, holotype USNM 

5547, 63.8 mm; (D) A. (Felipponea) dalli Cowie, Hayes, and Strong, nomen novum pro Ampullaria (Felipponea) elongata Dall, 1921, lectotype 

USNM 333024, 27.6 mm; (E) A. gracilis Lea, 1856, lectotype USNM 106323, 35.4 mm; (F) A. (Felipponea) neritiniformis Dall, 1919, lectotype 

USNM 332780, 33.1 mm; (G) A. palmeri Marshall, 1930, holotype USNM 380696, 66.2 mm; (H) A. pinei Dall, 1898, lectotype USNM 

152699, 59.8 mm; (I) Pomacea cumingii sanjosensis Morrison, 1946, holotype USNM 542136, 44.4 mm; (J) Paludina sinistrosa Lea, 1838, 

lectotype USNM 106348, 32.0 mm; (K) A. superba Marshall, 1926, holotype USNM 362863, 52.0 mm; (L) A. tubaeformis Lea, 1856, 

holotype USNM 106329, 33.1 mm; (M) A. turbinis Lea, 1856, lectotype USNM 106341, 61.9 mm; and (N) Pomacea zeteki Morrison, 1946, 

holotype USNM 542137, 40.5 mm. Scale bar = 1 cm.
6 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

neavei Melvill and Standen, 1907

Lanistes neavei Melvill and Standen, 1907:6, unnumbered pl., fig. 1.

Current taxonomic status: Lanistes neavei Melvill and Standen, 1907; valid 

species (Cowie, 2015:21).

Type Material: Syntype USNM 708877 (1 specimen).

Type Locality: “Kapopo” [southwest of Ndola, Zambia (Neave, 1907: 

unnumbered pl.)].

Remarks: USNM 708877 is labeled “paralectotype,” but no lectotype appears to have been designated. Other syntypes are in various museums 

(Cowie, 2015:21), and the specimen figured in the original publication 

(MMUE EE.7815) is the best candidate for future designation as the 

lectotype (Code, Recommendation 74B).

neritiniformis Dall, 1919

FIGURE 1F

Ampullaria (Felipponea) neritiniformis Dall, 1919:10.

Current taxonomic status: Felipponea neritiniformis (Dall, 1919); valid 

species (Cowie and Thiengo, 2003:49).

Type Material: Lectotype USNM 332780, here designated; paralectotypes 

USNM 1270739 (formerly 332780X; 3 specimens).

Type Locality: “Rio Uruguay, Department of Paysandu” [Uruguay].

Remarks: The original description was based on more than one specimen 

and Dall did not designate a holotype, although he provided dimensions of only one shell and a single catalogue number (USNM 332780) 

of a lot (labeled “holotype”) that contains only one shell. Cowie and 

Thiengo (2003:49) listed this shell as the holotype, based on the label; 

their listing did not constitute fixation of the specimen as a lectotype. 

We designate the shell in USNM 332780 as the lectotype to stabilize 

the name. According to the handwritten ledger there was also a radula 

preparation derived from the original lot, but it could not be located.

oligista Pilsbry and Olsson, 1953

Pomacea (Effusa) oligista Pilsbry and Olsson, 1953:98, pl. 6, fig. 6.

Current taxonomic status: Incertae sedis in Pomacea (Effusa).

Type Material: Paratypes USNM 605847 (3 specimens).

Type Locality: “[O]n the road . . . from Cartagena to Barranquilla . . . a freshwater lake known as the Cienaga de Luruaco” [Colombia].

Remarks: The holotype (by original designation) and additional paratypes 

are in ANSP (Baker 1964:168; Cowie and Thiengo, 2003:55). Synonymized by Pain (1956:76–77) with Pomacea (Effusa) planorbula

(Philippi, 1852), which he treated as a valid species but which Cowie 

and Thiengo (2003:55) retained as only an infraspecific taxon following Baker (1930:12, 24) and Pilsbry (1933:72), the latter considering 

Ampullaria planorbula Philippi, 1852 “to be the young stage of some 

variety of P. (Effusa) glauca (L.).” Pending further research, we retain 

oligista Pilsbry and Olsson, 1953, as an infraspecific taxon of undetermined status and specific affinity.

gracilis Lea, 1856

FIGURE 1E

Ampullaria gracilis Lea, 1856:110.

Current taxonomic status: Pila gracilis (Lea, 1856); valid species (Cowie, 

2015:37).

Type Material: Lectotype USNM 106323 (1 specimen), here designated; paralectotype USNM 1272906 (formerly 106323a) (1 

specimen).

Type Locality: “Siam” [Thailand].

Remarks: No holotype was designated and the description was based on two 

specimens (Lea, 1866:113). The specimen (USNM 106323), figured 

subsequent to the original description by Lea (1866, pl. 22, fig. 1), is 

labeled “holotype”; based on the banding pattern of the shell, it is the 

one illustrated, and it bears the corresponding figure number inked on 

the body whorl. We here designate this specimen (USNM 106323) as 

the lectotype to stabilize the name.

meridaensis Pain, 1950

Pomacea (Limnopomus) meridaensis Pain, 1950:109.

Current taxonomic status: Synonym of Pomacea camena Pain, 1949 (Pain, 

1957:175; Cowie and Thiengo, 2003:69).

Type Material: Paratypes USNM 595032 (2 specimens).

Type Locality: “Merida, Venezuela.”

Remarks: The holotype, designated by Pain (1950:110), is the specimen 

figured by Alderson (1925: pl. 11, fig. 7), the whereabouts of which 

is not known to us. Additional paratypes are in various museum 

collections (Cowie and Thiengo, 2003:69); one of them may in fact 

be the holotype.

miamiensis Pilsbry, 1899

Ampullaria miamiensis Pilsbry, 1899:365 [1927:252, pl. 22, figs. 5

(lectotype), 6, 7].

Current taxonomic status: Synonym of Pomacea paludosa (Say, 1829).

Type Material: Possible paralectotypes USNM 160193 (2 specimens).

Type Locality: “[I]n a creek flowing from the Everglades near Miami, Dade 

County, in southeastern Florida.”

Remarks: Use of the term “type” by Pilsbry (1927:253, pl. 22, figs. 5–7) 

for a specimen in the ANSP constitutes a valid lectotype designation according to the Code (Article 74.5); see also Baker (1964:168) 

and Cowie and Thiengo (2003:69). Additional paralectotypes are 

in various museum collections (Cowie and Thiengo, 2003:69). 

Treated by Clench and Turner (1956:122) as a “local population” 

of Pomacea paludosa (Say, 1829), which we here consider to constitute synonymy, although Cowie and Thiengo (2003:69) retained it 

noncommittally as an infraspecific taxon of that species. Cowie and 

Thiengo (2003:69) erroneously noted only one specimen in USNM 

160193.
NUMBER 645 • 7

palmeri Marshall, 1930

FIGURE 1G

Ampullaria palmeri Marshall, 1930:4, pl. 1, figs. 5, 8.

Current taxonomic status: Pomacea palmeri (Marshall, 1930); valid species 

(Cowie and Thiengo, 2003:71).

Type Material: Holotype USNM 380696, by original designation; paratypes 

USNM 380697 (4 specimens).

Type Locality: “[S]mall stream in a dense jungle, 13 kilometers south of 

Puerto Santos, Province of Santander del Norte, Republic of Colombia.”

Remarks: The USNM catalogue number and dimensions of the holotype 

were given by Marshall (1930:5), who also noted three paratypes as 

USNM 380697 (in fact there are four specimens in this lot). The lot 

(USNM 380698; 133 specimens) described by Marshall (1930:5) 

as “embryonic shells” is not type material, as its listing separately 

from the “type” and “paratypes” expressly excludes it from the type 

series (Code, Article 72.4.6).

pinei Dall, 1898

FIGURE 1H

Ampullaria pinei Dall, 1898:75.

Current taxonomic status: Synonym of Pomacea paludosa (Say, 1829) 

(Clench and Turner, 1956:120; Cowie and Thiengo, 2003:71).

Type Material: Lectotype USNM 152699 (1 specimen), here designated; 

paralectotypes USNM 1270740 (formerly 152699X; 11 specimens).

Type Locality: “Homosassa River, Florida.”

Remarks: A label, noted by Cowie and Thiengo (2003:72), indicated that 

USNM 152699 was the figured “type,” although Dall (1898:75–76) 

did not designate or figure a type, and there are multiple type specimens. We designate the shell in USNM 152699 as the lectotype to 

stabilize the name.

sanjosensis Morrison, 1946

FIGURE 1I

Pomacea cumingii sanjosensis Morrison, 1946:6, pl. 1, fig. 1.

Current taxonomic status: Pomacea cumingii sanjosensis Morrison, 1946; 

valid subspecies (Cowie and Thiengo, 2003:74).

Type Material: Holotype USNM 542136, by original designation; paratypes 

USNM 588458 (19 specimens), USNM 588459 (10 specimens, 4 operculae), USNM 588511 (3 specimens), USNM 598924 (25 specimens), 

USNM 599037 (12 specimens).

Type Locality: “[T]hree small streams (not of contiguous drainage) on the 

west side of San José island” [Panama; the catalogue ledger entry associated with USNM 599037 indicates “about 1⁄4 mi. from mouth” but does 

not specify which stream.]

Remarks: Morrison (1946:7) gave the USNM catalogue number and 

dimensions of the “type”; although he also gave dimensions of 

the “largest of many specimens seen” (according to the ledger, the 

“largest specimens measured” were from USNM 588511) and of 

the “smallest adult shell,” he was not more explicit in identifying 

the paratypes. There are 10 specimens in USNM 588459 but only 9 

recorded in the ledger, although the lot includes several hatchlings, 

one of which could have been easily overlooked. The catalogue ledger notes 27 specimens in USNM 598924; the two missing specimens were not located in adjacent lots.

sinistrosa Lea, 1838

FIGURE 1J

Paludina sinistrosa Lea, 1838: 90, pl. 23, fig. 78.

Current taxonomic status: Lanistes sinistrosus (Lea, 1838); valid species 

(Cowie, 2015:26).

Type Material: Lectotype USNM 106348 (1 specimen), here designated.

Type Locality: “East Indies.”

Remarks: No holotype was designated, although USNM 106348 was 

labeled “holotype.” Lea obtained from a Miss Hodges “one of the 

two specimens which were in her fine cabinet” and noted both his 

own and her cabinet as repositories. Thus, the description may or 

may not have been based on just the one specimen actually given to 

him. We avoid assumption of a holotype (Code, Recommendation 

73F) and designate USNM 106348 as the lectotype to stabilize the 

name. This rather nondescript shell has the catalogue number and 

“type” written on the inside of its aperture and is consistent with 

the original figure of Lea (1838:fig. 78), but it bears no distinguishing features or figure number inked on the body whorl to conclusively deem it as such. Pilsbry and Bequaert (1927:195) placed it in 

Lanistes, which is African only, and considered the original locality 

erroneous, having seen specimens from Gabon and Mozambique 

(though it seems unlikely that these would belong to a single species) and mentioning possible records from Tanzania. The genus 

Lanistes in Latin is masculine.

superba Marshall, 1926

FIGURE 1K

Ampullaria superba Marshall, 1926:3, pl. 1, fig. 9.

Current taxonomic status: Pomacea superba (Marshall, 1926); valid species 

(Cowie and Thiengo, 2003:75).

Type Material: Holotype USNM 362863, by original designation.

Type Locality: “Cienaga Totuma, Department of Atlantico, United States of 

Columbia” [Colombia].

Remarks: The “type” was figured and its USNM catalogue number and 

dimensions were given by Marshall (1926:4, pl. 1, fig. 9). The shell 

has the catalogue number and “type” written on the inside of its 

aperture. The original handwritten ledger indicates that it is the figured type, and the pattern of banding on the shell matches that in 

the illustration precisely.
8 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Type Material: Lectotype USNM 106341, here designated; paralectotypes 

USNM 1270744 (formerly 106341X; 2 specimens).

Type Locality: “Siam” [Thailand].

Remarks: No holotype was designated, although USNM 106341 was labeled 

“holotype,” and the description was based on three specimens (one 

adult, two young), according to the later description of Lea (1866:114). 

The shell in USNM 106341 is the shell illustrated by Lea (1866: pl. 22, 

fig. 2), based both on the banding pattern and the very clear repair scars 

or growth arrests, especially that at the start of the body whorl; it bears 

the corresponding figure number inked on the body whorl. We here 

designate this shell as the lectotype to stabilize the name. The paralectotypes (USNM 1270744, formerly 106341X), which are indeed smaller 

than the lectotype, were considered not conspecific with the lectotype, 

according to a note associated with the lot by J. P. E. Morrison, but this 

issue requires further research as the paralectotypes could simply be 

young individuals of the same species, as stated by Lea (1866:14).

zeteki Morrison, 1946

FIGURE 1N

Pomacea zeteki Morrison, 1946:8, pl. 1, fig. 3.

Current taxonomic status: Pomacea zeteki Morrison, 1946; valid species 

(Cowie and Thiengo, 2003:77).

Type Material: Holotype USNM 542137, by original designation; paratypes 

USNM 542138 (36 specimens).

Type Locality: “Chagres River near Gatuncilla, Republic of Panamá.”

Remarks: Additional material, listed as paratypes by Cowie and Thiengo 

(2003:77) and Cowie (2015:65), is present in various other museum 

collections.

tubaeformis Lea, 1856

FIGURE 1L

Ampullaria tubæformis Lea, 1856:110.

Current taxonomic status: Incertae sedis in Ampullariidae; valid species 

(Cowie, 2015:56).

Type Material: Holotype USNM 106329, by monotypy.

Type Locality: Unknown.

Remarks: The original publication neither stated nor implied that the description was based on a single specimen. However, Lea (1866:114) subsequently indicated that it was indeed based on only a “single specimen” 

and USNM 106329 (labeled “holotype”), based on the banding pattern 

and the clear repair scar, is the specimen illustrated by Lea (1866: pl. 22, 

fig. 3); it bears the corresponding figure number inked on the body whorl. 

We accept this specimen as the holotype by monotypy (Code, Article 

73.1.2). Although Lea (1856:110) gave no locality, he subsequently (Lea, 

1866:114) said that the specimen “came with many other shells from India,” and notes associated with the specimen and the original accession 

ledger (as “India?”) also suggest this to be the case. However, the original 

ledger entry was crossed out and replaced with “Brazil” by J. P. E. Morrison and dated September 1955; a note with the specimen written by 

Morrison at the same time indicates, “probably from Rio di [sic] Janeiro 

or Brazil near Bahia.” The location also has been suggested as “Siam” 

(Gaudion 1880:41). The shell appears more like that of a species of Pomacea (New World) than of Pila (Old World), but its correct generic placement and possible synonymy require further research. Not listed by Sowerby (1909:358, 1910:61, 1916:73) or by Cowie and Thiengo (2003).

turbinis Lea, 1856

FIGURE 1M

Ampullaria turbinis Lea, 1856:110.

Current taxonomic status: Synonym of Pila ampullacea (Linnaeus, 1758) 

(Habe, 1964:49 [as “ampullaria”], Brandt, 1974:49; Ng et al., 2014:32; 

Cowie, 2015:50).
zischkai Blume and Pain, 1952

Pomacea zischkai Blume and Pain, 1952:267, pl. 7.

Current taxonomic status: Pomacea zischkai Blume and Pain, 1952; valid species (Cowie and Thiengo, 

2003:77).

Type Material: None.

Type Locality: “Chapara Region, at 400 m. Bolivia tropica.”

Remarks: The holotype, designated by Blume and Pain (1952:268), was reported by them as in Blume’s 

collection; it is now in the ZSM (Cowie and Thiengo, 2003:77). Blume and Pain (1952:268) also 

listed five paratypes, one in Pain’s collection (now in NMW) and four in the FMNH (Cowie and 

Thiengo, 2003:77). Blume and Pain (1952:268) noted that “the whole series examined” (they did 

not say “type” series) included “some 340 examples.” Many of these were curated or cited as paratypes and noted as present in the ZSM and other museums by Cowie and Thiengo (2003:77) and as 

possible paratypes by Cowie (2015:65). However, all this additional material (presumably including 

those in USNM 667036, 2 specimens) was mentioned separately by Blume and Pain (1952:268) 

from their explicit designation of the holotype and five paratypes in their table, thereby expressly 

excluding them from the type series (Code, Article 72.4.6).

Material Previously Considered as Types

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