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Type Specimens of Hawaiian Land Snails in the Smithsonian Institution, National Museum of Natural History, with Lectotype Designations

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Pacific island land snail faunas are among the most threatened faunas in the world, having suffered a higher rate of extinction than any other major animal group. The Hawaiian land snails are among the most species-rich and most severely effected of these faunas, yet the current status of most of the Hawaiian species is unknown. Most of the major taxonomic studies on the fauna were undertaken 50–100 years ago and only certain groups were comprehensively studied. New research is uncovering undescribed species, both extant and extinct. The need for rigorous taxonomic treatment of the group is acute if the taxonomic and conservation status of the many species is to be ascertained, and the basis for such research is comprehensive study of type material. The Smithsonian's National Museum of Natural History holds type material of 39 nominal species-group taxa of Hawaiian land snails belonging to eight families; this annotated catalog provides details of this material. Of these taxa, 29 were described by Augustus Addison Gould from material collected by the U.S. Exploring Expedition of 1838–1842. For completeness, we also provide details for one lot of purported paralectotypes that are here inferred not to have been syntypes and one lot representing an unavailable infrasubspecific name. We designate lectotypes for 12 species-group taxa. Yeung, Norine W., Robert H. Cowie, Kenneth A. Hayes, and Ellen E. Strong. Type Specimens of Hawaiian Land Snails in the Smithsonian Institution, National Museum of Natural History, with Lectotype Designations. Smithsonian Contributions to Zoology, number 647, vi + 34 pages, 11 figures, 2017.
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A Chronology of
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By Craig M. Johnson
with contributions by
Stanley A. Ahler, Herbert Haas, and Georges Bonani
Smithsonian Institution
Scholarly Press
Smithsonian Institution
Scholarly Press
smithsonian contributions to zoology • number 647
Type Specimens of Hawaiian
Land Snails in the Smithsonian
Institution, National Museum
of Natural History, with
Lectotype Designations
Norine W. Yeung, Robert H. Cowie,
Kenneth A. Hayes, and
Ellen E. Strong
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smithsonian contributions to zoology • numbe r 6 4 7
Type Specimens of Hawaiian
Land Snails in the Smithsonian
Institution, National Museum
of Natural History, with
Lectotype Designations
Norine W. Yeung, Robert H. Cowie,
Kenneth A. Hayes, and
Ellen E. Strong
WASHINGTON D.C.
2017
ABSTRACT
Yeung, Norine W., Robert H. Cowie, Kenneth A. Hayes, and Ellen E. Strong. Type Specimens of Hawaiian
Land Snails in the Smithsonian Institution, National Museum of Natural History, with Lectotype Designa-
tions. Smithsonian Contributions to Zoology, number 647, vi + 34 pages, 11 figures, 2017. — Pacific island
land snail faunas are among the most threatened faunas in the world, having suffered a higher rate of extinc-
tion than any other major animal group. The Hawaiian land snails are among the most species-rich and most
severely affected of these faunas, yet the current status of most of the Hawaiian species is unknown. Most of
the major taxonomic studies on the fauna were undertaken 50–100 years ago and only certain groups were
comprehensively studied. New research is uncovering undescribed species, both extant and extinct. The need
for rigorous taxonomic treatment of the group is acute if the taxonomic and conservation status of the many
species is to be ascertained, and the basis for such research is comprehensive study of type material. The
Smithsonian’s National Museum of Natural History holds type material of 39 nominal species-group taxa
of Hawaiian land snails belonging to eight families; this annotated catalog provides details of this material.
Of these taxa, 29 were described by Augustus Addison Gould from material collected by the U.S. Exploring
Expedition of 1838–1842. For completeness, we also provide details for one lot of purported paralectotypes
that are here inferred not to have been syntypes and one lot representing an unavailable infrasubspecific
name. We designate lectotypes for 12 species-group taxa.
Cover images (left to right): Left of main Hawaiian Islands, top row (dimensions = shell height): Newcombia
cinnamomea honomuniensis (17.6 mm), Pupa peponum (2.7 mm), Achatinella radiata (17.8 mm), Amastra
obesa aurora (10.9 mm), Laminella concinna (8.0 mm); bottom row (dimensions = shell height [first three
shells] and shell width [last two shells]): Achatinella nubilosa (22.1 mm), A. nucleola (11.4 mm), Pupa lyrata
(2.3 mm), Helix rubiginosa (5.6 mm), Amastra rex (13.7 mm). Right of main Hawaiian Islands, top row (di-
mensions = shell width): Helix cryptoportica (5.6 mm), H. exaequata (10.9 mm), Vitrina caperata (11.2 mm),
Helicina uberta (4.8 mm); bottom row (dimensions = shell height): Succinea canella (9.7 mm), S. explanata
(9.7 mm), S. lumbalis (11.8 mm).
Published by SMITHSONIAN INSTITUTION SCHOLARLY PRESS
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Library of Congress Cataloging-in-Publication Data
Names: National Museum of Natural History (U.S.) | Yeung, Norine W., author. | National Museum of Natural History
(U.S.), issuing body.
Title: Type specimens of Hawaiian land snails in the Smithsonian Institution, National Museum of Natural History, with
lectotype designations / Norine W. Yeung [and three others].
Other titles: Hawaiian land snails in the Smithsonian Institution, National Museum of Natural History, with lectotype
designations | Smithsonian contributions to zoology ; no. 647.
Description: Washington, D.C. : Smithsonian Institution Scholarly Press, 2017. | Series: Smithsonian contributions to
zoology, ISSN 0081-0282 ; number 647 | Includes bibliographical references.
Identifiers: LCCN 2016045907
Subjects: LCSH: Snails—Hawaii—Type specimens—Catalogs and collections—Washington (D.C.) | National Museum of
Natural History (U.S.)—Catalogs. | Endangered species—Hawaii.
Classification: LCC QL430.4 .N38 2017 | DDC 594/.3074753—dc23 | SUDOC SI 1.27:647
LC record available at https://lccn.loc.gov/2016045907
ISSN: 1943-6696 (online); 0081-0282 (print)
ZooBank registration: 15 March 2017 (LSID: urn:lsid:zoobank.org:pub:9AADB713-5454-4DEE-A0DB-
9A558A6FAC99)
Publication date (online): 29 March 2017
Ó The paper used in this publication meets the minimum requirements of the American National Standard for Perma-
nence of Paper for Printed Library Materials Z39.48–1992.
Contents
LIST OF FIGURES v
INTRODUCTION 1
Approach and Format of Accounts 2
The Species of Augustus Addison Gould 2
Lectotype Fixation and Designation 3
Acronyms and Abbreviations 4
SYSTEMATIC CATALOG 5
Family Achatinellidae 5
honomuniensis Pilsbry and Cooke, 1912; Newcombia cinnamomea 5
marmorata Gould, 1847; Achatinella 5
peponum Gould, 1847; Pupa 7
radiata Gould, 1845; Achatinella 7
Family Amastridae 7
abberans Hyatt and Pilsbry, 1911; Amastra affinis bigener 7
acuminata Gould, 1847; Achatinella 9
aurora Pilsbry and Cooke, 1914; Amastra obesa 9
auwahiensis Pilsbry and Cooke, 1914; Amastra subsoror 9
cerealis Gould, 1847; Achatinella 9
circumcincta Hyatt and Pilsbry, 1911; Laminella concinna 10
ellipsoidea Gould, 1847; Achatinella 10
evelynae Cooke and Kondo, 1952; Carelia 10
guttula Gould, 1847; Achatinella 12
microstoma Gould, 1845; Achatinella 12
nubilosa Mighels, 1845; Achatinella 12
nucleola Gould, 1845; Achatinella 12
rex Sykes, 1904; Amastra 13
rubens Gould, 1845; Achatinella 13
rubinia Hyatt and Pilsbry, 1911; Amastra 16
Family Endodontidae 16
rubiginosa Gould, 1846; Helix 16
setigera Gould, 1844; Helix 16
Family Helicarionidae 17
cicercula Gould, 1846; Helix 17
cryptoportica Gould, 1846; Helix 19
iv
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
exaequata Gould, 1846; Helix 19
subtilissima Gould, 1846; Helix 21
Family Helicinidae 21
uberta Gould, 1847; Helicina 21
Family Pupillidae 21
lyrata Gould, 1843; Pupa 21
Family Succineidae 23
aperta Lea, 1838; Succinea 23
canella Gould, 1846; Succinea 23
cepulla Gould, 1846; Succinea 23
explanata Gould, 1852; Succinea 24
lumbalis Gould, 1846; Succinea 24
oregonensis Lea, 1841; Succinea 24
rotundata Gould, 1846; Succinea 26
venusta Gould, 1846; Succinea 26
vesicalis Gould, 1846; Succinea 28
Family Zonitidae 28
caperata Gould, 1846; Vitrina 28
pauxilla Gould, 1852; Helix 28
pusilla Gould, 1846; Helix 28
tenella Gould, 1846; Vitrina 30
ACKNOWLEDGMENTS 31
REFERENCES 33
1. Newcombia cinnamomea honomuniensis; Achatinella marmorata;
Pupa peponum 6
2. Achatinella radiata; Amastra affinis bigener var. abberans; Achatinella
acuminata; Amastra obesa aurora; Amastra subsoror auwahiensis;
Achatinella cerealis 8
3. Laminella concinna; L. concinna color-var. circumcincta; Achatinella
ellipsoidea; Carelia evelynae; A. guttula; A. microstoma; A. nubilosa;
A. nucleola 11
4. Amastra rex 14
5. Achatinella rubens; Amastra rubens var. rubinia; Helix rubiginosa 15
6. Helix setigera 18
7. Helix cicercula; H. cryptoportica; H. exaequata 20
8. Helix subtilissima; Helicina uberta; Pupa lyrata; Succinea aperta;
S. canella 22
9. Succinea cepulla; S. explanata; S. lumbalis 25
10. Succinea oregonensis; S. rotundata; S. venusta; S. vesicalis 27
11. Vitrina caperata; Helix pusilla; V. tenella 29
Figures
INTRODUCTION
Habitat destruction and the impacts of invasive species are the primary causes of
biodiversity loss and species extinction across many taxa, particularly on Pacific islands
(Cox and Elmqvist, 2000; Lydeard et al., 2004; Duncan et al., 2013). The spectacularly
diverse assemblages of land snails on these islands have been particularly heavily af-
fected, with many species already extinct and the remaining fauna disappearing rapidly
(Lydeard et al., 2004; Régnier et al., 2009, 2015; Richling and Bouchet, 2013; Sartori et
al., 2014). Among the Pacific islands, the most species-rich land snail fauna is that of the
Hawaiian Islands, with more than 750 described species, over 99% of them endemic to
the archipelago and many to single islands (Cowie et al., 1995). It has been suggested that
up to 90% of these species may already be extinct (Lydeard et al., 2004).
The current biodiversity crisis, exemplified by this fauna, emphasizes the urgent
need for taxonomic research to describe species before they vanish unknown (Solem,
1990; Hopkins and Freckleton, 2002; Rodman and Cody, 2003; Wheeler, 2004; Hawk-
sworth and Cowie, 2013). The major taxonomic research on Hawaiian land snails was
undertaken more than 50 years ago (e.g. Neal, 1934; Baker, 1940; Cooke and Kondo,
1960) and in some cases a century ago (e.g., Hyatt and Pilsbry, [1910]–1911; Pilsbry
and Cooke, 1912–1914). It is therefore difficult to assess the number of species still ex-
tant, especially as some groups have yet to be studied in detail (e.g., Endodontidae and
Punctidae; Solem, 1976, 1983), and because modern molecular and microscopy tech-
niques (e.g., scanning electron microscopy) are discovering numerous undescribed and
sometimes cryptic species, both extinct and extant. This lack of taxonomic clarity and
the dearth of recent studies of the Hawaiian land snails hinder attempts to assess their
conservation status accurately.
To begin conserving any fauna, a comprehensive compilation of information about
type material must be developed to provide the framework for the necessary systemat-
ics assessments. Natural history museum collections play a vital role in the study of
biodiversity and its loss by providing an indispensable resource of historical and current
Type Specimens of Hawaiian Land Snails
in the Smithsonian Institution, National
Museum of Natural History, with
Lectotype Designations
Norine W. Yeung,1,2,3* Robert H. Cowie,2 Kenneth A. Hayes,2,3,4
and Ellen E. Strong3
1 Bishop Museum, 1525 Bernice Street, Hono-
lulu, Hawaii, 96817, USA.
2 Pacific Biosciences Research Center, University
of Hawaii, 3050 Maile Way, Gilmore 408, Ho-
nolulu, Hawaii, 96822, USA.
3 Smithsonian Institution, National Museum
of Natural History, PO Box 37012, MRC 163,
Washington, D.C., 20013, USA.
4 Department of Biology, Howard University, 415
College St. NW, Washington, D.C., 20059, USA.
* Correspondence: nyeung@hawaii.edu
Manuscript received 8 August 2014; accepted
10 July 2016.
2
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
biological records (Davis, 1996; Ponder et al., 2001; Suarez and
Tsutsui, 2004). Hawaiian land snail type materials have been de-
posited in several national and international malacological col-
lections including the United States National Museum (USNM)
collection at the National Museum of Natural History (NMNH),
Smithsonian Institution.
Although the USNM malacological collection was not for-
mally established as the Department of Mollusks until 1880,
the museum began acquiring molluscan material almost as soon
as the Smithsonian Institution was founded in 1846. The col-
lection now contains more than 900,000 lots (Sturm, 2006)
of which over 12,000 are primary type lots. The large size of
this collection is partly due to it being the official repository for
national governmental agencies and expeditions including the
United States Exploring Expedition (1838–1842). The Mollusca
of the Exploring Expedition, which were collected or obtained
by J. P. Couthouy, were initially sent to the Peale museum in
Philadelphia and subsequently transferred to the National In-
stitution before being moved to the Smithsonian Institution in
1856 (Johnson, 1964). The Exploring Expedition Mollusca were
described primarily by Augustus Addison Gould, including 32
Hawaiian land snail species described between 1843 and 1862
(Cowie et al., 1995), and many of the specimens on which the
descriptions were based are housed within the USNM. Gould’s
material constitutes the greatest part of the Hawaiian land snail
type material in the USNM. The primary objective of this catalog
is to document this Hawaiian type material, as one in a series
of catalogs of museum types representing this highly threatened
fauna (e.g., Cowie et al., 2016).
ApproAch And FormAt oF Accounts
This catalog is a work of nomenclature and clarification of
the status of type material; it is not a work of taxonomy and
we have avoided making any new taxonomic judgments. All in-
terpretations follow the International Code of Zoological No-
menclature (ICZN, 1999), hereafter, the Code. Primary types
(i.e., holotype, syntype, lectotype; there are no neotypes) and
secondary types (i.e., paratype, paralectotype) are included in
this catalog.
The list is arranged alphabetically by family. Within each
family, taxa are arranged alphabetically by species-group name.
The heading of each entry consists of the name, author(s) and
date of description, followed by the genus of the original com-
bination, and the species for infraspecific taxa. The next line of
the entry then consists of the name as given with the original
genus (and species for subspecies, varieties, etc.) in which it was
described, verbatim as published by the author, including subge-
nus if in the original description, using the original orthography,
even if now considered incorrect according to the Code (e.g., dia-
critical marks, ligatures, incorrect gender ending, species name
beginning with a capital), except that genus and species names
are in italic even if printed otherwise in the original publication,
and with the original status indicated (e.g., subspecies, “var.”) as
necessary, with upper/lower case and italic/plain font as in the
original description. The name is followed by its author(s), date
of publication, page number, and plate/figure number(s). Sub-
sequent literature by the same author(s) bearing directly on the
original description follows immediately after the bibliographic
information, separated by a semicolon. Next the current taxo-
nomic status is given, including generic and subgeneric placement,
whether a valid taxon, and if not, the appropriate synonymy,
with one or more citations supporting the status. Current status
is taken to be that given by Cowie et al. (1995), with the excep-
tion of one taxon (rubinia Hyatt and Pilsbry, 1911; Amastridae)
described as a variety and treated as “infraspecific” by Cowie et
al. (1995) that is here cataloged as a subspecies in the light of the
Code (Article [Art.] 45.6.4). This is followed by a listing of type
material with USNM catalog number(s) and the number of speci-
mens in each lot; all specimens are dry shells. The type locality
follows within quotation marks, with the original orthography
as provided in the original description, or as clarified by refer-
ence to other sources (e.g., original labels, localities within the
known range of the taxon; Code, Recommendation [Rec.] 76A),
or as restricted by the designation of a lectotype. Additional type
locality information (e.g., clarifications, corrections, information
from subsequent publications) is given in square brackets. In-
formation on type material at other institutions (not necessarily
comprehensive), corrections or additional information, changes
in type status, information on lost or destroyed specimens, and
so on, is included in a remarks section. In these remarks, species-
group taxa are generally referred to in the generic combination
of their original description.
the species oF Augustus Addison gould
Some confusion, or at least ambiguity, has arisen in the lit-
erature regarding certain original numbers such as “A1197” (see
also Johnson, 1964) associated with some of Gould’s specimen
lots (e.g., Achatinella radiata Gould). These appear to be catalog
numbers originally given by Gould to lots containing specimens
he described. These lots were probably loaned to Gould in Bos-
ton by the National Institution (see above), and on their return
to the newly formed Smithsonian Institution in Washington
these numbers were entered into the catalog ledgers some time
after 1860 by P. P. Carpenter (Carpenter, 1864:530; Johnson,
1964:15). Some of these lots, with labels with Gould’s original
numbers, were distributed as duplicates to various museums, no-
tably the New York State Museum, the type material of which is
now on permanent loan to the Museum of Comparative Zool-
ogy of Harvard University (Johnson, 1964), although some of
this material appears to have been lost during the transfer among
museum collections (Hall, 1875).
The vast majority of Gould’s U.S. Exploring Expedition
type material of Hawaiian land snails in the USNM is cataloged
under two sets of numbers, one in the USNM 5000 series and
one in the USNM 20000 series. In the original handwritten
USNM catalog ledger, many 20000 series lots are accompanied
NUMBER 647
3
by yet another “original number” preceded by the letter “f”
(e.g., “f20”; Succinea canella Gould); the corresponding lots
in the 5000 series have the same “original number” but with
no prefix (e.g., “20”). Unlike the original catalog numbers,
these “original numbers” correspond to the figure numbers of
Gould’s illustrations published in 1856, with the “f” appearing
to indicate “figure.” In most cases, the 20000 series lots contain
multiple specimens and appear to have been separated from the
original lots cataloged in the 5000 series, which now often con-
tain only a single specimen that matches Gould’s (1856) figure.
It is generally the latter specimen that is selected as the lectotype,
when appropriate.
Most lots in the 20000 series are noted in the original hand-
written USNM catalog ledger as having been received from the
U.S. Exploring Expedition, but a number are either noted as also
having been received from William Harper Pease or as having
been received only from Pease. Born in New York in 1824, Pease
moved to Hawaii in December 1849 (Kay and Clench, 1975:2–
3), after most of Gould’s species had been described (Gould,
1844, 1845, 1846, 1847). So it seems most likely that Pease
would have received the material on loan or on exchange from
Gould after he had arrived in Hawaii and become interested in
Hawaiian land snails, returning it subsequently, perhaps directly
to the USNM. There, along with the material returned by Gould
from Boston, it was cataloged by Carpenter, who, perhaps inad-
vertently, noted a few lots as having been received from Pease
only, without noting their original receipt from the U.S. Explor-
ing Expedition. Many of the 20000 series lots are annotated with
original numbers (as explained above). Carpenter (1864:530)
explained that “a considerable part of the shells professing to
be the figured types of the new species were found together, with
the artist’s marks corresponding with the plates and figures.” Ac-
cordingly, we consider 20000 series lots annotated with Gould’s
original catalog and/or figure numbers to be type material. In
several cases, lots with no original numbers are considered to be
possible type material, as explained below.
lectotype FixAtion And designAtion
Additional confusion, regarding possible lectotype
designation/ fixation, has arisen as a result, in particular, of the
catalog of Gould’s material by Johnson (1964). In a number
of cases Johnson used the term “lectotype” clearly and validly,
thereby designating the particular specimen as such. However,
his use of the terms “holotype,” “figured holotype,” and “fig-
ured neoholotype” may not be valid lectotype designations/
fixations, based on the relevant articles of the Code (Art. 74.5,
74.6), which are difficult to interpret. We have interpreted these
two articles of the Code as follows in the context of this paper.
A holotype can be fixed only in the original publication
when the nominal taxon is established (Code, Art. 73.1.3, Glos-
sary). There are two ways in which an incorrect use of “holo-
type” can be a lectotype fixation: (1) under Art. 74.5, when the
original description reveals that the taxon was based on more
than one specimen and a subsequent author made an explicit,
intentional statement of selection; or (2) under Art. 74.6, when
the original description does not imply that the taxon was based
on more than one specimen, and when an author published be-
fore the year 2000 an inference that a syntype is the “holotype”
or “the type”; if it is discovered that there was more than one
syntype, this assumption becomes a lectotype designation, but
only if the author had assumed that the original description was
based on only one specimen.
An author writing “holotype” or “figured holotype” when
no holotype was originally designated and when he or she knows
that the type series had more than one specimen is in error: no
specimen has been “unambiguously selected” under Art. 74.5,
because in fact no selection has occurred, as the author consid-
ers the holotype to have been fixed by the original author. For
example, Johnson (1964:148), in his treatment of Helix setigera,
explicitly noted the “figured neoholotype, selected by Gould,
USNM 5453”; this is not a lectotype designation under Art.
74.5. Furthermore, such usages cannot be lectotype designa-
tions under Art. 74.5 because Johnson (1964) did not misuse
the term “holotype”—that is, intending it as a novel selection of
the name-bearing type—as he also used “lectotype” in the same
paper, clearly understanding the distinction.
Thus when Johnson used the terms “figured holotype” or
“holotype,” such usages cannot be lectotype designations under
Art. 74.5, but potentially can be lectotype fixations under Art.
74.6, although only if he accepted that the taxon was based on
a single type specimen. In no case did Johnson explicitly do so
when he used these terms and in most cases he also listed other
type (“paratype”) material. Therefore, these are not lectotype
designations under Art. 74.6 and thus lectotypes were neither
designated under Art. 74.5 nor fixed under Art. 74.6.
Johnson (1994) clarified his former usage of the terms “fig-
ured holotype” or “holotype,” stating that if he could “locate
the single figured or measured syntype, it was usually regarded
as the holotype.” But he also acknowledged that this practice
was no longer tenable under the 1985 Code (ICZN, 1985), and
neither is it under the current Code (ICZN, 1999). Whether a
specimen was part of the type series and therefore eligible for
lectotype designation may be judged using external information
(Code Art. 72.4.1.1), but whether the wording of a putative des-
ignation qualifies as a lectotype designation must be judged on
the basis of information contained in the publication; one can-
not retroactively change the status of such statements. However,
Johnson (1964), in multiple places as cited herein, wrote “fig-
ured holotype” and also listed paratypes. This shows that he did
not mean by “figured holotype” that the type series had only a
single specimen, and therefore he did not designate lectotypes
under Art. 74.6.
It is also necessary to clarify our interpretation of pos-
sible lectotype designations by Baker (1963). He often used
“TOM,” which means “type because only one example was
included in the original description, or was indicated by only
one set of dimensions (of course the first) or by reference to a
4
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
(cited) illustration(s) of only one shell, in the definition proper,
exclusive of additional remarks” (Baker, 1963:191). Therefore
TOM does not mean that Baker necessarily accepted that there
was a single specimen. However, Baker went a step further than
Johnson in stating that any use of his abbreviations for type
designations was a “TSD,” which means “type by subsequent
selection, followed by ‘now’ if apparently first designated in
these lists and/or preceded by name and reference, especially
when selected previously. Of course, every usage of any of these
abbreviations is a TSD in this list” (Baker, 1963:191). A “TSD”
is broader than the current concept of a new lectotype designa-
tion, but in cases in which the “type” was “first designated
in these lists” we treat “TSD” as a valid lectotype designation
under Art. 74.5.
Additional details and explanations of our conclusions re-
garding the status of specimens as lectotypes are provided under
the individual taxon entries, including cases involving other au-
thors (Hyatt and Pilsbry, [1910]–1911; Pilsbry and Cooke, 1908,
1914–1916, 1918–1920; Baker, 1940, 1941).
In addition to clarifying possible lectotype designations of
previous authors, we make a number of designations herein.
This paper is part of an ongoing effort to update the systemat-
ics of the Hawaiian land snails, and appropriate designation of
lectotypes is part of this overarching program of research (see
Rec. 74G; ICZN, 2003). However, we have been circumspect
and have not designated lectotypes in situations in which it is
possible that more appropriate specimens may be present in
other collections.
Acronyms And AbbreviAtions
alt. altitude
ANSP Academy of Natural Sciences of Drexel
University, Philadelphia
Art. Article of the Code
BPBM Bernice Pauahi Bishop Museum, Honolulu
BSNH Boston Society of Natural History, Boston
Code International Code of Zoological Nomenclature
(ICZN, 1999)
lat. latitude
long. longitude
MCZ Museum of Comparative Zoology, Harvard
University, Cambridge, Mass.
n. comb. new combination
NHMUK the Natural History Museum, London, UK
(former acronym BMNH)
NMNH National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
no. number
n. var. new variety
NYSM New York State Museum, Albany
Rec. Recommendation of the Code
spm(s) specimen(s)
subg. subgenus
USNM U.S. National Museum (collections now held in
the Smithsonian’s NMNH)
var. variety
Family achatinellidae
honomuniensis Pilsbry and Cooke, 1912;
Newcombia cinnamomea
Newcombia cinnamomea var. honomuniensis Pilsbry and Cooke, 1912:12, pl. 14, figs. 6, 7.
Current taxonomic status: Newcombia pfeifferi honomuniensis Pilsbry and Cooke, 1912.
Valid subspecies (Pilsbry and Cooke, 1914:355; Cowie et al., 1995:62).
Type material: Paralectotypes USNM 673317 (8 spms; Figure 1A).
Type locality: “Honomuni” [which is on Molokai (Pilsbry and Cooke, 1912:11)].
Remarks: No holotype was designated but figures of two specimens were provided with
the original description, captioned as “cotypes.” According to the original descrip-
tion, the type material was collected by D. Thaanum (Pilsbry and Cooke, 1912:12).
However, the original handwritten USNM ledger indicates that USNM 673317 was
received from Kuhns and Thaanum and the specimens were “paratypes.” The ledger
confirms the locality as Honomuni, Molokai, Hawaii. Baker (1963:194) validly (as
“TOM”) designated ANSP 110071a as the lectotype (see section in the Introduc-
tion regarding lectotype designations). An additional paralectotype, BPBM 36858 (1
spm), was collected and donated by D. Thaanum. The original species combination
was listed incorrectly by Cowie et al. (1995:62) as Newcombia pfeifferi var. hono-
muniensis, which is the current placement of this taxon.
marmorata Gould, 1847; Achatinella
Achatinella marmorata Gould, 1847:200; 1852:85; 1856: pl. 7, figs. 94, 94a; 1862:34.
Current taxonomic status: Partulina (Partulina) marmorata (Gould, 1847). Valid species
(Pilsbry and Cooke, 1912:42; Cowie et al., 1995:70).
Type material: Lectotype USNM 5496 (Figure 1B), here designated; paralectotype USNM
1418213 (1 spm, Figure 1C; ex USNM 5496).
Type locality: “Haleakala Mountains, Maui, Sandwich Islands” [=Hawaiian Islands].
Remarks: No holotype was designated nor a figure provided with the original descrip-
tion, although figures were provided subsequently by Gould (1856: pl. 7, figs. 94,
94a). Johnson (1964:109) noted “2 syntypes USNM 5496 slightly smaller than mea-
sured type.” There are no remarks in the original handwritten USNM ledger or on
the printed labels identifying either of the specimens as figured. Neither the color
Systematic Catalog
6
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 1. A. Paralectotypes (USNM 673317; 17.6 mm) of Newcombia cinnamomea honomuniensis. B.
Lectotype (USNM 5496; 14.7 mm), here designated, of Achatinella marmorata. C. Paralectotype (USNM
1418213; 16.0 mm) of A. marmorata. D. Gould’s (1856: pl. 7, figs. 94, 94a) figured Achatinella marmorata. E.
Lectotype (USNM 5506; 2.5 mm) of Pupa peponum. F. Gould’s (1856: pl. 7, fig. 104) figured Pupa peponum.
G. Paralectotype (USNM 5506a; 2.7 mm) of P. peponum. H. Gould’s (1856: pl. 7, fig. 104a) figured Pupa
peponum. Dimensions are shell height (length). If more than one shell in the lot is figured, dimension is of the
shell indicated by an asterisk. Scale bars: A–C, 5 mm; E and G, 1 mm.
NUMBER 647
7
patterning, which is highly stylized, nor the shell shape in
Gould’s (1856: figs. 94, 94a) illustrations (Figure 1D) is a
perfect match for either of the shells and it is possible that
they are composites incorporating elements from both. We
here designate USNM 5496 as the lectotype (Figure 1B).
The type material was collected by J. D. Brackenridge and
J. Drayton (Gould, 1852:86) and the original handwritten
ledger indicates that it was received from the U.S. Exploring
Expedition, with “original number” 94, corresponding to
Gould’s (1856) illustration.
peponum Gould, 1847; Pupa
Pupa peponum Gould, 1847:197; 1852:93; 1856: pl. 7, figs. 104, 104a–e;
1862:34, 244.
Current taxonomic status: Lamellidea (Lamellidea) peponum
(Gould, 1847). Valid species (Pilsbry and Cooke, 1914:156;
Cowie et al., 1995:80).
Type material: Lectotype (Pilsbry and Cooke, 1914:157, pl. 35,
figs. 1, 2) USNM 5506 (Figure 1E); paralectotype USNM
5506a (Figure 1G).
Type locality: Hilo or Oahu (Gould, 1852:93).
Remarks: No holotype was designated nor a figure provided
with the original description. In the original description,
Gould (1847:197) stated the type locality as “Sandwich
Islands.” Gould (1847:197) described the species as hav-
ing “very variable characters” and subsequently (Gould,
1852:94; 1856: pl. 7, figs. 104, 104a–e) provided figures of
multiple specimens, stating that he had specimens collected
in Hilo, Hawaii, by C. Wilkes and on Oahu by J. D. Brack-
enridge. Sykes (1903:382) determined that Gould’s illustra-
tions of P. peponum represented three species, identifying
the shells (note the plural) in Gould’s figs. 104 and 104d as
Pupa peponum. Pilsbry and Cooke (1914:157), aware of
the multiple syntypes, noted “the type specimen, no. 5506
Smithsonian Institution” and indicated that this was the
specimen figured by Gould (1856: pl. 7, fig. 104), thereby
selecting it as the lectotype of Pupa peponum (and avoiding
the problematic use of the term “holotype,” as discussed
in the section on lectotypes in the Introduction; Code, Art.
74.5). Johnson (1964:125) noted this lot as the “holotype.”
The original handwritten ledgers indicate that there were
originally two specimens in this lot, received from the U.S.
Exploring Expedition, with “original number” 104. Gould
(1856: pl. 7, figs. 104, 104a) figured two entire shells. Pils-
bry and Cooke (1914: pl. 35, figs. 1, 2) considered their fig-
ured shell (USNM 5506) to be that of Gould’s fig. 104, but
the other shell in USNM 5506 is not a definitive match for
Gould’s fig. 104a (see Figures 1F, 1H, respectively). A note
with the lot, written by Y. Kondo in 1956, indicates that the
smaller specimen is probably an immature specimen of Paci-
ficella variabilis Odhner, 1922 (as Tornatellinops), though
this species is not known from the Hawaiian Islands. This
smaller specimen was subsequently removed from the lot
and designated as USNM 5506a (Figure 1G). USNM 5506
is listed in the ledger simply as originating from the “Sand-
wich Islands,” but Gould (1852:94) stated that he had ma-
terial from Hilo (island of Hawaii) and Oahu, indicating
that the type locality could be on either of these islands.
Johnson (1964:125) noted an additional “paratype” (MCZ
216798, ex Peabody Museum, Salem, Massachusetts, prob-
ably collected by J. P. Couthouy). However, this specimen
is in fact a species of Eulimidae, which are marine. It is not
clear why Johnson considered it a paratype of P. peponum
and we do not consider it type material of this species.
radiata Gould, 1845; Achatinella
Achatinella radiata Gould, 1845:27; 1862:195.
Current taxonomic status: Partulina (Partulina) radiata (Gould,
1845). Valid species (Pilsbry and Cooke, 1912:49; Cowie et
al., 1995:72).
Type material: Lectotype USNM 712806 (Figure 2A), here desig-
nated; paralectotypes USNM 1418215 (3 spms, Figure 2B;
ex USNM 712806).
Type locality: “Sandwich Islands” [=Hawaiian Islands].
Remarks: No holotype was designated nor a figure provided
with the original description. The “Type lot” (Johnson,
1964:138) appears to have been one of a number of lots
of multiple taxa intended for distribution to the New York
State Museum (NYSM) in 1863 (Hall, 1875:13; Johnson,
1964:15). Under the number A1197, this type lot was re-
ported as “not to be found in the collection” in the NYSM
annual report for 1873 (Hall, 1875:13) and was treated as
“lost” by Johnson (1964:138). It has now been found in the
USNM type collection as USNM 712806 (originally 4 spms)
and the USNM handwritten ledger confirms the original
number as A1197 (“type series 1845. PBSNH. 2:27” [i.e.,
Gould, 1845:27]). We here designate USNM 712806 as the
lectotype, with the other three specimens of the original lot
becoming paralectotypes and given the new number USNM
1418215. Four additional paralectotypes are in the MCZ
(Johnson, 1996:195; MCZ 2982814, as “syntypes”).
Family amastridae
abberans Hyatt and Pilsbry, 1911;
Amastra affinis bigener
Amastra affinis bigener var. abberans Hyatt and Pilsbry, 1911:300, pl. 44,
fig. 8.
Current taxonomic status: None; unavailable name.
Type material: None; unavailable name.
Type locality: None; unavailable name.
Remarks: We include this name in this catalog for complete-
ness and to preempt potential confusion, as it was listed
8
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 2. A. Lectotype (USNM 712806; 17.8 mm), here designated, of Achatinella radiata. B. Para-
lectotypes (USNM 1418215; ex 712806; 18.0 mm) of A. radiata. C. Non-type material (unavailable
name; USNM 117264; 14.2 mm) of Amastra affinis bigener var. abberans. D. Hyatt and Pilsbry’s (1911:
pl. 44, fig. 8) figured Amastra affinis bigener var. abberans. E. Lectotype (USNM 5502; 10.5 mm), here
designated, of Achatinella acuminata. F. Gould’s (1856: pl. 7, fig. 100) figured Achatinella acuminata. G.
Paralectotypes (USNM 308750; 10.9 mm) of Amastra obesa aurora. H. Paralectotypes (USNM 308754;
18.9 mm) of Amastra subsoror auwahiensis. I. Syntype (USNM 5501; 9.2 mm) of Achatinella cerealis.
J. Gould’s (1856: pl. 7, fig. 99) figured Achatinella cerealis. Dimensions are shell height (length). If more
than one shell in the lot is figured, dimension is of the shell indicated by an asterisk. Scale bars: 5 mm.
NUMBER 647
9
as available by Cowie et al. (1995:90). The name was in-
troduced under the heading “A. a. bigener Hyatt, n. var.”
(i.e., infrasubspecific), although in the body of the text the
Hyatt manuscript name “A. bigener var. abberans” was
mentioned, leading Cowie et al. (1995:90) to treat it as
available, although invalid as a synonym of Achatinella
affinis Newcomb, 1854 (now placed in Amastra). How-
ever, it is clear that the name was used to denote an infra-
subspecific entity, as Hyatt and Pilsbry (1911:383) in the
plate legend listed it as “Amastra a. bigener,” and to our
knowledge it was neither adopted prior to 1985 as the
valid name for a species or subspecies nor treated as a se-
nior homonym. Therefore, the name is unavailable (Code,
Art. 10.2, 45.6).
Lot USNM 117264 represents the material on which
the name was based, at least in part. According to the origi-
nal USNM ledger, eight specimens were present when the
lot was accessioned from the Lea collection, but only two
specimens remain. Although the exact collection locality is
unknown, the original description was placed in the section
of the publication that described Amastra spp. from Maui
(Hyatt and Pilsbry, 1911). The ledger indicates the locality
simply as the “Sandwich Islands.” One of the two shells in
USNM 117264 (Figure 2C) is the specimen figured (Figure
2D) by Hyatt and Pilsbry (1911: pl. 44, fig. 8).
acuminata Gould, 1847; Achatinella
Achatinella acuminata Gould, 1847:200; 1852:88; 1856: pl. 7, figs. 100,
100a.
Current taxonomic status: Leptachatina (Leptachatina) acumi-
nata (Gould, 1847). Valid species (Cooke in Hyatt and Pils-
bry, 1910:6; Cowie et al., 1995:120).
Type material: Lectotype USNM 5502 (Figure 2E), here des-
ignated.
Type locality: “Kauai, Sandwich Islands.”
Remarks: No holotype was designated nor a figure provided
with the original description, although multiple figures of a
single shell were provided subsequently by Gould (1856: pl.
7, figs. 100, 100a) and the colored figure (fig. 100) is here re-
produced as Figure 2F. Cooke (in Hyatt and Pilsbry, 1910:6)
noted the presence of a single broken shell in the Smithson-
ian collections, and failed to find additional material despite
a careful search in the Gould collection of the New York
State Museum. The original handwritten USNM ledger con-
firms that the specimen was already broken when it was
accessioned; the ledger does not have any additional infor-
mation regarding type status. It gives the locality simply as
the “Sandwich Islands,” although the original description
restricted it to Kauai, and indicates that the specimen was
received from the U.S. Exploring Expedition, with “origi-
nal number” 100. We here designate this single broken shell
(USNM 5502) as the lectotype.
aurora Pilsbry and Cooke, 1914;
Amastra obesa
Amastra obesa aurora Pilsbry and Cooke, 1914:18, pl. 4, figs. 9–12.
Current taxonomic status: Amastra (Cyclamastra) obesa aurora
Pilsbry and Cooke, 1914. Valid subspecies (Cowie et al.,
1995:100).
Type material: Paralectotypes USNM 308750 (3 spms, Figure 2G).
Type locality: “East Maui: Auwahi at about 4200 ft. elevation.”
Remarks: No holotype was designated, but figures of three speci-
mens were provided with the original description. Pilsbry
and Cooke (1914:18) reported “cotypes” in ANSP, BPBM,
and Thaanum’s personal collection. The original handwrit-
ten USNM ledger indicates the status of the specimens in
USNM 308750 as “cotypes” and that the collectors were
D. Thaanum and D. B. Kuhns, and that they were received
from A. Busck. August Busck was a microlepidopterist at
the NMNH, and apparently had a shell collection. Without
any information in the original description about collectors,
it seems reasonable to conclude that these were the speci-
mens noted by Pilsbry and Cooke as in Thaanum’s collec-
tion. This weight of evidence leads us to consider them as
paralectotypes. Baker (1963:197) validly (as “TSD now”)
designated ANSP 109838a as the lectotype (see section in
the Introduction regarding lectotype designations).
auwahiensis Pilsbry and Cooke, 1914;
Amastra subsoror
Amastra subsoror auwahiensis Pilsbry and Cooke, 1914:48, pl. 5, figs. 8–10.
Current taxonomic status: Amastra (Heteramastra) subsoror
auwahiensis Pilsbry and Cooke, 1914. Valid subspecies
(Cowie et al., 1995:103).
Type material: Paralectotypes USNM 308754 (3 spms, Figure 2H).
Type locality: “East Maui: Auwahi, at 4200 ft.”; “Auwahi is on
the slope of Haleakala facing Hawaii, just above Ulupal-
akua.”
Remarks: No holotype was designated but figures of three speci-
mens were provided with the original description. Pilsbry
and Cooke (1914:48) did not mention where type material
was deposited. The USNM ledger indicates the collectors of
USNM 308754 as D. Thaanum and D. B. Kuhns, that they
were received from A. Busck (see aurora Pilsbry and Cooke,
1914), and that the locality was “Auwahi, East Maui.”
Baker (1963:197) validly (as “TSD now”) designated ANSP
109836a as the lectotype (see section in the Introduction re-
garding lectotype designations). We consider the specimens
in USNM 308754 to be paralectotypes.
cerealis Gould, 1847; Achatinella
Achatinella cerealis Gould, 1847:201; 1852:90; 1856: pl. 7, figs. 99, 99a;
1862:35, 244.
10
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Current taxonomic status: Leptachatina (Leptachatina) cerealis
(Gould, 1847). Valid species (Cooke in Hyatt and Pilsbry,
1910:13; Cowie et al., 1995:121).
Type material: Possible syntype USNM 5501 (Figure 2I).
Type locality: “Waianai, Oahu” [sic, Waianae].
Remarks: No holotype was designated nor a figure provided with
the original description, although figures of a single specimen
were provided subsequently by Gould (1856: pl. 7, figs. 99,
99a). It has not been determined that the description was
based on a single specimen, and the original description does
not imply or require that it was based on more than one
specimen. The specimen in USNM 5501 is not a good match
for Gould’s (1856: pl. 7, fig. 99) figure and is probably not
the figured specimen (Figure 2J), suggesting that there were
in fact multiple syntypes, or that this specimen was not part
of the type series. By noting the “Figured holotype USNM
5501,” Johnson (1964:53) inferred a “holotype,” but this
statement is not a valid lectotype fixation (see section in the
Introduction regarding lectotype designations). Additionally,
because it probably is not the figured specimen and may not
have been part of the type series (perhaps a different species),
we refrain from designating it as the lectotype, treating it
as only a possible syntype. The original handwritten ledger
for USNM 5501 confirms Oahu as the locality and indicates
that one specimen was received from the U.S. Exploring Ex-
pedition, with “original number” 99.
circumcincta Hyatt and Pilsbry, 1911;
Laminella concinna
Laminella concinna Color-var. circumcincta Hyatt and Pilsbry, 1911:337, pl.
54, figs. 12, 13.
Current taxonomic status: Laminella concinna circumcincta
Hyatt and Pilsbry, 1911. Valid subspecies (Cowie et al.,
1995:114).
Type material: Lectotype USNM 31404 (Figure 3A), here des-
ignated.
Type locality: “Lanai.”
Remarks: Hyatt and Pilsbry (1911:337) noted “no. 31404 U. S.
Nat. Mus., from the Dall coll.” as a “typical example” of
a unique color pattern of Laminella concinna, having three
bands, whereas another specimen “in C. M. Cooke’s collec-
tion (no. 2201) lacks the broad median band.” Although
both specimens are figured, Hyatt and Pilsbry (1911:337)
did not designate a holotype. USNM 31404 appears to be
Hyatt and Pilsbry’s (1911: fig. 12) figured specimen (Figure
3B). We here designate USNM 31404 as the lectotype. The
original handwritten USNM ledger indicates that the type
material was received from W. H. Dall. Hyatt and Pilsbry
(1911:337) described this as a “Color-var.” and “form” but
did not expressly give it infrasubspecific rank, nor does the
content of the work reveal clearly that the name was pro-
posed for an infrasubspecific entity. It is subspecific accord-
ing to the Code (Art. 45.6.4).
ellipsoidea Gould, 1847; Achatinella
Achatinella ellipsoidea Gould, 1847:200; 1852:87; 1856: pl. 7, figs. 96, 96a;
1862:35.
Current taxonomic status: Synonym of Helix textilis Férussac,
1825 (now placed in Amastra subg. Metamastra) (Hyatt
and Pilsbry, 1911:167; Cowie et al., 1995:106).
Type material: Lectotype (Hyatt and Pilsbry, 1911:167) USNM
5498 (Figure 3C).
Type locality: Oahu [possibly restricted to Nuuanu (Hyatt and
Pilsbry, 1911:167)].
Remarks: No holotype was designated nor a figure provided with
the original description, although figures were provided sub-
sequently by Gould (1856: pl. 7, figs. 96, 96a). The origi-
nal description does not imply or require that it was based
on more than one specimen. Hyatt and Pilsbry (1911:167)
stated that “. . . we have examined Gould’s figured type, is
no. 5498 U.S. Nat. Mus.” and “the type is figured” (Hyatt
and Pilsbry, 1911: pl. 40, figs. 17, 18), thereby designating
a lectotype (Code, 74.5). Johnson (1964:72) subsequently,
while also noting the “holotype” as USNM 5498, identified
additional type material as MCZ 156364 (1 spm; ex Smith-
sonian Institution; paralectotype). Gould (1847:200) gave
the type locality as “Maui.” However, Hyatt and Pilsbry
(1911:167) concluded that USNM 5498 could not have been
collected from Maui, being a form of Amastra textilis “ex-
actly like some of the Nuuanu specimens.” Indeed, Gould
(1852:87) acknowledged the close affinity of Achatinella el-
lipsoidea to A. ventulus (=Amastra textilis). On this basis we
consider Oahu, as stated by Hyatt and Pilsbry (1911:167),
although not explicitly, to be the type locality. The type ma-
terial was collected by J. D. Brackenridge and Hale (Gould,
1852:87). USNM 5498 closely matches Gould’s (1856: pl. 7,
fig. 96) figured specimen (Figure 3D). The original handwrit-
ten ledger indicates the lectotype was received from the U.S.
Exploring Expedition, with “original number” 96.
evelynae Cooke and Kondo, 1952; Carelia
Carelia evelynae Cooke and Kondo, 1952:331, figs. 2a–f.
Current taxonomic status: Carelia evelynae Cooke and Kondo,
1952. Valid species (Cowie et al., 1995:111).
Type material: Paratypes USNM 666051 (2 spms, Figure 3E),
666052 (1 spm, Figure 3F).
Type locality: “Kauai, Polihale, base of Polihale Ridge: 500 ft.
inland from ocean, 150 ft. alt.”
Remarks: Holotype BPBM 9092, by original designation; para-
types BPBM 212325–212329 (115 spms), 212298 (137
spms) (Cooke and Kondo, 1952:333). The original hand-
written USNM ledger indicates that USNM 666051 and
666052 were separated from BPBM 212326 and 212298,
respectively. The ledger confirms the locality as “Kavai [sic]
Polihale Hawaii,” that the specimens were received from
NUMBER 647
11
FIGURE 3. A. Lectotype (USNM 31404; 8.0 mm), here designated, of Laminella concinna. B. Hyatt and
Pilsbry’s (1911: pl. 54, fig. 12) figured Laminella concinna color-var. circumcincta. C. Lectotype (USNM
5498; 15.9 mm) of Achatinella ellipsoidea. D. Gould’s (1856: pl. 7, figs. 96, 96a) figured Achatinella el-
lipsoidea. E. Paratypes (USNM 666051; 37.9 mm) of Carelia evelynae. F. Paratype (USNM 666052; 32.0
mm) of C. evelynae. G. Possible lectotype (USNM 5500; 7.3 mm) of Achatinella guttula. H. Gould’s (1856:
pl. 7, fig. 98) figured Achatinella guttula. I. Paralectotypes (USNM 611217; 13.2 mm) of Achatinella mi-
crostoma. J. Possible paralectotype (USNM 5497; 22.1 mm) of Achatinella nubilosa. K. Paralectotypes
(USNM 611221; 11.4 mm) of Achatinella nucleola. Dimensions are shell height (length). If more than one
shell in the lot is figured, dimension is of the shell indicated by an asterisk. Scale bars: 5 mm.
12
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Bishop Museum, collected by G. F. Arnemann, and their
status as paratypes.
guttula Gould, 1847; Achatinella
Achatinella guttula Gould, 1847:201; 1852:89; 1856: pl. 7, figs. 98, 98a;
1862:35, 244.
Current taxonomic status: Leptachatina (Leptachatina) guttula
(Gould, 1847). Valid species (Cooke in Hyatt and Pilsbry,
1910:36; Cowie et al., 1995:123).
Type material: Possible lectotype (Cooke in Hyatt and Pilsbry,
1910:37) USNM 5500 (Figure 3G).
Type locality: East Maui (Cooke in Hyatt and Pilsbry, 1910:36).
Remarks: No holotype was designated and no figure was provided
with the original description, although figures were provided
subsequently by Gould (1856: pl. 7, figs. 98, 98a). Gould
(1847:201) indicated “Maui, Sandwich Islands” as the type
locality. The original description includes the phrase “inter-
dum castaneo zonata,” which Gould (1852:89) rendered in
English as “sometimes [a faint appearance of] revolving ru-
fous bands,” indicating that there was more than one speci-
men in the type series. However, Cooke in Hyatt and Pilsbry
(1910:37, pl. 2, fig. 34) noted “the type specimen in the Na-
tional Museum, Washington” and reproduced “Gould’s fig-
ure of the type” (Gould, 1856: pl. 7, figs. 98, 98a), thereby
designating the shell illustrated as the lectotype (Code, Art.
74.5). Johnson (1964:87) subsequently identified the figured
“holotype” as USNM 5500 but also noted additional type
material in the MCZ, including MCZ 169182 (22 spms, ex
NYSM 195, original USNM no. A1175) and MCZ 142956
(2 spms, ex Smithsonian Institution). The original handwrit-
ten USNM ledger noted that the material was received from
the U.S. Exploring Expedition, with “original number” 98.
However, USNM 5500 does not closely match the illustra-
tions of Gould (Figure 3H) and of Cooke, and Johnson may
therefore have been incorrect in identifying it as the lectotype
designated by Cooke. We therefore remain uncertain as to
whether USNM 5500 is indeed the lectotype; however, no
other possible type material could be found.
microstoma Gould, 1845; Achatinella
Achatinella microstoma Gould, 1845:28; 1852:87; 1862:196.
Current taxonomic status: Synonym of Helix textilis Férussac, 1825
(now placed in Amastra subg. Metamastra) (Gould, 1862:196;
Hyatt and Pilsbry, 1911:165; Cowie et al., 1995:106).
Type material: Paralectotypes USNM 611217 (2 spms, Figure 3I).
Type locality: Oahu (Johnson, 1964:110, pl. 41, fig. 6).
Remarks: No holotype was designated nor a figure provided
with the original description. Gould (1845:28) indicated
the type locality as “Sandwich Islands.” Hyatt and Pilsbry
(1911:166) reported that “the type of A. microstoma has
been lost” without further elaboration; this is not a valid
lectotype designation, as a lectotype designation must have
as its object the definition of that taxon (Code, Art. 74.3).
Johnson (1964:110) validly (Code, Art. 74.5) designated
MCZ 169244 as the lectotype and identified MCZ 169245
(10 spms) and USNM 611217 (2 spms, in the series received
on permanent loan from the New York State Museum, via
W. J. Clench) as paralectotypes, all derived from the same
original lot (original number A1167 from “Oahu”).
nubilosa Mighels, 1845; Achatinella
Achatinella nubilosa Mighels, 1845:20.
Current taxonomic status: Amastra (Amastra) nubilosa (Mighels,
1845). Valid species (Hyatt and Pilsbry, 1911:259; Cowie et
al., 1995:94).
Type material: Possible paralectotype USNM 5497 (1 spm, Fig-
ure 3J).
Type locality: Molokai (Hyatt and Pilsbry, 1911:259).
Remarks: No holotype was designated nor a figure provided with
the original description. Mighels’ collection was sold to the
Portland Society of Natural History but was destroyed by
fire in 1854, although some material survived, having been
previously donated to the collections of the AMNH, MCZ,
and NHMUK (Johnson, 1949:214; Dance, 1966:294). John-
son (1949:227) identified two “poorly preserved cotypes” in
the MCZ derived, as previously stated by Hyatt and Pils-
bry (1911:260), from the collection of the Portland Society
of Natural History (original catalog number 220). Johnson
(1949:227, pl. 27, fig. 22) validly (Code, Art. 74.5) desig-
nated MCZ 165606 as the lectotype, although this may not
have been one of the aforementioned “cotypes” as his fig-
ure suggests a shell in good condition, contrary to his own
statement and to that of Hyatt and Pilsbry (1911:259). John-
son (1996:192) subsequently identified MCZ 156098 (10
specimens recorded in the MCZ collection ledger) as para-
lectotypes. Mighels (1845:20) indicated the type locality as
“Oahu.” However, Hyatt and Pilsbry (1911:259) considered
this to be an error and that the type locality is “Molokai.”
Like other valid type material, USNM 5497 is cataloged in
the 5000 series with material from the U.S. Exploring Expe-
dition and has an “original number” 95 corresponding to
Gould’s illustration (1856, pl. 7, fig. 95). However, there is
no information for this lot in the original handwritten USNM
ledger regarding locality or type status and there is no indica-
tion why it was interpreted as type material. Therefore, we
consider USNM 5497 as only a possible paralectotype.
nucleola Gould, 1845; Achatinella
Achatinella nucleola Gould, 1845:28; 1862:196.
Current taxonomic status: Amastra (Amastrella) nucleola
(Gould, 1845). Valid species (Hyatt and Pilsbry, 1911:153;
Cowie et al., 1995:98).
NUMBER 647
13
Type material: Paralectotypes USNM 611221 (3 spms, Figure 3K).
Type locality: “Sandwich Islands.”
Remarks: No holotype was designated nor a figure provided
with the original description. Hyatt and Pilsbry (1911:154)
noted NYSM 1172 as “type,” but this is not a lectotype des-
ignation as they did not say “the type” and the lot contained
multiple specimens. Johnson (1964:117) validly (Code, Art.
74.5) designated MCZ 169265 (ex NYSM 194, original no.
A1172) as the lectotype and identified MCZ 169226 (30
spms) and USNM 611221 (3 spms, in the series received
on permanent loan from the New York State Museum, via
W. J. Clench) as paralectotypes, both separated from the
same original lot as the lectotype. The original handwrit-
ten USNM ledger indicates the locality as Kauai, Hawaiian
Islands, that the specimens were collected by Newcomb, and
that they are paratypes.
rex Sykes, 1904; Amastra
Amastra (Kauaia) rex Sykes, 1904:159, two unnumbered text figures
(Figure 4A).
Current taxonomic status: Tropidoptera rex (Sykes, 1904).
Valid species (Hyatt and Pilsbry, 1911:126 [as Pterodiscus];
Cowie et al., 1995:118).
Type material: Syntypes USNM 499926 (1 spm, Figure 4B),
USNM 180852 (4 spms, Figure 4C).
Type locality: “Summit of Konahuanui, Oahu, Hawaiian Islands.”
Remarks: No holotype was designated but a figure was provided
with the original description, in which Sykes (1904:160)
stated that “this very interesting shell was collected by Mr.
Ernest Lyman, and was kindly sent to me by Prof. H. W.
Henshaw.” Although the term “shell” is used in the singular,
it may not have been used to connote a single specimen, but
rather in the sense of the species, as there is more than one
lot of possible type material in the USNM as well as one
lot in the NHMUK. The label of USNM 499926 notes this
specimen as “Lyman Collection from Henshaw.” The origi-
nal handwritten USNM ledger entry for USNM 499926
(recorded 19 January 1938) indicates that the specimen lo-
cality is Konahuanui, Oahu, received from Henshaw, col-
lected by E. Lyman, and its status as “co-type.” A second lot
in the USNM general collection (USNM 180852) also ap-
pears to be type material from “Konahuanui, Oahu, Sandw.
Is.” received from Lyman through Henshaw and recorded in
the USNM ledger on 30 December 1904. Whereas USNM
180852 was cataloged the same year as the publication of
the original description, there is no type status indicated for
it in the ledger as there is for USNM 499926. The signifi-
cance of the lengthy interval between the entries of the two
lots in the ledger is unclear. An additional syntype is in the
Sykes collection of the Natural History Museum, London
(NHMUK 1962195). No lectotype is here designated, pend-
ing study of the NHMUK syntype.
rubens Gould, 1845; Achatinella
Achatinella rubens Gould, 1845:27; 1862:195.
Current taxonomic status: Amastra (Amastrella) rubens (Gould,
1845). Valid species (Hyatt and Pilsbry, 1911:193; Cowie et
al., 1995:99).
Type material: None found. Invalid lectotype designation by
Johnson (1964:142). See remarks.
Type locality: “Sandwich Islands.”
Remarks: No holotype was designated nor a figure provided
in the original description. Johnson (1964:142, pl. 42, fig.
6) designated MCZ 169350 as the lectotype and identi-
fied MCZ 169351 (20 spms) and USNM 611220 (2 spms,
Figure 5A) as paralectotypes, all derived from the same
lot (original no. A1471). An early handwritten label ac-
companying MCZ 169350 indicates that this lot bore the
original number A1432 and, correctly, the species name
Achatinella viridans. USNM 611220 is also A. viridans. A
much later label (the most recent one) now associated with
MCZ 169350 bears the original number A1432 but the
species name Achatinella rubens. It has a subsequent an-
notation “[A 1471],” the original number for the Achati-
nella rubens lot. We consider this latter label to be in error,
inasmuch as it is associated with the Achatinella viridans
lot (A1432) but identifying the material as Achatinella ru-
bens. The subsequent annotation on this label (“A1471”)
reflects an assumption that the lot is the original Acha-
tinella rubens lot. Furthermore, Hall (1875:13) indicated
that A1471 was not found in the NYSM collection and
so could not have been transferred from the NYSM to
the MCZ.
Achatinella viridans and Achatinella rubens are very
different species, the latter now placed in Amastra. Gould
(1845:27) described A. rubens as possessing six whorls and a
chestnut apex, with the remainder of the shell straw colored
and irregularly covered with brown epidermis. In contrast,
Achatinella viridans was described by Mighels (1845:20) as
possessing five whorls, green in color with lighter streaks,
with an aperture stained pink just within the margin, and a
slightly thickened lip. Johnson (1964:142) apparently took
the specimens in the A. viridans lot (A1432) to be A. rubens
on the basis of the incorrect later label. His lectotype des-
ignation (Johnson, 1964:142) for Achatinella rubens of a
specimen that was not a syntype is therefore invalid (Code,
Art. 74.2).
A careful search of the USNM collections yielded
no type material of A. rubens. The specimens in USNM
611220, considered to be paralectotypes by Johnson
(1964:142), are also A. viridans. A thorough search of the
MCZ collection is needed to determine if type material of
this species is still there, although it appears unlikely. Des-
ignation of a neotype of Achatinella rubens Gould may be
warranted.
14
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 4. A. Sykes’ (1904) unnumbered text figures of Amastra rex. B. Syntype (USNM 499926; 13.7 mm) of A. rex.
C. Syntypes (USNM 180852; 11.0 mm) of A. rex. Dimensions are shell diameter (width). If more than one shell in the
lot is figured, dimension is of the shell indicated by an asterisk. Scale bars: 5 mm.
NUMBER 647
15
FIGURE 5. A. Johnson’s (1964) invalidly designated “lectotype” (USNM 611220; 18.5 mm) of Achatinella rubens.
B. Paralectotypes (USNM 4710; 18.6 mm) of Amastra rubinia. C. Possible syntype (USNM 5449; 5.6 mm) of Helix
rubiginosa. D. Gould’s (1856: pl. 4, figs. 49, 49a,b) figured Helix rubiginosa. Dimensions are shell height (length)
(A, B) and width (C). If more than one shell in the lot is figured, dimension is of the shell indicated by an asterisk.
Scale bars: A and B, 5 mm; C, 1 mm.
16
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
rubinia Hyatt and Pilsbry, 1911; Amastra
Amastra (Amastrella) rubens var. rubinia Hyatt and Pilsbry, 1911:193, pl.
32, fig. 16.
Current taxonomic status: Amastra (Amastrella) rubens rubinia
Hyatt and Pilsbry, 1911. Valid subspecies (cf. Cowie et al.,
1995:99, who were noncommittal about its status).
Type material: Paralectotypes USNM 4710 (3 spms, Figure 5B).
Type locality: “Kukuiala,” “Oahu.”
Remarks: Hyatt and Pilsbry (1911:193, pl. 32, fig. 16) mentioned
“a series from Kukuiala” and provided dimensions for two
specimens with catalog numbers ANSP 92481, which they
illustrated, and USNM 4710. They did not designate a ho-
lotype. Baker (1963:199) designated ANSP 92481 as the
lectotype (“TSD now”). There are two lots in the USNM:
USNM 4710 and USNM 4710a. The original handwritten
catalog ledger lists only USNM 4710, with two specimens
from “Sandw Is” received from Dr. Newcomb (who col-
lected shells in Hawaii between 1850 and 1856; Clarke,
1960:136), “original number” 92. USNM 4710a is not
listed in the ledger. It is probable that two lots were inadver-
tently assigned the same catalog number, the second distin-
guished from the first by the addition of the suffix “a” when
the error was discovered, but this addition was not noted
in the ledger. The specimens in USNM 4710 (three of them,
in contrast to the ledger entry), match the original descrip-
tion: “the outer layer of cuticle is almost wholly wanting,
leaving the shell whitish or yellowish with more or less pink
suffusion, most pronounced on the latter half of the last
whorl.” The two specimens in USNM 4710a do not match
the description, as the periostracum of both is much more
intact. The significance of the “original number” 92 in the
ledger and on one of the labels of USNM 4710a (which also
says “rubinia”) is not clear. For other species in this catalog,
these “original numbers” refer to figures of Gould (1856).
However, Gould’s (1856) pl. 7, fig. 92 is an illustration of
Partula varia Broderip, 1832 from Huahine, Society Islands.
It is possible that one or more labels have been placed with
the wrong shells. Despite this confusion we treat the three
specimens labeled as USNM 4710 as paralectotypes of ru-
binia Hyatt and Pilsby, and the two specimens labeled as
USNM 4710a as having no type status.
Family endodontidae
rubiginosa Gould, 1846; Helix
Helix rubiginosa Gould, 1846:173; 1852:50; 1856: pl. 4, figs. 49, 49a–c;
1862:21, 243.
Current taxonomic status: Synonym of Helix jugosa Mighels,
1845, now placed in Cookeconcha (Solem, 1976:222;
Cowie et al., 1995:144).
Type material: Possible syntype USNM 5449 (Figure 5C).
Type locality: “Kauai, Sandwich Islands.”
Remarks: No holotype was designated nor a figure provided with
the original description, although figures were provided sub-
sequently by Gould (1856: pl. 4, figs. 49, 49a–c). It has not
been determined that the description was based on a single
specimen, and the original description does not imply or re-
quire that it was based on more than one specimen. Solem
(1976:222) noted that USNM 5449 “is quite different from
Gould’s . . . type figures” of Helix rubiginosa and considered
(Solem, 1976:220) it to be a mislabeled specimen of Helix
hystrix Pfeiffer, 1846 (synonym of Helix setigera Gould, see
next entry), indicating that either there were multiple syn-
types (i.e., the type series contained at least USNM 5449
and the shell illustrated), or the shell illustrated was seen
only subsequent to the 1846 publication, or Gould mixed up
his specimens prior to publishing the illustrations. Johnson
(1964:53) inferred a “holotype” by “noting the “Figured ho-
lotype USNM 5449”; this is not a valid lectotype fixation (see
section in the Introduction regarding lectotype designations).
However, Solem (1976:222) regarded Johnson’s lectotype fix-
ation as valid and that therefore Helix rubiginosa Gould was
a synonym of Helix hystrix; but he nonetheless recommended
“that the historical usage of these names be continued,” that
is, that the synonymy not be recognized. Because Johnson’s
action is here not considered to be a valid lectotype fixation,
it is not necessary to consider the two taxa synonyms.
Cowie et al. (1995:144) followed Solem’s (1976:222)
placement of Helix rubiginosa as a synonym of Helix jugosa
Mighels, 1845 (now placed in Cookeconcha). We refrain
from designating USNM 5449 as the lectotype of Helix ru-
biginosa, given that Solem (1976:220, 222) considered it
mislabeled, and treat it as a probable syntype, pending fur-
ther research to establish its true identity. Helix rubiginosa
Gould, 1846 is a primary junior homonyn of Helix sericea
form rubiginosa Rossmässler, 1838.
The type material was collected by J. P. Couthouy (Gould,
1852:51) and the USNM ledger notes that it was received
from the U.S. Exploring Expedition, with “original number”
49. Gould’s (1856: figs. 49, 49a, 49b) figures of the species
he described as Helix rubiginosa are reproduced here as Fig-
ure 5D, although they do not appear to illustrate the species
represented by USNM 5449 (Figure 5C). An additional lot of
possible type material, USNM 20926, with “original num-
ber” f49, is cataloged in the original handwritten USNM led-
ger as “rubiginosa Gld” from “Sandw Is,” received from the
U.S. Exploring Expedition, but it could not be found.
setigera Gould, 1844; Helix
Helix setigera Gould, 1844:174; 1852:55; 1856: pl. 4, figs. 52*, 52*a–c (as
Helix hystrix); 1862:194.
Current taxonomic status: Cookeconcha setigera (Gould, 1844).
Valid species; n. comb.
NUMBER 647
17
Type material: Lectotype USNM 20930 (Figure 6A), here desig-
nated; paralectotypes USNM 1418218 (2 spms, Figure 6B).
Type locality: “Sandwich Islands.”
Remarks: No holotype was designated nor a figure provided
with the original description, although figures were pro-
vided subsequently by Gould (1856: pl. 4, figs. 52*, 52*a–
c) for Helix hystrix Pfeiffer, 1846, which he treated as a
synonym of setigera Gould, 1844 and the valid name, as
he incorrectly considered the latter to be preoccupied (see
below). The locality in the original description was “Sand-
wich Islands” (Gould, 1844:174), with the specific island
not indicated. Gould (1852:56) mentioned specimens from
East Maui that “differ somewhat from those originally
examined” and are therefore not part of the type series,
but whether the original specimens were also from East
Maui is not clear. Johnson (1964:148) noted the “figured
neoholotype, selected by Gould, USNM 5453,” although
Gould did not select any name-bearing type specimen in
any of his publications. The original description does not
imply or require that it was based on more than one speci-
men, but subsequently Gould (1852:56) used the phrase
“those originally examined,” which demonstrates that it
was indeed based on more than one specimen. Further-
more, Johnson (1964:148–149) identified additional type
material in the MCZ, including MCZ 169367 (6 spms; ex
NYSM 278, original no. A 765) and MCZ 87861 (1 spm;
ex BSNH 4197). Therefore, Johnson’s identification of a
“neoholotype” is not a valid lectotype fixation according
to either Art. 74.5 or Art. 74.6 of the Code. The type ma-
terial was provided by J. D. Brackenridge and J. Drayton
(Gould, 1852:56) and the original handwritten ledger indi-
cates that it was received from the U.S. Exploring Expedi-
tion, “original number” 52. As discussed, these “original
numbers” correspond to the numbers of Gould’s (1856) il-
lustrations; Gould’s figs. 52 and 52a–m are labeled as Helix
bursatella Gould, 1846, although they probably represent
a number of species and forms (Solem, 1976:395). They
are not the figures of Helixhystrix,” which are figs. 52*,
52*a–c. Solem (1976:220), under the heading Cookecon-
cha hystrix (Pfeiffer, 1846), indicated a lectotype, USNM
5453, without explicitly stating that it was the lectotype
of Helix hystrix. Two paragraphs further down the page
he stated that the “lectotype of Helix setigera Gould, 1844
(not Sowerby, 1841) is juvenile.” Thus it is clear that Solem
(1) accepted the homonymy noted by Gould, (2) accepted
that Johnson’s identification of a “neoholotype” was a
valid lectotype designation, and (3) when referring to the
“lectotype” was in fact referring to setigera and not hys-
trix. Despite the logic of this interpretation, nowhere did
Solem explicitly state that USNM 5453 was the lectotype of
Helix setigera Gould, and therefore no lectotype was des-
ignated. Although Johnson (1964:148) referred to USNM
5453 (Figure 6C) as Gould’s “figured” specimen, it does
not match Gould’s illustrations of Helix hystrix, which
are figs. 52*, 52*a–c (Figure 6D), not 52a–c as incorrectly
cited by Johnson. Thus USNM 5453 is not type material of
Helix setigera. An additional lot of possible type material,
USNM 20930, listed as Pitys hystrix, “original number”
f52*, from “Sandw Is” and received from “Pse” (= W. H.
Pease) was found. Three specimens are in this lot (1 adult, 2
juvenile); the largest specimen is a good match for Gould’s
figures of Helixhystrix” and is here selected as the lecto-
type of Helix setigera Gould, 1844.
The name Helix setigera Gould, 1844 has been con-
sidered a junior primary homonym of Helix setiger Sow-
erby, 1841 (Gould, 1852:56; Solem, 1976:220; Cowie et
al., 1995:144). Gould (1852:55) synonymized it with Helix
hystrix Pfeiffer, 1846. Helix hystrix was described from
“Ins. Sandwich” by Pfeiffer (1846:67), who attributed it to
“Mighels (mss?)” and listed “H. setigera Gould in sched.”
in synonymy. Thus, given the supposed homonymy, Gould
(1852:56) used hystrix as the valid name, it being the next-
oldest available name for this species. However, setiger
Sowerby and setigera Gould are not homonyms, by the fol-
lowing reasoning. The name setiger Sowerby may be either
a noun in apposition or an adjective in the masculine gender
(Code, Art. 31.2.2). If treated as an adjective in combina-
tion with Helix (feminine), its gender would require manda-
tory change (Code, Art. 34.2.1) to setigera, rendering setiger
Sowerby and setigera Gould homonyms. However, in the
absence of evidence that it has been treated as an adjective,
and the fact that it was originally introduced in combination
with a feminine genus name, suggesting that it was intended
as a noun in apposition, setiger Sowerby should be consid-
ered as such (Code, Art. 31.2.2). The names setiger and se-
tigera are then deemed to be spelled differently (Code, Art.
57, Art. 58), the replacement of setigera Gould, 1844 by
hystrix Pfeiffer, 1846 was not necessary, and setigera Gould
is the valid name.
Family helicarionidae
cicercula Gould, 1846; Helix
Helix cicercula Gould, 1846:171; 1852:43; 1856: pl. 5, figs. 73, 73a–c;
1862:20, 243.
Current taxonomic status: Philonesia (Philonesia) cicercula
(Gould, 1846). Valid species (Cowie et al., 1995:159).
Type material: Syntype USNM 20948 (Figure 7A).
Type locality: “Mountains of Hawaii.”
Remarks: No holotype was designated nor a figure provided
with the original description, although figures were pro-
vided subsequently by Gould (1856: pl. 5, figs. 73, 73a–c).
Johnson (1964:54) noted the “Figured holotype USNM
20948” but also noted MCZ 169080 (8 spms; ex NYSM
264, original no. A 754) as additional type material. Al-
though the original description does not imply or require
18
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 6. A. Lectotype (USNM 20930; 5.6 mm), here designated, of Helix setigera. B. Paralectotypes (USNM
1418218; 4.0 mm) of H. setigera. C. Non-type material (USNM 5453; 4.5 mm) of H. setigera. D. Gould’s (1856: pl.
4, figs. 52*, 52*a,b) figured Helix setigera. Dimensions are shell diameter (width). If more than one shell in the lot is
figured, dimension is of the shell indicated by an asterisk. Scale bars: 1 mm.
NUMBER 647
19
that it was based on more than one specimen, the fact
that additional type material was recognized by Johnson
means this is not a valid lectotype fixation (see section in
the Introduction regarding lectotype designations). There
are notable differences between Gould’s figured specimen
(Figure 7B) and USNM 20948: the shell is more tightly
coiled and hence the width of the last whorl is smaller in
USNM 20948, the periphery is higher and less carinate in
USNM 20948, and the basal body whorl half a whorl back
from the aperture is more inflated in USNM 20948. USNM
20948 may therefore not be Gould’s (1856: pl. 5, figs. 73,
73a, 73b) figured specimen and we refrain from designat-
ing it as the lectotype and treat it as a syntype only. It is
possible that a specimen in MCZ 169080 may be a bet-
ter match to Gould’s illustrations, although it is unclear
whether MCZ 169080 was derived from the same lot as
USNM 20948. The type material was collected by J. Dray-
ton and J. D. Brackenridge (Gould, 1852:44) and the origi-
nal handwritten ledger indicates that USNM 20948 was
received from the U.S. Exploring Expedition, with “origi-
nal number” f73. The ledger also lists USNM 5475, with
“original number” 73, as “cicercula” from “Hawaii” from
the U.S. Exploring Expedition; this lot is therefore the most
likely to have contained the figured specimen, but it could
not be found.
cryptoportica Gould, 1846; Helix
Helix cryptoportica Gould, 1846:171; 1852:44; 1856: pl. 5, figs. 72, 72a–c;
1862:20, 243.
Current taxonomic status: Philonesia (Philonesia) cryptoportica
(Gould, 1846). Valid species (Baker, 1940:120; Cowie et al.,
1995:159).
Type material: Lectotype USNM 5474 (Figure 7C), here desig-
nated.
Type locality: Mountains of Oahu, Sandwich Islands (Gould,
1852:45).
Remarks: No holotype was designated nor a figure provided
with the original description, although figures were pro-
vided subsequently by Gould (1856: pl. 5, figs. 72, 72a–c).
No locality was given in the original description but the
locality was indicated in subsequent elaborations by Gould
(1852:44). Johnson (1964:63) noted the “Figured holotype
USNM 5474” but he also noted MCZ 169110 (2 spms; ex
NYSM 233, original no. A 747) as “paratypes.” Although
the original description does not imply or require that it
was based on more than one specimen, the fact that addi-
tional type material was recognized by Johnson means this
is not a valid lectotype fixation (see section in the Introduc-
tion regarding lectotype designations). We therefore here
designate USNM 5474 as the lectotype. The handwritten
USNM ledger confirms that the type material was received
from the U.S. Exploring Expedition. USNM 5474, with
“original number” 72, closely matches Gould’s (1856: pl.
5, figs. 72, 72a–c) figured specimen (Figure 7D). An addi-
tional lot of possible type material, USNM 20958 (1 spm),
but with no original numbers noted, which was listed in the
original handwritten USNM ledger as “Conulus cryptopor-
ticus Gld (non Pse)” from “Sandw Is” and received from
the U.S. Exploring Expedition, could not be found.
exaequata Gould, 1846; Helix
Helix exæquata [sic] Gould, 1846:171; 1852:47; 1856: pl. 5, figs. 61, 61a–c;
1862:19, 243.
Current taxonomic status: Hiona (Nesocyclus) exaequata
(Gould, 1846). Valid species (Baker, 1940:186; Cowie et al.,
1995:156).
Type material: Possible paralectotypes USNM 5463 (Figure 7E;
the larger of two specimens originally in this lot, incorrectly
identified as exaequata), USNM 20947 (2 spms).
Type locality: Kauai, Wailua Valley (Baker, 1940:187).
Remarks: No holotype was designated nor a figure provided
with the original description, although figures were pro-
vided subsequently by Gould (1856: pl. 5, figs. 61, 61a–c).
Gould (1846:171) indicated the type locality as “Kauai,
Sandwich Islands.” The type material was collected by J.
P. Couthouy (Gould, 1852:47) and the original handwrit-
ten ledger indicates that USNM 5463 was received from
the U.S. Exploring Expedition, with “original number” 61.
Baker (1940:188, pl. 42, figs. 7–9) illustrated the largest
shell in NYSM 259 (original number A748; now MCZ
169134, 13 spms) and unambiguously and explicitly se-
lected this specimen as the name-bearing type (lectotype)
by stating that it “is taken as the type” (Code, Art. 74.5).
Johnson (1964:73) noted a “Figured holotype USNM
5463,” but since Baker had already designated a lectotype,
this statement is irrelevant from the perspective of lectotype
designation/fixation.
The original handwritten USNM ledger indicates that
there were two specimens in USNM 5463 when it was cata-
loged. Baker (1940:188) also noted two shells but stated
that “USNM 5463 contains the base of a broken shell that
appears to have been a smaller specimen of this species
[Hiona exaequata], possibly that figured in the U.S. Explor-
ing Expedition shells, and a larger specimen of a very differ-
ent shell (casually examined, it looked like my H. pilsbryi).”
The single specimen remaining in USNM 5463 (Figure 7E)
has a domed shell, not a discoidal one as in the original
description of Helix exaequata, and is indeed a good match
for the three illustrations of Hiona pilsbryi (Baker, 1940: pl.
38, fig. 4). It differs from the specimen in Gould’s (1856:
pl. 5, figs. 61, 61a, 61b) figure (Figure 7F), which is indeed
discoidal and which also matches Baker’s (1940: pl. 42, figs.
7–9) figures of H. exaequata. Consequently, the specimen
remaining in USNM 5463 is probably the “larger specimen
20
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 7. A. Syntype (USNM 20948; 5.7 mm) of Helix cicercula. B. Gould’s (1856: pl. 5, figs. 73, 73a,b) fig-
ured Helix cicercula. C. Lectotype (USNM 5474; 5.6 mm), here designated, of Helix cryptoportica. D. Gould’s
(1856: pl. 5, figs. 72, 72a,b) figured Helix cryptoportica. E. Possible paralectotype (USNM 5463; 10.9 mm) of
Helix exaequata. F. Gould’s (1856: pl. 5, figs. 61, 61a,b) figured Helix exaequata. Dimensions are shell diameter
(width). If more than one shell in the lot is figured, dimension is of the shell indicated by an asterisk. Scale bars:
1 mm.
NUMBER 647
21
of a very different shell” (Baker, 1940:188). If it indeed was
part of the original lot, then it is a paralectotype of H. exae-
quata, though misidentified as that species. However, as it is
so distinct from true H. exaequata, it is difficult to imagine
Gould making such an error and it is more likely that it
was misplaced in the box some time later and was not part
of the original type series. A careful search of the USNM
collections failed to produce the figured specimen or any
additional type material of Helix exaequata, including the
broken shell noted by Baker (1940:188). An additional lot
of possible type material, USNM 20947 (2 spms), listed in
the original handwritten USNM ledger as “exaequata Gld,”
from “Hawaii” but with only one specimen indicated, was
collected by the U.S. Exploring Expedition and received
from “Pse” (=W. H. Pease). However, there are no original
numbers associated with this lot and it is considered pos-
sible type material only. Johnson (1964:73) noted the pres-
ence of additional type material in the collections of the
MCZ, including MCZ 169134 (13 spms; ex NYSM 259,
original no. A748; the lectotype is the largest shell in this
lot, as designated by Baker—see above) and MCZ 87862 (2
spms; ex BSNH 4368).
subtilissima Gould, 1846; Helix
Helix subtilissima Gould, 1846:177; 1852:48; 1856: pl. 5, figs. 62, 62a–c;
1862:24.
Current taxonomic status: Euconulus (Nesoconulus) subtilissi-
mus (Gould, 1846). Valid species (Baker, 1941:215; Cowie
et al., 1995:154).
Type material: Holotype (by monotypy) USNM 5464 (Figure 8A).
Type locality: “Maui, Sandwich Islands” [probably east Maui
(Baker, 1941:215)].
Remarks: No holotype was designated but by saying “this lit-
tle pellucid shell, though imperfect” Gould (1846:177,
1852:49) indicated that his description was based on just
this single shell, which is therefore the holotype by mono-
typy. No figure was provided with the original description,
although figures were provided subsequently by Gould
(1856: pl. 5, figs. 62, 62a–c). Baker (1941:216) noted that
“USNM 5464 and NYSM A-5881 consist of shells of the
more depressed form, such as occurs at lower elevations on
Haleakala (BBM [i.e., BPBM] 11286; [Baker, 1941:] pl. 53,
figs. 1–3) and is represented in Gould’s figure; the USNM
specimen is taken as type.” Because Gould (1846:177,
1852:49) based his description on just one specimen, the
additional material noted by Baker in NYSM and BPBM
can only be topotypical. The holotype was collected by
J. Drayton (Gould, 1852:49) and the original handwritten
ledger indicates that it was received from the U.S. Exploring
Expedition. The specimen in USNM 5464, with “original
number” 62, although badly damaged, matches Gould’s
(1856: pl. 5, figs 62, 62a, 62b) figured specimen (Figure 8B).
Family helicinidae
uberta Gould, 1847; Helicina
Helicina uberta Gould, 1847:202; 1852:94; 1856: pl. 7, figs. 114, 114a–c;
1862:37.
Current taxonomic status: Orobophana uberta (Gould, 1847).
Valid species (Neal, 1934:19; Cowie et al., 1995:18).
Type material: Lectotype (Pilsbry and Cooke, 1908:13) USNM
5516 (Figure 8C); possible paralectotype USNM 20202
(8 spms).
Type locality: “Maui, and Oahu Mountains” [possibly restricted
to the “back of Leilehua, in the Waianae mountains” (Pils-
bry and Cooke, 1908:13)].
Remarks: No holotype was designated nor a figure provided
with the original description, although figures were pro-
vided subsequently by Gould (1856: pl. 7, figs. 114, 114a–
c). However, Gould stated that he had specimens collected
from Maui and Oahu, indicating that there were multiple
syntypes. Pilsbry and Cooke (1908:13) noted “the type shell
(No. 5516, Smithsonian Institution),” thereby designating
the lectotype (Code, Art. 74.5), and Johnson (1964:162)
recognized this specimen as the “Figured holotype USNM
5516.” Pilsbry and Cooke (1908:13) did not formally re-
strict the type locality, though stated that the “only local-
ity at which typical forms are collected at present is in the
Waianae mountains” and “Specimens from back of Leile-
hua, in the Waianae mountains, agree very closely with the
type shell.” Johnson (1964:162) identified additional type
material in the MCZ, including MCZ 169411 (5 spms), and
two lots obtained from the Smithsonian Institution, MCZ
216585 (3 spms) and MCZ 186722 (1 spm), all derived
from NYSM 292 (original no. G2626). Johnson (1964:172)
noted that the locality on the original label, “Taheiti,” was
an error. The type material was collected by Pickering and
Case (Oahu) and Drayton (Maui) (Gould, 1852:95). USNM
5516, with “original number” 114, closely matches Gould’s
(1856: pl. 7, figs 114, 114a, 114b) figured specimen (Fig-
ure 8D). An additional lot of possible type material, USNM
20202 (8 spms), is listed in the original handwritten USNM
ledger as “Helicina uberta” from “Maui & Oahu” with 11
specimens from the U.S. Exploring Expedition. However,
there are no original numbers associated with this lot and
therefore we consider it only as possible type material.
Family PuPillidae
lyrata Gould, 1843; Pupa
Pupa lyrata Gould, 1843:139; 1844: pl. 16, fig. 16; 1862:189.
Current taxonomic status: Lyropupa (Lyropupa) lyrata (Gould,
1843). Valid species (Pilsbry and Cooke, 1920:233; Cowie
et al., 1995:131).
22
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 8. A. Holotype (USNM 5464; 2.6 mm) of Helix subtilissima. B. Gould’s (1856: pl. 5, figs. 62,
62a,b) figured Helix subtilissima. C. Lectotype (USNM 5516; 4.8 mm) of Helicina uberta. D. Gould’s
(1856: pl. 7, figs. 114, 114a,b) figured Helicina uberta. E. Paralectotypes (USNM 64344; 2.3 mm) of Pupa
lyrata. F. Syntypes (USNM 117931; 11.6 mm) of Succinea aperta. G. Lectotype (USNM 5420; 9.7 mm),
here designated, of Succinea canella. H. Paralectotypes (USNM 20853; 9.0 mm) of S. canella. I. Gould’s
(1856: pl. 2, figs. 20, 20a) figured Succinea canella. Dimensions are shell diameter (C) and height (length;
E–I). If more than one shell in the lot is figured, dimension is of the shell indicated by an asterisk. Scale bars:
1 mm.
NUMBER 647
23
Type material: Paralectotypes USNM 64344 (2 spms, Figure 8E).
Type locality: Oahu [possibly restricted to Nuuanu (Pilsbry and
Cooke, 1920:234)].
Remarks: No holotype was designated nor a figure provided
with the original description, although a figure was provided
subsequently by Gould (1844: pl. 16, fig. 16). The locality
given in the original description was listed simply as “Ha-
waiian Islands.” Gould (1862:189) subsequently refined the
locality to “Maui, Sandwich Islands.” Pilsbry and Cooke
(1920:234) noted that the original material was recorded
from Kauai. However, they concluded that both Maui and
Kauai were incorrect and that “P. lyrata was doubtless from
Oahu, where typical examples have been taken in Nuuanu
valley.” Pilsbry and Cooke (1920:235, pl. 19, figs. 4, 5)
identified the “Type and paratypes no. 219, G. 2687” in
NYSM and, by illustrating one of these specimens (fig. 4) as
the “Type,” designated it as the lectotype (Code, Art. 74.5),
with the shell illustrated as a “Paratype” (fig. 5) becoming
a paralectotype. Johnson (1964:107) noted the “figured ho-
lotype MCZ 169233, ex NYSM 219, original no. G2687,”
that is, the same specimen identified by Pilsbry and Cooke
as the type, which by then had been transferred from NYSM
to MCZ (Johnson, 1964:15). Johnson (1964:107) identified
additional type material in the collections of the MCZ, in-
cluding MCZ 169234 (1 spm; also ex NYSM 219, original
no. G2687). The original USNM handwritten ledger indi-
cates that USNM 64344 (2 spms) was also received from the
U.S. Exploring Expedition, and we therefore consider these
specimens as additional paralectotypes.
Family succineidae
aperta Lea, 1838; Succinea
Succinea aperta Lea, 1838:101, pl. 23, fig. 107.
Current taxonomic status: Succinea aperta Lea, 1838. Valid spe-
cies (Cowie et al., 1995:152).
Type material: Syntypes USNM 117931 (2 spms, Figure 8F).
Type locality: “Banks of Columbia River.”
Remarks: No holotype was designated but a figure was provided
with the original description. Lea did not mention where
type material was deposited. According to the original hand-
written USNM ledger, two type specimens were received
from the Lea collection, collected by Nuttall in “Oregon?”
We refrain from designating either of these two specimens
as a lectotype, which would be better done in the context
of a study of North American Succineidae and following a
search for additional syntype material.
Gould (1846:182) stated that Succinea rotundata
Gould resembled S. aperta Lea. In a footnote, Gould (in
Binney, 1851:66–67) stated that “S. aperta is undoubtedly
a species belonging to the Sandwich Islands, described by
me under the name S. rotundata.” Cowie et al. (1995:152)
included Succinea aperta as among taxa questionably in-
cluded in the Hawaiian fauna and noted it as a possible
synonym of S. rotundata Gould, 1846, on the authority of
Baldwin (1893:24). For these reasons, we include S. aperta
in this catalog of Hawaiian taxa, while acknowledging that
it is probably not a Hawaiian species.
canella Gould, 1846; Succinea
Succinea canella Gould, 1846:184; 1852:27; 1856: pl. 2, figs. 20, 20a,b;
1862:29.
Current taxonomic status: Succinea canella Gould, 1846. Valid
species (Cowie et al., 1995:150).
Type material: Lectotype USNM 5420 (Figure 8G), here desig-
nated; paralectotypes USNM 20853 (2 spms, Figure 8H).
Type locality: “Maui, Sandwich Islands.”
Remarks: No holotype was designated nor a figure provided with
the original description, although figures were provided
subsequently by Gould (1856: pl. 2, figs. 20, 49a,b). John-
son (1964:51) noted the “Figured holotype USNM 5420”
but also identified additional type material in the collections
of the MCZ, including MCZ 169072 (14 spms; ex NYSM
175, original no. G2643) and MCZ 161661 (2 spms; ex
Smithsonian Institution). Although the original description
does not imply or require that it was based on more than
one specimen, because additional type material was recog-
nized by Johnson, his noting of the “Figured holotype” is
not a valid lectotype fixation (see section in the Introduc-
tion regarding lectotype designations). We therefore here
designate USNM 5420 as the lectotype. It is unclear if the
additional type material noted above was from the same lot
as the lectotype. The original handwritten ledger indicates
the lectotype as being received from the U.S. Exploring Ex-
pedition, with “original number” 20. USNM 5420 closely
matches Gould’s (1856: pl. 2, figs. 20, 20a) figured specimen
(Figure 8I). The two specimens in an additional lot, USNM
20853, original numbers “=G546 =f20,” received from the
U.S. Exploring Expedition are considered paralectotypes.
cepulla Gould, 1846; Succinea
Succinea cepulla Gould, 1846:182; 1852:16; 1856: pl. 2, figs. 15, 15a,b;
1862:27, 244.
Current taxonomic status: Succinea cepulla Gould, 1846. Valid
species (Cowie et al., 1995:150).
Type material: Syntypes USNM 5415 (1 spm, Figure 9A), USNM
20868 (2 spms, Figure 9B).
Type locality: “Hawaii.”
Remarks: No holotype was designated nor a figure provided
with the original description, although figures were pro-
vided subsequently by Gould (1856: pl. 2, figs. 15, 15a,b).
Johnson (1964:53) noted several syntypes in the MCZ and
USNM, including USNM 5415, MCZ 169114 (8 spms, ex
24
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
NYSM 176, original no. G2644), MCZ 39646 (1 spm),
and MCZ 216754 (1 spm), the last two lots received from
the Smithsonian Institution. The type specimens were col-
lected by C. Pickering and J. Drayton (Gould, 1852:17).
The specimen in USNM 5415 is similar to Gould’s (1856:
pl. 2, figs. 15, 15a, 15b) figured specimen (Figure 9C) but
is smaller than the measurements provided in the original
description (Johnson, 1964:53), which were “Long. ½, lat.
720, alt. 15 poll.” [approximately 13 mm long, 9 mm wide,
5 mm high]. The respective dimensions of the USNM 5415
specimen are 10 mm, 7 mm, and 5 mm. This specimen is
therefore not designated as a lectotype, pending further
research, notably of the MCZ collections. The original
handwritten ledger indicates that the type material was
received from the U.S. Exploring Expedition, with “origi-
nal number” 15. An additional lot, USNM 20868, origi-
nal numbers “=G303, =–544, f15,” is listed in the original
handwritten ledger from “Hilo Hawaii” received from the
U.S. Exploring Expedition. Ditto marks below the preced-
ing line indicate that this is “cepulla Gld.” We treat these
as syntypes. Additionally, H. W. Henshaw noted on a label
that he considered one of these specimens to not be conspe-
cific but possibly an immature specimen of Succinea vesi-
calis Gould, 1846.
explanata Gould, 1852; Succinea
Succinea explanata Gould, 1852:13; 1856: pl. 2, figs. 31, 31a–c; 1862:223.
Current taxonomic status: Catinella explanata (Gould, 1852).
Valid species (Cowie et al., 1995:148).
Type material: Lectotype USNM 5431 (Figure 9D), here desig-
nated; paralectotype USNM 20870 (1 spm, Figure 9E).
Type locality: “Kauai, Sandwich Islands.”
Remarks: The type material was collected by J. P. Couthouy and
Gould (1852:13) noted that “Mr. Couthouy has labeled the
shell as ‘Testacella’.” This might suggest that Gould had
only a single shell, which therefore would be the holotype
by monotypy. However, there are two lots, USNM 5431 and
20870, received from the U.S. Exploring Expedition. John-
son (1964:73), while noting the “Figured holotype USNM
20870,” also identified additional type material as USNM
5431 (1 spm; “original number” 31) and MCZ 169135 (1
spm; ex NYSM 182, original number A1523), and MCZ
155126 (1 spm; ex Smithsonian Institution). Consequently,
Johnson’s (1964: 73) inference does not constitute valid
lectotype fixation (see section in the Introduction regarding
lectotype designations) and we therefore interpret Gould’s
statement above as using “shell” in the sense of the species.
The shell in USNM 20870, with original numbers “=G405,
445” and f31, is missing its apex. It was not noted as broken
in the original handwritten USNM ledger, so the specimen
may have been damaged subsequently. We here designate
USNM 5431, which matches Gould’s figured specimen
(1856: pl. 2, 31, 31a, 31b; Figure 9F), as the lectotype and
USNM 20870 is a paralectotype.
lumbalis Gould, 1846; Succinea
Succinea lumbalis Gould, 1846:183; 1852:17; 1856: pl. 2, figs. 18, 18a,b;
1862:28, 244.
Current taxonomic status: Succinea lumbalis Gould, 1846. Valid
species (Cowie et al., 1995:151).
Type material: Possible syntype USNM 5418 (Figure 9G).
Type locality: “Kauai, Sandwich Islands.”
Remarks: No holotype was designated nor a figure provided with
the original description, although figures were provided sub-
sequently by Gould (1856: pl. 2, figs. 18, 18a,b). The locality
provided in the original description was “Kauai, Sandwich Is-
lands” (Gould, 1846:183) but both Kauai and the additional
locality “Mauna Kea, Hawaii” were noted subsequently
(Gould, 1852:18), perhaps because Gould saw additional
material in the interval between his 1846 and 1852 publi-
cations. Johnson (1964:105) noted the “Figured holotype
USNM 5418” but also identified additional “paratypes MCZ
169231, ex NYSM 177, original no. G2646.” However,
Johnson’s lectotype fixation is not valid (see section in the
Introduction regarding lectotype designations). The original
handwritten USNM ledger indicates the locality as “Hawaii”
and that it was received from the U.S. Exploring Expedition.
This probably refers to the island of Hawaii, not the Hawai-
ian Islands more generally, which probably would have been
referred to as the Sandwich Islands, as in Gould’s publica-
tions and usually in the original handwritten USNM ledger.
The shell in USNM 5418 closely matches Gould’s (1856: pl.
2, figs. 18, 18a) figured specimen (Figure 9H), and no other
specimens of Succinea lumbalis could be found in the USNM.
However, it is cataloged as from the island of Hawaii, not
the type locality of Kauai, so it is probably not type material
unless (1) “Kauai” in the original description was a mistake,
or (2) Gould (1846:183) in the original description inadver-
tently failed to include “Mauna Kea, Hawaii” as a locality,
or (3) “Mauna Kea, Hawaii” in the subsequent publication
(Gould, 1852:18) was a mistake and our interpretation of the
ledger “Hawaii” is wrong, or (4) the ledger “Hawaii” was
also a mistake. We retain it as a possible syntype only.
oregonensis Lea, 1841; Succinea
Succinea oregonensis Lea, 1841:32; 1844:5.
Current taxonomic status: Succinea oregonensis Lea, 1841.
Valid species (Cowie et al., 1995:152).
Type material: Holotype (by monotypy) USNM 117935 (Figure
10A).
Type locality: “Oregon.”
Remarks: No holotype was designated nor a figure provided
with the original, very brief, Latin description. However,
NUMBER 647
25
FIGURE 9. A. Syntype (USNM 5415; 10.1 mm) of Succinea cepulla. B. Syntypes (USNM 20868; 10.0 mm)
of S. cepulla. C. Gould’s (1856: pl. 2, figs. 15, 15a) figured Succinea cepulla. D. Lectotype (USNM 5431; 9.7
mm), here designated, of Succinea explanata. E. Paralectotype (USNM 20870; 9.5 mm) of S. explanata. F.
Gould’s (1856: pl. 2, figs. 31, 31a,b) figured Succinea explanata. G. Possible syntype (USNM 5418; 11.8 mm)
of Succinea lumbalis. H. Gould’s (1856: pl. 2, figs. 18, 18a,b) figured Succinea lumbalis. Dimensions are shell
height (length). If more than one shell in the lot is figured, dimension is of the shell indicated by an asterisk.
Scale bars: 1 mm.
26
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
the subsequent elaboration (Lea, [1844]:5) stated that a
“single specimen only of this small species was given to me
by Mr. Nuttall” while at the same time stating that material
was present in “My Cabinet, and Cabinets of Prof. Nuttall,
and Dr. Jay.” Under the assumption that Lea did not see
the specimens of Nuttall or Jay, we interpret this to mean
that the original description was based on the “single speci-
men,” as did Pilsbry (1948:842), and that Lea was simply
acknowledging the presence of the species in the collections
of Nuttall and Jay. The specimen is therefore the holotype by
monotypy (Code, Art. 73.1.2). It was noted as “Lea’s type”
and identified as USNM 117935 by Pilsbry (1948:842, fig.
457a). In the original handwritten USNM ledger, USNM
117935 is noted as being in the “type coll.” and collected
by Nuttall.
The Succinea oregonensis of all authors subsequent
to Lea is not the same species as that of Lea, according to
Pilsbry (1948:842), who considered Lea’s S. oregonensis to
have “not been collected again.” Several specimens of Suc-
cinea collected on Oahu in the BPBM collection have been
labeled with an unpublished name attributed to C. F. Ancey,
Succinea oostoma.” (A label name is not nomenclaturally
available, and mentioning the name in the present publi-
cation does not make it available; Code, Arts. 9.6, 16.1.)
C. M. Cooke Jr. compared these specimens with material of
Succinea oregonensis Lea, 1841, sent to him by H. A. Reh-
der, with “Hawaii” as the probable locality (C. M. Cooke
Jr., unpublished notes; correspondence, 1947–1948, with
H. A. Rehder, NMNH). Both Cooke and Rehder considered
S. oostoma and S. oregonensis as the same species. It is
possible that S. oregonensis Lea is actually a Hawaiian spe-
cies. It is included here, pending further research.
rotundata Gould, 1846; Succinea
Succinea rotundata Gould, 1846:182; 1852: 15; 1856: pl. 2, figs. 14, 14a–c;
1862:27, 244.
Current taxonomic status: Catinella rotundata (Gould, 1846).
Valid species (Cowie et al., 1995:149).
Type material: Syntypes USNM 5414 (3 spms, Figure 10B),
USNM 20866 (1 spm, Figure 10C).
Type locality: “Mountains of Oahu, Sandwich Islands.”
Remarks: No holotype was designated nor a figure provided
with the original description, although figures were pro-
vided subsequently by Gould (1856: pl. 2, figs. 14, 14a–c).
Johnson (1964:142) noted the “Figured holotype USNM
5414” in fragments, but also noted a “paratype” (USNM
20866; incorrectly cited as USNM 208666), also damaged.
Although the original description does not imply or require
that it was based on more than one specimen, the fact that
additional type material was recognized by Johnson means
this is not a valid lectotype fixation (see section in the In-
troduction regarding lectotype designations). According to
the original handwritten USNM ledger, the specimens were
already broken at the time the lot was cataloged and the
number of specimens uncertain, as indicated by “?”. Three
broken specimens but with intact embryonic whorls were
found in USNM 5414, the largest badly damaged and the
two smaller specimens in fragments. Due to their condi-
tion, it is difficult to assess whether any one of these three
specimens matches Gould’s (1856: pl. 2, figs. 14, 14a, 14b)
figured specimen (Figure 10D), Johnson’s noting of the “Fig-
ured holotype” notwithstanding. We therefore refrain from
designating a lectotype and treat the specimens in USNM
5414, with “original number” 14, and 20866, with “origi-
nal number” f14, as syntypes. The type material was col-
lected by C. Pickering (Gould, 1852:15) and the original
handwritten ledger indicates that it was received from the
U.S. Exploring Expedition.
Gould (in Binney, 1851:66–67) considered Succinea
rotundata Gould to be a synonym of Succinea aperta Lea.
Baldwin (1893:24) considered them only possibly synony-
mous and was followed by Cowie et al. (1995:152), who
also did not definitively synonymize them, considering ap-
erta as questionably belonging to the Hawaiian fauna.
venusta Gould, 1846; Succinea
Succinea venusta Gould, 1846:186; 1852:22; 1856: pl. 2, figs. 25, 25a,b;
1862:30.
Current taxonomic status: Succinea venusta Gould, 1846. Valid
species (Cowie et al., 1995:152).
Type material: Lectotype USNM 5425 (Figure 10E), here desig-
nated; paralectotypes USNM 20860 (3 spms, Figure 10F).
Type locality: “Hawaii.”
Remarks: No holotype was designated nor a figure provided with
the original description, although figures were provided
subsequently by Gould (1856: pl. 2, figs. 25, 25a,b). John-
son (1964:165) noted the “figured holotype USNM 5425”
but also identified additional type material in the MCZ,
including MCZ 169417 (4 spms; ex NYSM 292, original
no. G2648, erroneously labeled “Taheiti”) and MCZ 39647
(2 spms; ex Smithsonian Institution). Although the original
description does not imply or require that it was based on
more than one specimen, the fact that additional type ma-
terial was recognized by Johnson means this is not a valid
lectotype fixation (see section in the Introduction regarding
lectotype designations). We therefore here designate USNM
5425 as the lectotype. The original handwritten USNM led-
ger confirms the locality as “Hawaii” and that the speci-
men was received from the U.S. Exploring Expedition, with
“original number” 25. The specimen in USNM 5425 closely
matches Gould’s (1856: pl. 2, figs. 25, 25a,b) figured speci-
men (Figure 10G). An additional lot, USNM 20860, origi-
nal numbers “=G575, =4977, =f25,” is listed in the original
handwritten USNM ledger as collected by the U.S. Explor-
NUMBER 647
27
FIGURE 10. A. Holotype (USNM 117935; 6.7 mm) of Succinea oregonensis. B. Syntypes (USNM 5414; 10.1
mm) of Succinea rotundata. C. Syntype (USNM 20866; 10.9 mm) of S. rotundata. D. Gould’s (1856: pl. 2, figs.
14, 14a,b) figured Succinea rotundata. E. Lectotype (USNM 5425; 9.5 mm), here designated, of Succinea venusta.
F. Paralectotypes (USNM 20860; 7.9 mm) of S. venusta. G. Gould’s (1856: pl. 2; fig. 25) figured Succinea ve-
nusta. H. Syntype (USNM 5417) of Succinea vesicalis. Dimensions are shell height (length). If more than one shell
in the lot is figured, dimension is of the shell indicated by an asterisk. Scale bars: 1 mm.
28
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
ing Expedition and received from “Pse” (=W. H. Pease); we
treat these specimens as paralectotypes.
vesicalis Gould, 1846; Succinea
Succinea vesicalis Gould, 1846:183; 1852:21; 1856: pl. 2, figs. 17, 17a;
1862:28, 244.
Current taxonomic status: Succinea vesicalis Gould, 1846. Valid
species (Cowie et al., 1995:152).
Type material: Syntype USNM 5417 (Figure 10H).
Type locality: Mauna Kea, elevation of 7000 ft., Hawaii, Sand-
wich Islands (Gould, 1852:21).
Remarks: No holotype was designated nor a figure provided
with the original description, although figures were pro-
vided subsequently by Gould (1856: pl. 2, figs. 17, 17a). In
the original description, the locality was given as “Mauna
Kea, Hawaii” (Gould, 1846:183) but it was subsequently
refined by Gould (1852:21). It has not been determined that
the description was based on a single specimen, and the
original description does not imply or require that it was
based on more than one specimen, so by noting the “Fig-
ured holotype USNM 5417,” Johnson (1964:53) inferred a
“holotype.” However, this was not a valid lectotype fixation
as he did not explicitly accept that the original description
was based on a single specimen, and it has not been consid-
ered subsequently that the original description was based
on more than one specimen (Code, Art. 74.6). As it is in
fragments, we refrain from designating USNM 5417 as the
lectotype, and following the Code (Rec. 73F), treat it as a
syntype, pending further research. According to the origi-
nal handwritten USNM ledger, the specimen was already
broken at the time it was cataloged, and was received from
the U.S. Exploring Expedition, with “original number” 17.
The type material was obtained by J. P. Couthouy (Gould,
1852:21).
Family Zonitidae
caperata Gould, 1846; Vitrina
Vitrina caperata Gould, 1846:181; 1852:10; 1856: pl. 1, fig. 9, 9a; 1862:26.
Current taxonomic status: Godwinia caperata (Gould, 1846).
Valid species (Cowie et al., 1995:164).
Type material: Lectotype USNM 5409 (Figure 11A), here des-
ignated.
Type locality: “Kauai, Sandwich Islands.”
Remarks: No holotype was designated nor a figure provided
with the original description, although figures were pro-
vided subsequently by Gould (1856: pl. 1, figs. 9, 9a). John-
son (1964:51) noted the “Figured holotype USNM 5409”
but also identified MCZ 135612 (5 spms; ex BSNH from
Smithsonian Institution) as additional type material. Al-
though the original description does not imply or require
that it was based on more than one specimen, the fact that
additional type material was recognized by Johnson means
this is not a valid lectotype fixation (see section in the In-
troduction regarding lectotype designations). We therefore
designate USNM 5409 as the lectotype. The type material
was collected by J. P. Couthouy (Gould, 1852:11). The
original handwritten ledger indicates that the type was re-
ceived from the U.S. Exploring Expedition, with “original
number” 9. The specimen in USNM 5409 closely matches
Gould’s (1856: pl. 1, figs. 9, 9a) figured specimen in shape,
size, whorl expansion, and faint longitudinal sculpture (Fig-
ure 11B).
pauxilla Gould, 1852; Helix
Helix pauxillus Gould, 1852:40; 1856: pl. 3, figs. 46, 46a–c; 1862:19, 243.
Current taxonomic status: Nesovitrea pauxilla (Gould, 1852).
Valid species (Cowie et al., 1995:165).
Type material: See pusilla Gould, 1846.
Type locality: See pusilla Gould, 1846.
Remarks: New name for Helix pusilla Gould, 1846, which is
a primary junior homonym of Helix pusilla Lowe, 1831.
Helix pauxilla Gould, 1852 therefore has the same name-
bearing type and type locality as Helix pusilla Gould, 1846.
pusilla Gould, 1846; Helix
Helix pusillus Gould, 1846:171; 1852:40; 1856: pl. 3, figs. 46, 46a–c (as
Helix pauxillus); 1862:19, 243.
Current taxonomic status: Nesovitrea pauxilla (Gould, 1852).
Valid species (Cowie et al., 1995:165).
Type material: Lectotype USNM 20964 (Figure 11C), here des-
ignated.
Type locality: “[M]ountains of East Maui, Sandwich Islands”
(Gould, 1852:41).
Remarks: No holotype was designated nor a figure provided
with the original description, although figures were pro-
vided subsequently by Gould (1856: pl. 3, figs. 46, 46a–
c). The locality provided in the original description was
“mountains of Maui, Sandwich Islands” and it was subse-
quently refined by Gould (1852:41). Johnson (1964:136)
noted the “Figured holotype USNM 20964” but also identi-
fied additional type material in the collections of the MCZ,
including MCZ 169288 (12 spms; ex NYSM, original no.
A781) and MCZ 169287 (3 spms; ex NYSM 240, original
no. A784). Although the original description does not imply
or require that it was based on more than one specimen, the
fact that additional type material was recognized by John-
son means this is not a valid lectotype fixation (see section
in the Introduction regarding lectotype designations). The
original handwritten USNM ledger lists an additional speci-
men, USNM 5446, with “original number” 46, identified as
Helix pauxillus” from “Sandwich I.” but notes that it was
NUMBER 647
29
FIGURE 11. A. Lectotype (USNM 5409; 11.2 mm), here designated, of Vitrina caperata. B. Gould’s (1856: pl.
1, figs. 9, 9a) figured Vitrina caperata. C. Lectotype (USNM 20964; 3.9 mm), here designated, of Helix pusilla.
D. Gould’s (1856 pl. 3, figs. 46, 46a,b) figured Helix pusilla. E. Lectotype (USNM 20874; 5.0 mm), here des-
ignated, of Vitrina tenella. F. Gould’s (1856: pl. 1, figs. 10, 10a,b) figured Vitrina tenella. Dimensions are shell
diameter (width). If more than one shell in the lot is figured, dimension is of the shell indicated by an asterisk.
Scale bars: 1 mm.
30
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
“Destroyed (Dec. ’75)” with the entry initialed “J. G. S.”
We therefore here designate USNM 20964 as the lectotype.
Although the destroyed USNM 5446 presumably was the
figured specimen, USNM 20964, with “original number”
f46, was received from the U.S. Exploring Expedition and
is a reasonable match to Gould’s (1856: pl. 3, figs. 46, 46a,
46b) figured specimen (Figure 11D), although the aperture
is damaged.
Helix pusilla Gould, 1846 is a primary junior homonym
of Helix pusilla Lowe, 1831, and was replaced by Helix
pauxilla Gould, 1852 (as pauxillus). The name-bearing type
(lectotype) of H. pusilla Gould, 1846 is therefore also the
lectotype of H. pauxilla Gould, 1852 (Code, Art. 72.7).
tenella Gould, 1846; Vitrina
Vitrina tenella Gould, 1846:181; 1852:11; 1856: pl. 1, figs. 10, 10a–c;
1862:26.
Current taxonomic status: Vitrina tenella Gould, 1846. Valid
species (Cowie et al., 1995:164).
Type material: Lectotype USNM 20874 (Figure 11E), here des-
ignated.
Type locality: “Kauai, Sandwich Islands.”
Remarks: No holotype was designated nor a figure provided with
the original description, although figures were provided sub-
sequently by Gould (1856: pl. 1, figs. 10, 10a–c). It has not
been determined that the description was based on a single
specimen, and the original description does not imply or re-
quire that it was based on more than one specimen. Johnson
(1964:53) noted a “Figured holotype USNM 20874,” thereby
inferring a “holotype.” However, he did not explicitly accept
that the original description was based on a single specimen
and, although we now know that there was additional type
material, his inference is not a valid lectotype fixation (see
section in the Introduction regarding lectotype designations).
The original handwritten USNM catalog ledger entry for
USNM 20874, with “original number” f10, states “(from di-
agnosis: type lost).” The entry for USNM 5410, with “origi-
nal number” 10, is annotated “specimen lost”; this specimen
was presumably the figured specimen. Although damaged,
USNM 20874 is a reasonable match to Gould’s (1856: pl. 1,
figs. 10, 10a, 10b) figured specimen (Figure 11F). The type
material was collected by J. P. Couthouy (Gould, 1852:12)
and the original handwritten ledger indicates that it was re-
ceived from the U.S. Exploring Expedition. We here desig-
nate USNM 20874 as the lectotype.
The taxonomic status of this species is not clear. Cooke
(1921:269) noted the same positioning of the right tenta-
cle retractor in Vitrina tenella and V. alaskana, but noted
that V. tenella was no doubt endemic to Hawaii and not
of recent introduction (Cooke, 1921:271). Similarly, Baker
(1941:322) did “not dare describe the Hawaiian species
[Vitrina tenella] as distinct from V. alasakana Dall” and
stated that there are “Apparently no specific differences be-
tween V. tenella from Hawaii and V. alaskana from Western
N.A.” He concluded that the Hawaiian species was intro-
duced by birds and that the lack of anatomical characters
differentiating the two supported the likelihood that this
was a recent event. He pointed out that tenella has priority,
clearly considering them to be synonyms although without
a formal synonymy (Baker, 1958:146). Roth and Sadeghian
(2006:62) synonymized V. alaskana with V. pellucida and,
hence, Christensen (2013) concluded that tenella is a syn-
onym of pellucida. However, none of these species has been
assessed using molecular data and we follow Cowie et al.
(1995) in considering V. tenella a valid species, pending fur-
ther research.
We thank Philippe Bouchet (Muséum national d’Histoire naturelle, Paris),
Neal Evenhuis (BPBM), and Gary Rosenberg (ANSP) for bibliographic and
nomenclatural advice, especially regarding interpretation of Art. 74 of the
Code. We thank Robert Hershler, Paul Greenhall, William Moser, and Cheryl Bright
(all NMNH, Smithsonian Institution), Carl Christensen and Regina Kawamoto (both
BPBM), Jaynee Kim, Kelley Leung, and Dylan Ressler (all University of Hawaii), Adam
Baldinger (MCZ), John Slapcinksy (University of Florida), Jonathan Ablett (NHMUK),
and Anne L’Ecuyer (Washington Writer’s Retreat) for their assistance and support. Tim
Pearce (Carnegie Museum of Natural History) and Gary Rosenberg (ANSP) provided de-
tailed reviews that greatly improved this contribution. This work was supported by Na-
tional Science Foundation grant DEB-1120906. This is contribution number 2016-011
to the Hawaii Biological Survey and number 9743 of the University of Hawaii School of
Ocean and Earth Science and Technology.
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... A second possibility of inadvertent designation of lectotype by Johnson (1989), by means of the wrong use of the term holotype (ICZN 1999: article 74.5) cannot be advocated, since Johnson (1989), when wrongly using that term ("figured holotype"), did not explicitly indicate that specimen to serve as the name-bearing type. This conclusion that this lectotype designation by Johnson (1989) is invalid is in accordance to the interpretations by Pimenta and Absalão (2004) and Yeung et al. (2017), both also regarding Johnson's designations. Yeung et al. (2017: 3) reinforced the rejection of such lectotype designation under ICZN (1999: article 74.5), adding that, when Johnson (1964) referred to the "figured holotype", he was not unambiguously selecting a specimen to serve as the name-bearing type, "because in fact no selection has occurred, as the author considers the holotype to have been fixed by the original author". ...
... This conclusion that this lectotype designation by Johnson (1989) is invalid is in accordance to the interpretations by Pimenta and Absalão (2004) and Yeung et al. (2017), both also regarding Johnson's designations. Yeung et al. (2017: 3) reinforced the rejection of such lectotype designation under ICZN (1999: article 74.5), adding that, when Johnson (1964) referred to the "figured holotype", he was not unambiguously selecting a specimen to serve as the name-bearing type, "because in fact no selection has occurred, as the author considers the holotype to have been fixed by the original author". The exact same interpretation is here taken regarding lectotypes designation by Johnson (1989). ...
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