gS rg ee NE ar
ed Dir eer

Pete
—s - Shen Tt SEN ea!
> ry o UNV Sere g
Pettey da * ‘ AN Mes
Mauss, :
etapa 3
“ v : tem t 5
tyes : On ; qe»
THT ee = Serica
Sea9 WAY Cae i : ae
eR a Ty fe eee

~
i SSN as as
NOEs es, 5 ; : ;
la whe Ae Me Aine ts 1, Hl .
SSeS ame se yo x jee,
Bot rare
Me eae

Vad i 02 wty,
Sib eee, VEN were
ent
vhvey
¢ Wards
Z Thana
Se tangs) oa fs
Ae ot ons. * ae) ;
=~ DERN Bie Be Bang ioe yy
Feild Wetter aay MR ena ete EES aon
tat te fed ye

Darya

Coe

Rhee ca
TENE Awe 6h
Beta, : ap
Sena ener 2 ;
MeeVee nt le oe sm.
; ‘ 4540 Stes :
: TS ew
: ere,
; Aad ones
; : : Cg tang,
} : Pais
; . AVIN Dey
PRAT Ue at
Pees Oras ne,
SD awenta gts
WiCanga es : :
DUCE a ezine : ANA
Seen tee Pad we
TE pe
Oat
EN arate trom,

ARIS AEE Layee,
Sas, Sots
tthe
PWR en oy .
Sete: Deer eet.
FON eR eaes
PRONE aS nysy,
Uda Seg des ;
6 Ces IN Gg oa :
Wiese Mea ay PhO nage :
SERN a ‘ ec
oe

Fer oh zis

INE EES. aug
AEN SY 35 tre

ote.

TSS eewanryy,
“ONE ea
Steere
da Deer,
dee
ORS IY oe
eee ey

Ss

A

NEMOURIA

Occasional Papers of the Delaware Museum of Natural History

NUMBER 31 ; DECEMBER 28, 1988

SPIDERS OF BERMUDA

Petra Sierwald!

ABSTRACT. Fifty-nine species from 22 spider families are currently known to occur
on Bermuda, the majority being widely distributed, cosmopolitan and cosmotropical
species. Collections of Bermudian spiders from the Peabody Museum, the Academy
of Natural Sciences of Philadelphia, and the Bermuda Department of Agriculture
& Fisheries were studied. Newly collected material (July 1983, May 1988), deposited
at the Natural History Museum of Bermuda, was incorporated. Lycosa atlantica
Marx, 1889, is a junior synonym of Trochosa ruricola (DeGeer, 1778); Anyphaena
verrilli Banks, 1902, and Oonops bermudensis Banks, 1902, are junior synonyms of
Aysha velox (Becker, 1879) and Heteroonops spinimanus (Simon, 1891) respectively.
A new anyphaenid species, Anyphaena bermudensis, is described.

INTRODUCTION

Bermuda consists of an isolated group of seven large and numerous
smaller islands, situated at 32°18'N, 64°46'W in the Atlantic Ocean, about
1000 km east-southeast of Cape Hatteras, North Carolina, USA. Due to
the nearby Gulf Stream, the climate is mild and frost-free. Annual rainfall
averages 1400 mm and is spread evenly throughout the seasons.

The spider fauna of Bermuda is known from three previous collections
by Blackwall (published 1868, nomenclature revised by Simon, 1883),
Heilprin (published by Marx, 1889) and Verrill (published by Banks,
1902). In total, twenty-eight species of spiders had been recorded from
Bermuda previously [additionally, Banks listed six unidentified species].
The present study adds twenty-six new species records. Together with one
salticid species recorded by Griswold and four unidentified species, a total
of 59 species of spiders from 22 families are known to occur on Bermuda.

An identification key to the spiders of Bermuda is not presented here.
The reader is referred to Levi (1987) for family-level determinations, and

lResearch Associate Serial Publication
Delaware Museum of Natural History ISSN 085-3887

2 NEMOURIA No. 31

to Roth (1985) for genus-level identification. Identification to species can
only be reached by comparison of copulatory organs. For this purpose, the
specialized, taxonomic literature cited under each species should be
consulted.

This paper is part of an informal series on the Bermudian fauna and
flora initiated by the Bermuda Department of Agriculture & Fisheries.

MATERIALS AND METHODS
Existing Collections of Bermudian Spiders

This study includes the following material:

1) the Heilprin collection, collected 1888; deposited in the Academy
of Natural Sciences of Philadelphia (12 species); identified and results
published by Marx, 1889 [only parts of the original Heilprin material
were found in the collection].

2) The Verrill collection, collected 1901-1902; deposited in the Peabody
Museum of Natural History, Yale University; identified and results
published by Banks, 1902 (23 species, plus six genus records based on
immature specimens).

3) Collections of the Natural History Museum of Bermuda: a)
collection of the Department of Agriculture & Fisheries [collections made
mainly by Dr. I. W. Hughes, Francis Monkman and Dr. D. J. Hilburn];
b) material collected in July 1983 and May 1988 by the author. Additional
duplicate material was deposited in the USNM as indicated.

Blackwall’s collection could not be located (Hope Entomological
Collections, Oxford, Great Britain).

Text conventions

Under each species, references are made to diagnostic figures of 2 and |

& for identification purposes. Synonyms in brackets are names used by —
other authors in previous publications on Bermudian spiders. If the species _
was originally described in a different genus, that generic name is added |

(e.g., sub Aranea). The specimen listings are ordered according to the
months (abbreviated, first three letters) in which adults were found.

Abbreviations

INSTITUTIONS:
AMNH American Museum of Natural History, New York
ANSP Academy of Natural Sciences of Philadelphia
BBS Bermuda Biological Station
BG Botanical Gardens, Paget, Bermuda
NHMB Natural History Museum, Bermuda

December 28, 1988 Sierwald 3

PMNH Peabody Museum of Natural History, Yale University, New
Haven, Connecticut

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

LOCALITY LISTINGS: FIGURE CITATIONS:
DP Devonshire Parish co copulatory organs
HP Hamilton Parish cp color pattern
PP Paget Parish ep eye pattern
PBP Pembroke Parish h habitus
SP Smith’s Parish sp spination

SAP Sandy’s Parish
SHP Southampton Parish
SGP St. George’s Parish
WP Warwick Parish
w/o without locality

a same locality as
previous sample

Each sample is indicated by a preceding ‘“**’’. “?”? and/or “‘o”’ indicate
adult females and/or males in the sample, “‘juv.” indicates juveniles, “‘sa”’
indicates subadults. New Bermuda records are indicated by “M@’’ in front
of species name.

LIST OF BERMUDIAN SPIDERS

Families and genera are listed alphabetically.

Agelenidae

Tegenaria domestica (Clerck, 1758), sub Araneus
[= Tegenaria derhami (Scopoli, 1763), see Roth, 1956]

* HP: Walsingham, May 1901, PMNH 2327 ¢. * w/o, PMNH 2326 2.
Distribution: All species of Tegenaria are considered cosmopolitan (Roth,
1968: 4). Figures: Roberts, 1985, 1: figs. 72b (co ?0°); 3, plate 92 (h).

Anyphaenidae

M@ Anyphaena fraterna (Banks, 1896), sub Gayenna

* SGP: Ferry Reach, BBS, garden, Jul 1983, NHMB 2.
Distribution and figures: North America: southern New York west to
eastern Kansas, south to western Florida and eastern Texas; Platnick, 1974:
234, figs. 52, 56, 60, 77, 78 (co 0’, he’).

4 NEMOURIA No. 31

M Anyphaena bermudensis n. sp.
* SAP: Ft. Scaur, Jan 1988, NHMB ? (paratype IX). * SAP: Heydon
Trust, sweeping, Jan 1988, USNM 2 (holotype), USNM @ (paratype
I). * PP: Point Finger Road, garden, at night, May 1988, AMNH 2
(paratype II). * HP: Walsingham jungle, sweeping, Jul 1983, NHMB
o” (paratype III), NHMB ¢ (paratypes IV, V, VI). * SP: Spittal Pond
Park, Oct 1987, NHMB juvenile. * PBP: Admiralty House Park,
sweeping, Oct 1987, NHMB 2 (paratype VIII). * PP: Camden, BG,
sweeping, Dec 1987, AMNH 2 (paratype VII).

Distribution: Known from Bermuda only. Figures: 1-5. See description

below.

Aysha velox (Becker, 1879), sub Anyphaena
[= Anyphaena verrilli Banks, 1902] NEW SYNONOMY
* SAP: Ft. Scaur, sweeping, Jan 1988, NHMB ¢. * HP: Walsingham,
May 1901, PMNH 2346 ? (holotype of Anyphaena verrilli). * SGP:
Tucker’s Town, sweeping, Jul 1983, NHMB juv. [Aysha cf. velox].
* PP: Berry Hill Road, in house, Oct 1987, NHMB 2. * several
juveniles from other Parishes.
Distribution and figures: North America and West Indies: North Carolina
west to Arkansas, south to eastern Texas and Florida; Cuba, Haiti,
Dominican Republic and Bermuda; Platnick, 1974: 259, figs. 110, 111,
122, 125 (co 2°’).
Note: Banks (1902: 270) mentioned that he had “... seen specimens [of
“‘Anyphaena verrillv’| from parts of the West Indies.”

Araneidae

Cyclosa turbinata (Walckenaer, 1841), sub Epezra

[= Cyclosa caudata Hentz, 1850]
* SHP: Horseshoe Bay, sweeping, Jan 1988, NHMB 2. * HP: Shelly
Bay, Mar 1988, sweeping, NHMB 2. * PP: Hinson’s Island, in web
in orchard, Apr 1969, NHMB ¢?o¢. * HP: Walsingham jungle, May
1901, PMNH 2329 ¢, csa. * May 1901, PMNH 2350 ¢.
* SAP: [Tucker’s Island Cave, now part of the U.S. Naval Annex],
May 1901, PMNH 2328 juv. * PMNH 2358 juv. * HP:
Walsingham jungle, on webs, Jul 1983, NHMB 20, juv. * SGP:
Tucker’s Town, sweeping, Jul 1983, NMHB ?c. * —— web in
mangroves, Jul 1983, NHMB juv. * HP: Flatts Village, on web, Jul
1983, NHMB °. * SGP: Hill Park, among rocks, Jul 1983, NHMB
?o°, * PP: Hinson’s Island, Jul 1969, NHMB ?o. * SP: Spittal Pond
Park, sweeping, Oct 1987, NHMB ¢. * SGP: Ferry Point Park,
sweeping, Oct 1987, NHMB ~. * HP: Shelly Bay, mangrove swamp,

December 28, 1988 Sierwald 5

Nov 1987, NHMB 20. * w/o, PMNH 2339 juv. * w/o, Heilprin
Coll., USNM @°.
Distribution and figures: North and Central America, West Indies; Levi,
1977: 82, figs. 20, 38-50 (co, cp, h ?¢’).

@ Eustala anastera (Walckenaer, 1841), sub Epezra
* SGP: Smith’s Island, May 1988, NHMB ?. * SGP: Tucker’s Town
Grove, Jul 1983, NHMB 8. * SP: Spittal Pond Park, NHMB juv.
Distribution and figures: North America: widely distributed; Levi, 1977a:
115, figs. 205-232, 280-285, 298-302, 314-315 (co, cp, h 20; variability of
CO).

Mi Gasteracantha cancriformis (Linné, 1767), sub Aranea
* SGP: Nonsuch Island, Feb 1967, NHMB ?. * PBP: Tulo Valley,
in web, May 1969, NHMB ~~. * PP: Camden, BG, May 1988,
NHMB @. * PP: Hinson’s Island, in web in cherry hedge, Jun 1969,
NHMB ¢. * SGP: Ferry Reach, BBS, garden, Jul 1983, NHMB 2.
* SGP: Hill Park, among rocks, Jul 1983, NHMB juv. * HP: Flatts
Village, sweeping in grass and shrubs, Jul 1983, USNM 0. * SGP:
Tucker’s Town Grove, Jul 1983, NHMB °. * SAP: Sep 1969,
NHMB @. * SAP, Shrewsbury, in web in banana patch, Sep 1969,
NHMB ¢’sa. * PP: Wreck Hill, Dec 1965, NHMB °. * PP: Point
Finger Road, garden, May 1988, NHMB juv. * several juveniles from
different locations throughout Bermuda.
Distribution and figures: North and South America: North Carolina west
to southern California, south to northern Argentina; Levi, 1978: 441, figs.
69-84 (co, cp, h 0). Note: In 1969, 1983 and 1988, Gasteracantha
cancriformis was found to be common and occurring in various habitats
on the islands (forests, mangroves, tall shrubs and grass, banana plantations,
gardens and among rocks). The species was first reported in 1934 in records
of the Bermuda Department of Agriculture & Fisheries.

Metepeira labyrinthea (Hentz, 1847), sub Epeira

* Heilprin Coll. [not in ANSP, non vidi].
Distribution and figures: widely distributed in eastern United States,
Ontario, Canada, south to Florida; Levi, 1977b: 198, figs. 14-20 (co, cp,
ep, h 2’),

cf. Neoscona moreli (Vinson, 1863)
* w/o, Blackwall Coll., det. by Blackwall as Epeira gracilipes Blackwall,
1862 [non vidi].

Note: Banks (1902: 267) suggested that Blackwall’s species Epeira gracilipes

6 NEMOURIA No. 31

is a synonym of “‘Araneus theis (Walckenaer, 1841)”. Grasshoff (1986: 69)
pointed out that Neoscona theisi occurs only in Asia (Malaysia, China, Japan
and Pacific Islands). The specimen collected by Blackwall might have been
a Neoscona moreli (Vinson, 1863). Grasshoff (1986: 56, figs. 71-78)
suggested that this widespread African species had been transported by
ships from West Africa to the West Indies. It is known to occur on the
Antilles. In the past, Neoscona moreli has often been misidentified as
Neoscona theisi (see citations in Grasshoff).

Clubionidae

Mi Clubiona reclusa O. Pickard-Cambridge, 1863

* SGP: Ferry Reach, BBS, garden, Jul 1983, NHMB 2.
Distribution: Northern and Middle Europe, Siberia (Roewer, 1954, 2a:
499). Figures: Roberts, 1985, 1: 30b (co 2°’).

HM Corinna abnormis Banks, 1930

* HP: Bailey’s Bay, in house, NHMB ¢.
Distribution and figures: previously only known from the male type
specimen from Puerto Rico [non vidi]; Petrunkevitch, 1930: 103, fig. 86
(co &).

Mi Corinna humilis (Keyserling, 1887), sub Hypsinotus
* PP: Camden, BG, under bark of Australian Pine (Casuarina
equisetifolia L.), May 1988, NHMB ?; USNM 3. * SHP: in grass,
Dec 1969, NHMB 2. * w/o, PMNH 2309 2, sa.

Distribution and figures: widely distributed in the West Indies; Petrunkevitch,

1930: 95, figs. 78, 79 (co 20°; vulva not figured).

Desidae

Paratheuma insulana (Banks, 1902), sub Eutichurus
* w/o, May 1901, PMNH 2362 ¢? (holotype). * SGP: Whalebone
Bay, in rock crevices above hightide line at shore, May 1988, NHMB
ae: Jul 1983, NHMB ¢.
Distribution and figures: West Indies: Cuba, Haiti; Bermuda, Florida
Keys; Platnick, 1977: 200, figs. 3, 4 (co ?); Beatty and Berry, 1988: 50,
figs. 1, 4, 7, 10 (co “?); Banks, 1902: fig. 3 (? epigynum).

Dictynidae

@ Dictyna altamira Gertsch & Davies, 1942
* DP: Middle Road, May 1988, USNM 20. * PP: Camden, BG,
May 1988, NHMB ?. * SGP: Whalebone Bay, rocks along shore, |
May 1988, NHMB ¢. * SGP: Tucker’s Town, sweeping shrubs, Jul |

December 28, 1988 Sierwald 7

1983, NHMB 3. * SGP: Ferry Reach; BBS, hedge, sweeping, Jul
1983, NHMB 2. * SP: Spittal Pond Park, Nov 1987, sweeping,
NHMB ¢.

Distribution and figures: North America and West Indies: widely distrib-
uted in eastern United States, Mexico and the West Indies; Chamberlin
& Gertsch, 1958: 117, figs. 1-4 (co ?¢).

Dysderidae

Dysdera crocata C. L. Koch, 1839
* PP: Camden, in rain catch bottle, BG, Mar 1966, NHMB 2. * SP:
Spittal Pond Park, May 1988, USNM ?. * SGP: Smith’s Island,
May 1988, USNM ¢. * HP: Walsingham, May 1901, PMNH 2347
juv. * SGP: Tucker’s Town, under stones, Jul 1983, NHMB juv.
* SP: Spittal Pond Park, under rocks, Jul 1983, NHMB 8 juv. * PP:
in house, Nov 1987, NHMB @. * w/o, PMNH 2308 8, °, juv. *
PMNH 2376 ¢. * PMNH 2371 2. * Heilprin Coll., ANSP,
several females, poor condition.

Distribution: cosmopolitan, widely distributed in Europe, Asia and the

Americas (Roewer, 1942: 296). Figures: Roberts, 1985, 1: figs. 19b, d, f,

h (co ?0°, sp femur).

Filistatidae

Kukulcania hibernalis (Hentz, 1842), sub Filistata

[= Filistata depressa (C.L. Koch, 1842)]
* HP: Walsingham, May 1901, PMNH 2322 ¢ [non vidi]. * SAP:
Tucker’s Island Cave, May 1901, under stones, PMNH 2321 2sa.
* PBP: Mount Hill, night, outside on wall, Jul 1969, NHMB 2.
* SGP: Ferry Reach, BBS, in house, Jul 1983, NHMB @. * w/o,
PMNH 2331 sa. * PMNH 2324 ¢. * PMNH 2332 juv.
* w/o, Blackwall Coll. [non vidi]. * several juveniles from various
localities.

Distribution: North and South America (Roewer, 1954, 2b: 1281). Figures:

Comstock, 1910: fig. 1 (co ); Lehtinen, 1967: fig. 19 (co <). Note: Lehtinen

(1967: 242) transferred all American species formerly assigned to Filistata

to a new genus Kukulcania Lehtinen, 1967. Banks (1902: 267) synonymized

F. depressa with hibernalis.

Gnaphosidae

Mi Camellina elegans (Bryant, 1940), sub Eilicina
* SGP: Tucker’s Town, under rocks, Jul 1983, NHMB ?. * SGP:
Whalebone Bay, under rocks, Jul 1983, NHMB juv.

Distribution and figures: North and Central America, Pacific, Africa:

8 NEMOURIA No. 31

Florida south to Curacao, Hawaii, Marshall Islands, Angola; Platnick &
Shadab, 1982: 6, figs. 1-4 (co 0’).

Callilepis sp.
* SAP: Tucker’s Island Cave, May 1901, PMNH 2356 ?sa (specimen
dessicated and damaged.

Note: Revision of the genus Callilepis by Platnick, 1975.

Linyphiidae
This large family (150 genera, 872 species; Roth, 1988) is in need of
revision. Identification is hampered and nomenclatural problems exist.

M@ Bathyphantes approximatus (O. Pickard-Cambridge, 1871), sub Linyphia
* WP: Warwick Pond, May 1988, NHMB 8. * SP: Spittal Pond Park,
under rocks, May 1988, USNM @°. * SP: Spittal Pond Park, under
rocks, Jul 1983, NHMB ?. * w/o, PMNH 2344 ¢. * PMNH
2341 sa. * PMNH 2338 &sa.

Distribution: Europe, Siberia (Ivie, 1969: 54; Roewer, 1942: 596). Figures:

Roberts, 1985, 3: plate 218; 1987: fig. 70a (co, cp 20"); Ivie, 1969: figs.

102-104. Note: Roewer (1942: 596) listed this species in the genus

Stylophora Menge, 1899.

MM Erigone autumnalis Emerton, 1882

* PP: Camden, BG, May 1988, NHMB ¢.
Distribution: throughout the United States (Roth, 1988). Figures: Kaston,
1981: pl. XXVIII figs. 616-618 (co, h @).

MM Erigone brevidentata Emerton, 1909
* PP: Camden, Dept. Agriculture, on floor in lab, Jul 1970, NHMB
%. * SP: Spittal Pond Park, sweeping, Nov 1987, USNM 2.
Distribution: eastern United States (Roth, 1988). Figures: Kaston, 1981:
pl. XXIX fig. 628, pl. XXXI figs. 662-663 (co, h @’).

Mi Erigone cf. promiscua (O. Pickard-Cambridge, 1872), sub Neriene

* w/o, PMNH 2335 ¢
Distribution: western Europe (Roewer, 1942: 720). Figures: Roberts, 1987:
figs 43b, 45c (co 20"). Note: The epigynum shows resemblance to Roberts’
figures of Erigone promiscua.

Mi Lepthyphantes leprosus (Ohlert, 1865), sub Linyphia

* SGP: Tucker’s Town, sweeping shrubs, Jul 1983, NHMB ?.
Distribution: holarctic: Europe, Siberia, North America (Roewer, 1942:
549). Figures: Zorsch, 1937: figs. 7-13 (co ?o); Roberts, 1985, 3: plate
229a, b; 1987: fig. 77b (co 2°, h 20’).

December 28, 1988 Sierwald 9

Mi Lepthyphantes obscurus (Blackwall, 1841), sub Linyphia

* SP: Spittal Pond Park, forest, on web, Jul 1983, NHMB ¢.
Distribution: Europe (Roewer, 1942: 552). Figures: Roberts, 1985, 3: plate
229d; 1987: fig. 78b (co, h ?’).

Loxoscelidae

Loxosceles rufescens (Dufour, 1820), sub Scytodes
* PP: Camden, Dept. Agriculture, laboratory, May 1988, NHMB ¢@.
* SGP: Ferry Reach, BBS, in house, Jul 1983, NHMB ?. * w/o,
Blackwall Coll., as Scytodes pallida Blackwall, 1865 [non vidz].
Distribution and figures: cosmopolitan: widespread in Europe, North
America, and Africa; of European-north African origin; Roewer, 1942:
319; Gertsch & Ennik, 1983: 354, figs. 341-343, 348-351 (co 20’, h?).

Lycosidae

Mi Gladicosa gulosa (Walckenaer, 1837), sub Lycosa
* Parish?: Outer Island [location unknown], Coll. Verrill 1902, PMNH
without number 2°.
Distribution and figures: southern Canada south to eastern Texas; Brady,
1986: 298, figs. 4, 6-9, 35, 36 (co, cp, h 2°’).

@ Lycosa cf. lenta-group

* w/o, G. Brown Goode leg., 1876-1877, PMNH 2304 c’sa.
Distribution and figures: spiders of the Lycosa lenta-group are widely
distributed in south-eastern United States; Wallace, 1942, 25 figs. (co ?¢’).
Note: Banks identified this specimen as Lycosa atlantica [= Trochosa
ruricola|. The specimen is too large to be conspecific with T. ruricola.
The black venter of the specimen allows the recognition of the species

group.

Trochosa ruricola (DeGeer, 1778), sub Aranea

[= Lycosa atlantica Marx, 1889] NEW SYNONYMY
* PP: Hungry Bay, Apr 1901, PMNH 2351 juv. * SAP: Tucker’s
Island Cave, May 1901, PMNH 2357 juv. * HP: Walsingham, May
1901, PMNH 2372 ¢&, ? damaged. * SGP: Smith’s Island, May 1988,
USNM 20, * SP: Spittal Pond Park, May 1988, NHMB 2. * SGP:
BBS, pitfall trap, Jul 1983, NHMB ¢ with egg sac. * SGP: Great
Head Park, forest litter, Jul 1983, NHMB @, ? with egg sac. * SGP:
Whalebone Bay, pitfall trap, Jul 1983, NHMB ¢, juv. * Parish?: Pen
Slow’s Cave [location unknown], PMNH 2375 ?. * w/o, under stones,

10 NEMOURIA No. 31

Heilprin Coll., ANSP 2 ? (syntypes Lycosa atlantica). * w/o, PANH

2307 ?, ¢sa. * PMNH 2325 ¢ with egg sac. * PMNH

2369 o, ?sa, juv. * PMNH 2405 ¢? adult. * w/o, USNM 32.
Distribution: palearctic (Roewer, 1954, 2a: 299). Figures: Roberts, 1985,
1: figs. 62c, 63a (co $0); Marx, 1889: fig. 4 (epigynum).
Note: The Trochosa specimens found on Bermuda are conspecific with
Trochosa ruricola of Europe. Aside from the structure of the copulatory
organs, a broad, low tooth on the fang of the male’s chelicerae (Roewer,
1928: 130) serves as an identification character. Roewer (1954, 2a: 247)
transferred Lycosa atlantica (without examination of type material) to a new
genus Hoggicosa Roewer, [1954 nom. nud.]| 1960 [1960: 772].

Oonopidae

Heteroonops spinimanus (Simon, 1891), sub Oonops
[= Oonops bermudensis Banks, 1902] NEW SYNONYMY

* w/o, PMNH 2340 & (holotype).
Distribution and figures: West Indies and northern South America, Florida;
Petrunkevitch, 1929: 67, figs. 53-57 (co ?o, ep, h, ? palp); Chickering,
1969: 156, figs. 28-32 (co, ep, spination 20°); Banks, 1902: figs. 1 a-c (ep,
co, leg 2).

M Orchestrina sp.
* SP: Spittal Pond Park, May 1988, NHMB juv.
Figures: Chickering, 1969; figs. 35-39 (ep).

Oxyopidae

Oxyopes salticus Hentz, 1845
* SP: Spittal Pond Park, sweeping, Jan 1988, NHMB 2c’. * SHP:
Horseshoe Bay, sweeping, Jan 1988, NHMB ?c’. * SP: Spittal Pond
Park, sweeping, Feb 1988, NHMB ~~. * SGP: Whalebone Bay,
sweeping grass, Jul 1983, NHMB ?o°. * SGP: Ferry Point Park,
sweeping grass, Jul 1983, NHMB 20°. * SGP: Great Head Park,
sweeping grass, Jul 1983, NHMB o. * SP: Spittal Pond Park,
sweeping grass, Jul 1983, NHMB ?; USNM 2c’. * HP: Flatts Village,
sweeping grass, Jul 1983, NHMB 2c, * SP: Spittal Pond Park,
sweeping, Oct 1987, NHMB ~. * PP: Camden, BG, sweeping, Dec
1987, NHMB 2. * w/o, PMNH 2345 juv.

Distribution and figures: North America: distributed throughout the United

States; Brady, 1964: 482, figs. 80-86, 91-96, 104-105 (co, cp, h, ep ?°’).

December 28, 1988 Sierwald 11

Pholcidae

Mi Pholcus phalangioides (Fuesslin, 1775), sub Aranea
* SAP: Tucker’s Island Cave, 3 May 1901, PMNH 2315 < (det. by
Banks as Pholcus tipuloides L. Koch, 1872).
Distribution: cosmopolitan (Roewer, 1942: 338; Emerton, 1902: 128).
Figures: Roberts, 1985, 1: figs. 21a; 3: plate 18 (co, cp, h ?’)

Smeringopus elongatus (Vinson, 1863), sub Pholcus

[= Pholcus tipuloides L. Koch, 1872]
* PP: Camden, in old building, Jan 1965, NHMB o, * Jan
1966, NHMB co. * PBP: BHSG-Gymnasium, on wall, Mar 1988,
NHMB 2. * SGP: Whalebone Bay, rocks close to shore, May 1988,
NHMB 3°. * SGP: Ferry Reach, BBS, in house, Jul 1983, NHMB °°;
USNM 20, * SGP: Tucker’s Town, on ground, NHMB sa. * SAP:
Tucker’s Island Cave, PMNH 2320 2 with egg sac. * —— PMNH
2361 ¢ damaged. * w/o, Heilprin Coll., ANSP sa.

Distribution: cosmotropical (Roewer, 1942: 343). Figures: Petrunkevitch,

1929: figs. 136-138; Marx, 1889: figs. 5a-b (co, ep, h @).

Salticidae

HM cf. Metaphidippus sp.
* SP: Spittal Pond Park, sweeping, Jan 1988, NHMB 8. * HP: Shelly
Bay, sweeping, Mar 1988, NHMB 8. * SP: Spittal Pond Park,
sweeping, Nov 1987, NHMB 32.
Note: Specimens are members of the Dendryphantinae. Specific identifica-
tion is currently not possible; the group is under revision (Griswold, pers.
comm.).

Mi Habronattus coecatus (Hentz, 1846)
* SAP: Heydon Trust, sweeping, Jan 1988, NHMB ¢. * DP:
Devonshire Marsh, Middle Road, May 1988, NHMB ~~. * SP:
Somerset, Jul 1971, AMNH © [non vidi]. * PP: Berry Hill Road, Nov
1987, NHMB sa. * several immature specimens from various
localities.
Distribution and figures: Eastern United States from New York to northern
Florida, west to edge of the Great Plains, north eastern Mexico; Griswold,
1987: 97, fig. 79 (¢ leg).

12 NEMOURIA No. 31

Habronattus nesiotus Griswold, 1987

* w/o, Jul 1905, MCZ ¢ (holotype) [non vidi].
Distribution and figures: known only from male holotype from Bermuda;
Griswold, 1987: 227, figs. 37, 174, 222 (cp, co ¢).

Hentzia vernalis (Peckham, 1893), sub Anoka

[= Wala vernalis]
* PP: Paget Marsh, sweeping, Jan 1988, NHMB o’sa. * SHP:
Horseshoe Bay, sweeping, Jan 1988, NHMB c’sa. * SP: Penhurst
Peak, sweeping, Jan 1988, USNM -.. * SAP: Heydon Trust, sweeping,
Jan 1988, NHMB ?. * SAP: Gilberts Nature Preserve, May 1988,
NHMB 2. * SP: Spittal Pond Park, May 1988, NHMB 32. * SAP:
Somerset Island, railway path, sweeping shrubs, Jul 1983, USNM 2.
* SAP: Ft. Scaur, sweeping, Nov 1987, NHMB ¢. * HP: Flatts
Village, Aquarium, Nov 1987, NHMB §°. * several immature speci-
mens from various localities.

Distribution and figures: common on Haiti, Puerto Rico, St. Vincent,

Cuba; Petrunkevitch, 1930: 139, figs. 120-129 (co, cp, h 2c’).

Menemerus bivittatus (Dufour, 1831), sub Salticus
[= Menemerus melanognathus (Lucas, 1838)]
* DP: south shore under stones, Dec 1968, NHMB 2, ¢sa. * w/o,
Heilprin Coll., ANSP ¢.
Distribution: cosmopolitan (Roewer, 1954, 2b: 1263). Figures: Petrunkevitch,
1930: figs. 165-168, Marx, 1889: figs 3a-b (co, cp 2°).

Plexippus paykulli (Audouin, 1827), sub Attus

[= Salticus diversus Blackwall, 1868]
* SGP: Ferry Point Park, under stones, Jul 1983, NHMB 29 in nest.
* SAP: Somerset Island, Black Bay, under stones, Jul 1983, NHMB
?. * SGP: Whalebone Bay, under rocks, Jul 1983, NHMB 2, juv.
* HP: Walsingham, PMNH 2373 juv. * SAP: Tucker’s Island Cave,
PMNH 2374 juv. * w/o, PMNH 2310 2. * PMNH 2311 ?
with nest. * PMNH 2312 ¢, * PMNH 2336 juv. *
PMNH 2337 damaged, juv. * PMNH 2359 2. * PMNH
2370 253. = Dec 1901, PMNH 2402 ¢. * Heilprin Coll.
(sub Menemerus) [not in ANSP, non vidi]. * Blackwall Coll. (¢,
osa, type of Salticus diversus) [non vidi].

Distribution: cosmopolitan (Roewer, 1954, 2b: 1086). Figures: Petrunkevitch,

1930: figs. 140-143; Marx, 1889: figs. 2a-b (co, cp, h 2°’; vulva not figured).

Note: Banks (1902) synonymized Salticus diversus Blackwall, 1868, with

Plexippus paykullt.

December 28, 1988 Sierwald 13

Scytodidae

Scytodes fusca Walckenaer, 1837
* HP: Jan 1976, NHMB 2. * PBP, Mount Hill, in house, Apr 1966,
NHMB 3. * Apr 1969, NHMB °. * SGP: Ferry Reach, BBS,
in house, Jul 1983, USNM @.

Distribution: cosmotropical (Roewer, 1942: 323). Figures: Valerio, 1981:

figs. 1, 10, 19 (co, cp 2°).

Scytodes longipes Lucas, 1845
* HP: under building, Jan 1970, NHMB 2c’. * PBP: Hamilton, under
ridge, Jan 1976, NHMB 2c. * SGP: Smith’s Island, May 1988,
USNM ?. * SHP: Seymour’s Pond, under tree trunk in woods, May
1988, NHMB ?. * SAP: Tucker’s Island Cave, May 1901, PMNH
2319 &. * PBP: Mount Hill, in house, Jul 1969, NHMB ? with
spiderlings. * w/o PMNH 2343 juv. * w/o PMNH 2360 ¢.

Distribution: cosmopolitan (Roewer, 1942: 323). Figures: Roberts, 1985,

1: 18c; 3: plate 14; Petrunkevitch, 1929: figs. 99, 100 (co, cp ?c’).

Sparassidae

Heteropoda venatoria (Linné, 1767), sub Aranea
[= Olios antillianus Walckenaer, 1837]
* SGP: BBS, in house, Jul 1983, NHMB @. * w/o, Dec 1901,
PMNH 2398 ¢. * PMNH 2317 2. * PMNH 2342 juv. *
— Heilprin Coll. ANSP juv. * —— Blackwall Coll., as Olios
antillianus [non vidi].
Distribution: cosmotropical (Roewer, 1954, 2a: 712). Figures: Jaérvi, 1914:
plate 8, figs. 3-6; text-fig. 61 (co ?); Pickard-Cambridge, 1905, II: pl VII,
figs. 22a-c, 23 a-f [non vidi]. Note: Hilburn (pers. comm.) has frequently
observed H. venatoria in bee hives on Bermuda.

Tetragnathidae

Leucauge, Metellina and Nephila are placed in Tetragnathidae (instead of
Araneidae) after Levi & Coddington (pers. comm.).

Leucauge venusta (Walckenaer, 1841), sub Epeira

[= Argyroepeira hortorum Hentz, 1847)
* HP: Walsingham woods, May 1901, PMNH 2330 ¢. * SGP: Ferry
Reach, BBS, garden, Jul 1983, NHMB ¢. * HP: Walsingham woods
May 1901, PMNH 2348 juv. * w/o, PMNH 2364 juv. * PMNH
2365 juv. * PMNH 2366 juv.

Distribution and figures: North and Central America: New Hampshire

west to South Dakota, south to Panama; coast of California; Levi, 1980:

28, figs. 44-59.

14 NEMOURIA No. 31

Mi Metellina menge (Blackwall, 1869), sub Epezra

* SGP: Ferry Reach, BBS, garden, Jul 1983, NHMB ?¢.
Distribution and figures: Europe; Levi, 1980: 38, figs. 105-107 (co 2);
Roberts, 1985, 1: figs. 90b,g, 91b; 3: plate 133, 134 (co, cp, h $0, sp
metatarsus, tarsus).

Nephila clavipes (Linné, 1767), sub Aranea
* SGP: Ferry Reach, BBS, garden, Jul 1983, NHMB @ and juv.
* DP: Arboretum, Sep 1969, NHMB ¢. * w/o, PMNH 2399 2.
* SP: Spittal Pond Park, forest, Jul 1983, NHMB juv. * w/o, Heilprin
Coll. [not in ANSP, non vidi].
Distribution and figures: North and South America: southeastern United
States south to northern Argentina; Levi, 1980: 23, figs. 23-43 (co, cp, h
90"),

M Tetragnatha laboriosa Hentz, 1850
* SGP: Stokes Point Nature Preserve, Jan 1988, NHMB ?¢. * SGP:
Ferry Reach, sweeping, Jan 1988, NHMB @. * SP: Spittal Pond Park,
sweeping, Jan 1988, NHMB 22. * HP: Shelly, Bay, sweeping, Mar
1988, NHMB 2. * DP: Devonshire Marsh, sweeping, May 1988,
NHMB 22’. * SP: Spittal Pond Park, May 1988, NHMB ?¢, USNM
?o°, * SGP: Ferry Reach, BBS, meadow, sweeping, Jul 1983, NHMB
Son hedge, sweeping, Jul 1983, NHMB $c. * HP: Walsingham,
sweeping, Jul 1983, NHMB juv. * HP: Shelly Bay, mangrove swamp,
Nov 1987, NHMB 2. * SP: Spittal Pond Park, sweeping, Nov 1987,
NHMB 2?¢. * SP: Spittal Pond Park, Nov 1987, sweeping, NHMB
?o°, * PP: Camden, BG, Dec 1987, sweeping, NHMB ?¢.

Distribution and figures: widespread, Alaska to Panama; Levi, 1981: 308,

figs. 16-22, 120-128, pl. 6 figs. h,i.

M@ Tetragnatha versicolor Walckenaer, 1841

* SGP: Ferry Reach, BBS, garden, Jul 1983, NHMB ?c¢’.
Distribution and figures: Alaska to Nicaragua and Cuba; Levi, 1981: 304,
figs. 90-109, pl. 3 figs. upper left and bottom, pl. 6 figs. a-f.

Theridiidae

Achaearanea tepidariorum (C.L. Koch, 1841), sub Theridium
* w/o, Dec 1901, PMNH 2401 2. * PMNH 2407 ¢. * Parish?:
Outer Island [location unknown], PMNH without number ?. * w/o,
Heilprin Coll., ANSP ?¢’.

Distribution and figures: cosmopolitan; Levi, 1955: 33, figs. 69-70, 83-84;

Roberts, 1985, 1: fig. 81a.

December 28, 1988 Sierwald 15

Anelosimus studiosus (Hentz, 1850), sub Theridion
* HP: Walsingham, May 1901, PMNH 2349 ¢. *
eggs and parts of nest, NHMB 3.
Distribution and figures: widely distributed in North and South America;
Levi, 1956: 420, figs. 21-23, 37-39 (co, cp, h ?¢).

Jul 1983, with

Argyrodes nephilae Taszanowski, 1872
* PP, Hinson’s Island, in “‘crab spider’ [= ? Gasteracantha] web,
Apr 1969, NHMB ~’. * HP: Flatts Village, sweeping grass and shrubs,
Jul 1983, NHMB °. * HP: Walsingham jungle, Jul 1983, NHMB 2,
&%. * SGP: Tucker’s Town, Jul 1983, NHMB ~. * SGP: Ferry Reach,
BBS, sweeping hedge, Jul 1983, NHMB ?o. * SGP: Hill Park, among
rocks, Jul 1983, NHMB ¢@. * SP Spittal Pond Park, forest, Jul 1983,
?. * SGP: Nonsuch Island, in web in low shrubs, Jul 1969, NHMB
¢¢, * SAP: Shrewsbury, in Nephila web, Sep 1969, NHMB 22.
* HP: Shelly Bay, mangrove swamp, Nov 1987, USNM $0. * w/o,
Heilprin Coll. [not in ANSP, non vidi].

Distribution and figures: Eastern Florida, Caribbean, South America;

Exline & Levi, 1962: 141, figs. 133-137 (co, h 2c’).

Mi Argyrodes trigonum (Hentz, 1850), sub Theridion

* SGP: Ferry Reach, BBS, garden, Jul 1983, NHMB 3.
Distribution and figures: eastern United States, Ontario, Canada, south
to Florida; Exline & Levi, 1962: 124, figs. 66-78 (co, h 2c’).

Mi Coleosoma floridana Banks, 1900
* SAP: Heydon Trust, sweeping, Jan 1988, NHMB 3°. * PP: Camden,
BG, Feb 1966, USNM °°; NHMB co. * SP: Spittal Pond Park,
May 1988, NHMB 3. * PP: Point Finger Road, garden, night, May
1988, NHMB ¢. * SGP: Tucker’s Town, sweeping shrubs, Jul 1983,
NHMB ?. * SGP: Whalebone Bay, under rocks, Jul 83, NHMB 2.
Distribution and figures: cosmotropical, probably distributed by man;
Levi, 1959b: 8, figs. 12-17.

Latrodectus geometricus C. L. Koch, 1841
* HP: Paynter’s Vale, Apr 1901, PMNH 2352 o. * PMNH 2353
?. * SHP: Zero, Oct 1968, NHMB 8. * w/o, PMNH 2323 2.
Distribution: cosmopolitan (Roewer, 1942: 425). Figures: Levi, 1959a: figs.
8-10, 28, 37 (co, h $0"). Note: Levi (1983) pointed out that there are
unresolved difficulties in the discrimination of Latrodectus species.

16 NEMOURIA No. 31

M@ Theridion chetmatos Gertsch & Archer, 1942
* SGP: Ferry Reach, BBS, sweeping meadow, Jul 1983, NHMB 2,
USNM 3.
Distribution and figures: southeastern United States; Levi, 1957: 98, figs.
335, 336, 354-357 (co, cp $c’).

Theridion rufipes Lucas, 1849
* PP: Camden, BG, Feb 1966, NHMB ?¢. * PBP: Feb 1988, NHMB
?. * PP: Camden, Dept. Agriculture, in laboratory, Mar 1988,
NHMB 8. * SAP: Tucker’s Island Cave, May 1901, PMNH 2354 2.
* PP: Berry Hill Road, in house, May 1988, NHMB 0. * SGP:
Ferry Reach, BBS, in house, Jul 1983, NHMB ¢?. * PP: Berry Hill
Road, in house, Oct 1987, NHMB ¢, USNM 2c. * PP: Berry Hill
Road, in house, Nov 1987, NHMB 3°.

Distribution and figures: cosmotropical; Levi, 1957: 58, figs. 188-193 (co

gon)

Mi Thymoites marxi (Crosby, 1906), sub Pazdisca

* SP: Spittal Pond Park, May 1988, NHMB ?.
Distribution and figures: eastern United States to central Mexico; Levi,
1957: 111, figs. 393-395 (co ?o). Note: Generic placement of marxi in
Thymoites is according to Levi (pers. comm.).

Thomisidae

Thomisus pallens Blackwall, 1868

* w/o, Blackwall Coll., 1 ?sa (holotype), lost [zon vidi].
Note: Roewer (1954, 2b: 1690) lists this species as nomen dubium (“‘nicht
zu deuten’’).

M Xysticus ferox (Hentz, 1847), sub Thomisus
* Parish?: Outer Island [location unknown], 1902, Verrill coll., PMNH
without number @.
Distribution and figures: Alaska to Nova Scotia, south to Utah, Texas and
Georgia; Turnbull et al., 1965: 1251, figs. 44, 47, 126, 129, 175 (co, cp ?¢).

Uloboridae

Zosis geniculatus (Olivier, 1789), sub Aranea
[= Uloborus zosis Walckenaer, 1841]
* w/o, PMNH 2404 2, o'sa, juv. *
Heilprin Coll. [not in ANSP, non vidi].
Distribution and figures: cosmopolitan; Opell, 1979: 510, figs. 443-549;
Marx, 1889: figs. la-c (co, h, cp, ep ?’).

PMNH 2406 &. * w/o,

December 28, 1988 Sierwald 17

DESCRIPTION OF ANYPHAENA BERMUDENSIS na. sp.
(Figures 1-5)

Female (7 specimens): overall color of animal pale yellow; head and
chelicerae, maxillae and labium darkened to light brown; black eye pigments
clearly visible. Life animal: prosoma and opisthosoma with green pattern
(Fig. 4); pattern fading in alcohol.

COPULATORY ORGANS: epigynum with central atrium-like area, copulatory
Openings in anterior part of epigynum; copulatory ducts wide; spermathecae
at posterior end of vulva.

MEASUREMENTS (in mm): holotype (USNM,): total length [tl] 3.08, prosoma
length [prosl] 1.32, prosoma width [pw] 0.92. Legs: I: Femur [Fe] 1.76,
Patella-Tibia [PaTi] 2.6, Metatarsus-Tarsus [MeTa] 2.4, total length [tl]
6.76. II: Fe 1.12, PaTi 1.6, MeTa 1.36, tl 4.08. III: Fe 0.84, PaTi 0.96,
MeTa 1.2, tl 3. IV: Fe 1.4, PaTi 1.72, MeTa 1.88, tl 5. Paratype IV: tl
3.4, prosl 1.36, pw 1. Legs: I: Fe 1.92, PaTi 2.4, MeTa 2.4, tl 6.72. II:
Fe 1.36, PaTi 1.48, MeTa 1.6, tl 4.44. III: Fe 1, PaTi 1.16, MeTa 1.28,
tl 3.44. IV: Fe 1.56, PaTi 1.8, MeTa 2, tl 5.36. Range (paratypes IV-IX):
tl 2.9-4.2, pros] 1.28-1.6, pw 1-1.2.

Male (3 specimens): color pattern as in female, chelicerae distinctly light
brown; prolateral spine of first ventral spine pair of metatarsus III thick
and thornlike, no modified spines or processes at coxae III and IV.

COPULATORY ORGANS: median apophysis and distal tegular projection
elaborate and distinct, presumably species-typical; conductor long, thin,
lamelliform and translucent, parallel to embolus.

MEASUREMENTS: Paratype I (USNM): tl 2.72, prosl 1.2, pw 0.92. Legs:
I: Fe 1.96, PaTi 2.72, MeTa 3, tl 7.68. II: Fe 1.12, PaTi 1.64, MeTa 1.52,
tl 4.28. III: Fe 0.96, PaTi 1, MeTa 1.4, tl 3.36. IV: Fe 1.48, PaTi 1.6,
MeTa 1.8, tl 4.88. Paratype II: tl 3 prosl 1.3, pw 1.08. Legs: I: Fe 1.96,
PaTi 2.68, MeTa 3.04, tl 7.68. II: Fe 1.28, PaTi 1.8, MeTa 1.76, tl 4.48.
III: Fe 1, PaTi 1.2, MeTa 1.4, tl 3.6. IV: Fe 1.72, PaTi 1.92, MeTa 2.2,
tl 5.84. Paratypes II and III: tl 2.96-3.6, prosl 1.68-1.32, pw 1.08-1.12.

Male and Female: Eyes: posterior eye row procurved, anterior eye row
recurved, posterior median eyes = posterior lateral eyes > anterior lateral
eyes > anterior median eyes.

CHELICERAE: ?: three teeth at anterior margin, the innermost the largest;
¢; no teeth, but pointed hump at anterior margin. Chelicerae of ~ distinctly
larger than those of ?.

No. 31

NEMOURIA

18

Svs

SNe

shy

December 28, 1988 Sierwald 19

SPINATION OF LEGs: Fe I: dorsal [d] 111, prolateral [pl] 1(long) 1(apical),
retrolateral [rl] l(apical). Fe II-IV: d 111, pl 1 (apical), rl l(apical). Pa
I-IV: d 11. Ti I,I,IV: d 11, pl 11, rl 11, ventral [v] 22. Ti III: d 11, pl
(apical), rl l(apical), v 2. Me I,II: pl 111, rl 111, v 22. Me III,IV: pl
111, rl 111, v 2 and brush of stiff hairs (apical). Me III @: pl 111, rl 111,
v thorn-like spine and 1 normal spine as pair, apically with brush as °.

SPINATION OF PEDIPALPS: Fe: d 1 2(apical), pl I(apical). Pa: d 11, pl
l(proximal). Ti: d 1(proximal), pl 2.

LEG FORMULA: I, IV, IJ, III.

NATURAL HIstTory: collected during both day and night at various localities
and habitats, mostly found while sweeping. Presumably living predomi-
nantly in low vegetation.

TYPE LOCALITY: Bermuda; Sandy’s Parish: Heydon Trust.

DISCUSSION: The male copulatory organs show resemblance to the species
of the Anyphaena pacifica species-group (Platnick, 1974). It is possible
that Anyphaena bermudensis is widely distributed in the West Indies. Most
genera with numerous species of the family Anyphaenidae are currently
unrevised, and it is beyond the scope of this study to review each described
species of the family. Nevertheless, it seems justified to describe these
specimens as a new species and make diagnostic figures available for future
taxonomic work in the Anyphaenidae.

DISCUSSION

Due to its small size (54 km?) and isolation, Bermuda’s terrestrial
fauna and flora are depauperate, with only few endemic species. With 59
species, the total number of recorded spider species is rather small.

[In comparison, on the Madeira Islands (32°45'N, 17°00'W) 145 spider species
occur (in 92 genera, 30 families; Wunderlich, 1987). The Madeira Islands, situated

Figures 1-5: Anyphaena bermudensis n. sp.
1. left male palp, ventral view [paratype III]
female epigynum [paratype IV]
female vulva [paratype IV]
female, color pattern [paratype VIII]
spination of leg II, lateral view [paratype II, male]

pe ete

Scale bars: Figs. 1-3 = 0.2 mm; Figs. 4-5 = 0.5 mm. Abbreviations: c = conductor,
cd = copulatory duct, co = copulatory opening, dtp = distal tegular projection, e
= embolus, fd = fertilization duct, ma = median apophysis, rta = retrolateral
tibial apophysis, s = spermatheca, st = subtegulum, t = tegulum.

20 NEMOURIA No. 31

at the same latitude as Bermuda, have a significantly drier climate (average of 660 —

mm rainfall, mainly in autumn and winter; average temperatures 17°C in winter
and 21°C in summer), are larger (797 km?), with much higher elevations (Pico
Ruivo 1,830 m) and the distance to the coast of Africa is only 600 km.]

The present study adds 26 new records to the 28 previously known
spider species of Bermuda. Sixteen species of the Bermudian spiders are
cosmopolitan or cosmotropical; 12 species are widely distributed in North,
Central and South America and in the West Indies; 15 species are of
North American origin; seven species are European or palearctic. Three
species of Bermudian spiders occur throughout the West Indies. Two
species, Corinna abnormis (Clubionidae) and Habronattus nesiotus (Salti-
cidae), are known from single specimens only. The high number of
widely-distributed species, which are often closely related to human
activities, lead to the conclusion that the majority of Bermudian spiders
were introduced by man. Apparently, some of the species recorded did
not establish permanent populations on Bermuda, since they were only
collected once [e.g., Tegenaria domestica, Neoscona moreli and Metepeira
labyrinthea].

Nephila clavipes and Smeringopus elongatus may be ‘native’ to Bermuda,
as these species were noted by early settlers (Jourdain, 1610, and Strachey,
1625, fide Kevan, 1981; Hughes, 1977). Apparently, Argyrodes nephilae
was already present in the webs of Nephila clavipes. Richard Norwood
[manuscript written about 1622, published in 1945; see Kevan, 1981: 10]
described in the web of the “‘yellow Spider ... her yong ... like little balls
of quick-silver,”’ thus mistaking Argyrodes nephilae for the spiderlings of
Nephila. Gasteracantha cancriformis was first recorded in 1934. No
mygalomorph spiders have been found on Bermuda so far.

Presumably due to the constant mild climate, several spider species
are active year-round. Adults of the following species were collected
throughout the year: Anyphaena bermudensis, Aysha velox; Cyclosa caudata,
Gasteracantha cancriformis, Oxyopes salticus, Hentzia vernalis, Tetragnatha
laboriosa, Argyrodes nephilae and Theridion rufipes. The frost-free climate
allows the occurrence of subtropical species (comprised by cosmotropical
and West Indian forms) that do not inhabit the mainland at the same
latitude.

ACKNOWLEDGMENTS

Travel to Bermuda in 1983 was supported by a grant from the
Studienstiftung des deutschen Volkes (Federal Republic of Germany);
spider collecting was supported by a Sydney L. Wright Fellowship from
the Bermuda Biological Station. Travel to Bermuda in 1988 was supported
by a grant of the Bermuda Department of Agriculture & Fisheries, Paget,

|
|
:

December 28, 1988 Sierwald 21

Bermuda. I wish to thank these institutions for their assistance. I thank
Dr. D. J. Hilburn for his generous hospitality and guidance on Bermuda
during my stay in May 1988. Dr. H. W. Levi and Dr. D. J. Hilburn read
earlier drafts and improved the manuscript through their suggestions. Dr.
H. W. Levi helped illuminate the identity of “‘Araneus thets’’.

I thank the following colleagues for loan of material: D. Azuma,
Academy of Natural Sciences of Philadelphia; Dr. D. Furth, Peabody
Museum of Natural History; and Dr. J. Coddington, National Museum
of Natural History. Dr. C. Griswold kindly assisted with the identifications
of Salticidae.

This paper is contribution No. 1189 from the Bermuda Biological
Station for Research.

LITERATURE CITED

Banks, N. 1902. Some spiders and mites from the Bermuda Islands. Transactions
of the Connecticut Academy of Arts and Sciences, 11: 267-275.

Beatty, J. A. & Berry, J. W. 1988. The spider genus Paratheuma Bryant (Araneae,
Desidae). The Journal of Arachnology, 16(1): 47-54.

Blackwall, J. 1868. Notice of several species of spiders supposed to be new or little
known to Arachnologists. The Annals and Magazine of Natural History (4)2:
403-410.

Brady, A. 1964. The lynx spiders of North America, north of Mexico (Araneae:
Oxyopidae). Bulletin of the Museum of Comparative Zoology, 131(13):
432-518.

—— 1986. Nearctic species of the new wolf spider genus Gladicosa (Araneae,
Lycosidae). Psyche, 93(3-4): 285-319.

Chamberlin, R. V. & W. J. Gertsch. 1958. The spider family Dictynidae in America
north of Mexico. Bulletin of the American Museum of Natural History,
116(1): 1-152.

Chickering, A. M. 1969. The Family Oonopidae in Florida. Psyche, 76(2): 144-162.

Comstock, J. H. 1910. The palpi of male spiders. Annals of the Entomological
Society of America, 3: 161-185.

Emerton, J. 1902. The common spiders of the United States. Dover Publications,
New York, 227 pp.

Exline, H. & H. W. Levi. 1962. American spiders of the genus Argyrodes (Araneae,
Theridiidae). Bulletin of the Museum of Comparative Zoology, 127(2):
75-202.

Gertsch, W. J. & F. Ennik. 1983. The spider genus Loxosceles in North America,
Central America and the West Indies (Araneae, Loxoscelidae). Bulletin of
the American Museum of Natural History, 175(3): 263-360.

2 NEMOURIA No. 31 —

Grasshoff, M. 1986. Die Radnetzspinnen-Gattung Neoscona in Afrika (Arachnida:
Araneae). Annales Musée royal de l’Afrique Central, Tervuren, Sciences
zoologiques, 250: 123 pp.

Griswold, C. E. 1987. A revision of the jumping spider genus Habronattus
F.O.P.-Cambridge (Araneae; Salticidae), with phenetic and cladistic analyses.
University of California Publications in Entomology, 107, 344 pp.

Hughes, I. W. 1977. The golden silk spider. Monograph Bulletin Bermuda
Department of Agriculture, 47(10): 79-80.

Ivie, W. 1969. North American spiders of the genus Bathyphantes (Araneae:
Linyphiidae). American Museum Novitates, 2364: 1-70.

Jarvi, T. H. 1914. Das Vaginalsystem der Sparassiden. Annales Academiae
Scientiarum Fennicae, (A) 4(1): 1-248, 11 pl.

Kaston, B. J. 1981. Spiders of Connecticut. State Geological and Natural History
Survey, Bulletin No. 70, revised edition. 1020 pp.

Kevan, D. K. McE. 1981. The terrestrial arthropods of the Bermudas: an historical
review of our knowledge. Archives of Natural History, 10(1): 1-29.

Lehtinen, P. 1967. Classification of the cribellate spiders and some allied families,
with notes on the evolution of the suborder Araneomorpha. Annales Zoologici
Fennici, 4: 199-468.

Levi, H. W. 1955. The spider genera Coressa and Achaearanea in America north
of Mexico (Araneae, Theridiidae). American Museum Novitates, 1718: 1-33.

—— 1956. The spider genera Neottiura and Anelosimus in America (Araneae:
Theridiidae). Transactions of the American Microscopical Society, 75(4):
407-422.

— 1957. The spider genera Enoplognatha, Theridion, and Paidisca in America
north of Mexico (Araneae, Theridiidae). Bulletin of the American Museum
of Natural History, 112(1): 5-123.

— 1959a. The spider genus Latrodectus (Araneae: Theridiidae). Transactions of
the American Microscopical Society, 78(1): 7-43.

— 1959b. The spider genus Coleosoma (Aranea, Theridiidae). Breviora, Museum
of Comparative Zoology, 110: 1-8.

— 1977a. The American orb-weaver genera Cyclosa, Metazygia and Eustala north
of Mexico. Bulletin of the Museum of Comparative Zoology, 148(3): 61-127.

—— 1977b. The orb-weaver genera Metepeira, Kaira and Aculepeira in America
north of Mexico (Aranea: Araneidae). Bulletin of the Museum of Comparative
Zoology, 148(5): 185-238.

— 1978. The American orb-weaver genera Colphepeira, Micrathena and Gastera- —
cantha north of Mexico (Araneae: Araneidae). Bulletin of the Museum of
Comparative Zoology, 148(9): 417-442.

— 1980. The orb-weaver genus Mecynogea, the subfamily Metinae and the genera
Pachygnatha, Glenognatha and Azilia of the subfamily Tetragnathinae north
of Mexico (Araneae: Araneidae). Bulletin of the Museum of Comparative
Zoology, 149(1): 1-75.

—— 1981. The American orb-weaver genera Dolichognatha and Tetragnatha north
of Mexico. Bulletin of the Museum of Comparative Zoology, 149(5): 271-318

December 28, 1988 Sierwald 23

-— 1983. On the value of genitalic structures and coloration in separating species
of widow spiders (Latrodectus sp.). Verhandlungen des naturwissenschaftli-
chen Vereins in Hamburg, (NF)26: 195-200.

—— 1987. Spiders and their kin. Golden Press, New York, 160 pp. [color figures].

Marx, G. 1889. A contribution to the knowledge of the spider fauna of the Bermuda
Islands. Proceedings of the Academy of Natural Sciences of Philadelphia,
1889: 98-101.

Norwood, R. [ca. 1622] 1945. The description of the Sommer Iands, once called
the Bermudas. Jn: Craven, W. C. & W. B. Hayward (eds.). The Journal of
Richard Norwood, Surveyor of Bermuda. Bermuda Historical Monument
Trust by Scholar’s Facsimiles & Reprints, New York, |xv-xcii. [non vidi,
fide Kevan, 1981].

Opell, B. 1979. Revision of the genera and tropical American species of the spider
family Uloboridae. Bulletin of the Museum of Comparative Zoology, 148(10):
443-549.

Petrunkevitch, A. 1929. The spiders of Puerto Rico. Part I. Transactions of the
Connecticut Academy of Arts and Sciences, 30: 158 pp.

—— 1930. The spiders of Puerto Rico, Part III. Transactions of the Connecticut
Academy of Arts and Sciences, 31: 191 pp.

Pickard-Cambridge, F. O. 1905. Araneida, part 2, pp. 561-610, pls. 52-54, in:
Biologia Centrali-Americana, Zoologia, 11, London.

Platnick, N. I. 1974. The spider family Anyphaenidae in America north of Mexico.
Bulletin of the Museum of Comparative Zoology, 146(4): 205-266.

— 1975. A revision of the holarctic genus Callilepis (Araneae, Gnaphosidae).
American Museum Novitates, 2573: 32 pp.

— 1977. Notes on the spider genus Paratheuma Bryant (Arachnida, Araneae).
The Journal of Arachnology, 3: 199-201.

Platnick, N. I. & Shadab, M. U. 1982. A revision of the American spiders of the
genus Camillina (Araneae: Gnaphosidae). American Museum Novitates, 2748:
1-38.

Roberts, M. 1985. The spiders of Great Britain and Ireland. Harley Books,

Colchester, Vol. 1, 229 pp. [Atypidae to Theridiosomatidae]; Vol. 3, 256
pp., 237 color pls. [Atypidae to Linyphiidae].

—— 1987. The spiders of Great Britain and Ireland. Harley Books, Colchester,
Vol. 2, 204 pp. [Linyphiidae].

Roewer, F. 1928. 4. Ordnung. Araneae. Echte oder Webspinnen. Jn: Brohmer, P.,
P. Ehrmann & G. Ulmer (eds.), Die Tierwelt Mitteleuropas, III(6): 144 pp.

— 1942. Katalog der Araneae. Kommissions-Verlag von ‘‘Natura”, Bremen,
Vol. 1, 1040 pp.

—— 1954. Katalog der Araneae. Institut Royal des Sciences Naturelles de Belgique,
Brussels, Vol. 2a, pp. 1-923, 2b, pp. 924-1751.

— 1960. Araneae, Lycosaeformia II, pp. 519-1040, im: Exploration du Parc
National de Upemba, 55. Institut des Parcs Nationaux du Congo Belge,
Brussels.

24 NEMOURIA No. 31 |

Roth, V. D. 1956. Taxonomic changes in Agelenidae. The Pan-Pacific Entomologist,
32(4): 175-180.

— 1968. The spider genus Tegenaria in the western hemisphere. American
Museum Novitates, 2323: 1-33.

—— 1985. Spider genera of North America. American Arachnological Society,
Portal, Arizona, 1-30 + A-1-1-B-47-1 pp.

—— 1988. Linyphiidae of America north of Mexico. American Arachnological
Society, 62 pp.

Simon, E. 1883. Etudes arachnologiques: 14° Mémoire. XXI. Matériaux pour
servir 4 la faune arachnologique des Iles de l’?Océan Atlantique (Acores,
Madére, Salvages, Canaries, Cap Vert, Sainte-Héléne et Bermudes). Annals
de la Société entomologique de France, (6)3: 259-314, pl. 8 [‘‘Iles Bermudes”’:
307; Liste générale: 308-313].

Turnbull, A. L., C. D. Dondale & J. H. Redner. 1965. The spider genus Xysticus
C. L. Koch (Araneae: Thomisidae) in Canada. The Canadian Entomologist,
97(12): 1233-1280.

Valerio, C. E. 1981. Spitting spiders (Araneae, Scytodidae), Scytodes from Central
America. Bulletin of the American Museum of Natural History, 170(1): 80-89.

Wallace, H. K. 1942. A study of the Jenta-group of the genus Lycosa, with
descriptions of new species (Araneae: Lycosidae). American Museum Novi-
tates, 1185: 1-21.

Wunderlich, J. 1987. Die Spinnen der Kanarischen Inseln und Madeiras (The
spiders of the Canary Islands and Madeira), 435 pp., Triops Verlag, Langen,
Germany.

Zorsch, H. 1937. The spider genus Leptyphantes in the United States. The American
Midland Naturalist, 18(5): 856-898.

Reviewers of this paper: Dr. James W. Berry, Butler University, Indianapolis, and
Dr. William K. Emerson, American Museum of Natural History.

President Managing Editor
JOHN E. DU PONT RUDIGER BIELER

Consulting Editors
WILLIAM K. EMERSON, American Museum of Natural History, New York, NY.
M. G. HARASEWYCH, National Museum of Natural History, Smithsonian Institu-
tion, Washington, DC.
KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh, PA.
RICHARD W. THORINGTON, JR., National Museum of Natural History, Smith-
sonian Institution, Washington, DC.

Published by the
DELAWARE MUSEUM OF NATURAL HISTORY
4840 Kennett Pike, Wilmington,
Delaware 19807, U.S.A.

Sb

NEMOURIA

Occasional Papers of the Delaware Museum of Natural History

NUMBER 32 FEBRUARY 28, 1989

NOTES ON THE BIRDS COLLECTED
IN THE PHILIPPINES DURING
THE STEERE EXPEDITION OF 1887/1888

Edward C. Dickinson!
Robert S. Kennedy?
Derek K. Read?
Frank G. Rozendaal?

ABSTRACT. Of the 5000 bird specimens obtained during the Steere Expedition to
the Philippines of 1887/1888, nearly 70% have been located. In this paper, we
show that some specimens from the Expedition obtained from E. L. Moseley, and
F. S. Bourns and D. C. Worcester, were retained by these collectors, and therefore
not available to Steere (1890), and should not be considered syntypes of taxa
described by Steere. We also recommend the deletion of 10 island records of 9
species due to error or lack of evidence; tabulate 9 records apparently based solely
upon the Steere Expedition’s findings for which we have been unable to locate the
supporting specimens; report 30 new and/or first island records of 26 species;
substantiate 14 earlier records by reference to the specimens; comment on 6 species
and 2 genera with doubtful records or confused histories; and finally, designate a
lectotype of Phylloscopus olivaceus.

INTRODUCTION

Joseph Beal Steere, Professor of Zoology at the University of Michigan
in Ann Arbor, Michigan first visited the Philippines in 1874 and 1875.
His collection of birds made on this trip was fully reported by Sharpe
(1877) and most of the type specimens are in Ann Arbor. Later Steere led

1190 Old Barn Road, Barrington, Illinois, 60010 U.S.A. Serial Publication
(For all correspondence regarding this paper). ISSN 0085-3887

2Cincinnati Museum of Natural History, 1720 Gilbert Ave.,
Cincinnati, Ohio 45202, U.S.A.

37 Park Street, Tring, Herts. HP23 6AW, United Kingdom.
4Prins Hendriklaan 58, 3721 AT Bilthoven, The Netherlands.

2 NEMOURIA No. 32

an Expedition to the Philippines that lasted from August 1887 until July
1888. He was accompanied by three students - Frank S. Bourns and Dean
C. Worcester, who were to have a close association with Philippine birds
for many years, and Edward L. Moseley - and by Mateo Francisco, a
Filipino who had been with Steere since 1875. Steere (1890, 1891a, 1894)
reported on the ornithological results of the second expedition. Although
he described 53 new avian taxa, his main account (Steere, 1890) was not
much more than a list with brief descriptions of the new forms. Additional
information on the expeditions appeared separately in more popular
accounts (Steere, 1888a, b, c, 1889, 1891b, c). In his chronology of islands
visited, Steere (1890) did not give the exact dates that the party was on
each island. Rather, he gave only the main months of such visits often
omitting parts of months at the beginning or end of a visit. For example,
he gave April 1888 for Samar but the party arrived there well before the
end of March. In fact, there were times when the party separated thus
adding to the confusion.

Over 5000 bird specimens were taken during the expedition (Steere,
1894). We have located about 70% of these. The largest share, about one
third of the original collection, is now in the British Museum of Natural
History. The rest is widely scattered (Table 1).

As we searched for the dispersed collection and its type material, we
discovered that Steere (1890) had made several errors in identification and
had omitted many important distributional records. In addition, we learned
that many museum curators were confused about the type status of
specimens obtained not from Steere but from other members of the
expedition.

In this paper, we address the errors and omissions and clarify the
type status of specimens from Steere’s colleagues. Finally, we comment
on a few species with doubtful records or confused history, and in one
case designate a lectotype. Latin names are almost all from duPont (1971).

ABBREVIATIONS USED

AMNH American Museum of Natural History, New York
BMNH _ British Museum (Natural History), London

ISB Institut Scientifique d’Histoire Naturelle, Brussels
OSUMZ _ Ohio State University Museum of Zoology, Columbus,
Ohio

USNM (United States) National Museum of Natural History,
Smithsonian Institution, Washington, DC

February 28, 1989 Dickinson, et al 3

Table 1. Present location of bird specimens from the 1887/1888 Steere
Expedition to the Philippines.

No. of
Museum Skins Origin
British Museum of Natural History 1773 Steere (1562)

Moseley (211)
National Museum of Natural History, 658 Worcester (625)!

Smithsonian Institution Bell Museum (33)
Field Museum of Natural History 178 Steere
Museum of Comparative Zoology 83 ? Moseley
Cambridge University 18 ? Moseley
Rijksmuseum van Natuurlijke 82 Moseley

Historie, Leiden
Carnegie Museum of Natural History 16 Bell Museum (5)

Purdue (10)
Ohio Wesleyan (1)

American Museum of Natural History 542, ? Moseley ex Rothschild
Royal Ontario Museum 32 Moseley
University of Michigan Museum ?3—- Steere

of Zoology
Purdue University, Indiana 154 Steere
Denver Museum of Natural History 298 Chicago Academy

of Science

Chicago Academy of Science 15 Moseley (12),

Steere (287)
Bowling Green State University Museum 88 Moseley

Bell Museum of Natural History 96 > Worcester
Ohio State University Museum 65 Moseley
of Zoology

'Bell Museum now holds the rest and the Menage Expedition catalogue into which
these were entered, but acquisition was from the Minneapolis Public Library.
“The AMNH may hold some 500 or more specimens, this is the number of relevant
types held.

3Steere taught here, so many may be here.

4Remaining after exchanges to the Carnegie Museum.

>Remaining after sending 298 to Denver.

®Remaining from at least 53 of which 33 went to USNM and 5 to the Carnegie
Museum.

4 NEMOURIA No. 32
ELIGIBILITY FOR TYPE STATUS

Steere originally intended that all specimens should be given to him
to write up (Hachisuka, 1931). However, some skins were retained by
Moseley and others by Bourns and Worcester.

Evidence that Moseley had held onto some skins came when he
described two new forms (Moseley, 1891). In addition, had Steere had
Moseley’s material, he would certainly have included the male, as well as
the female, when he (Steere, 1890) described Philemon [= Arachnothera
clarae| philippensis. Moseley’s male spiderhunter and the types of his two
forms were among 211 specimens that were catalogued at the BMNH as
early as 1890 and 1891. The catalogue dates are important because they
further show that these skins could not have been in front of Steere (1890)
when he described his new forms. Moseley appears to have retained most
of his specimens and to have sold these directly to various institutions and
none of these can be considered syntypes of Steere’s new taxa.

After the Steere Expedition, Bourns and Worcester returned to the
Philippines from 1890-1893 on the Menage Expedition. When they
published their results, however, they (Bourns & Worcester, 1894) included
a few new records based on 1887-1888 material and not upon specimens
taken in 1890-1893 (Worcester & Bourns, 1898). This clearly suggests that
they had kept these skins from Steere. In 1897, the ownership and
possession of Steere Expedition material by Bourns and Worcester was
confirmed when Worcester presented 625 skins from that expedition to
the United States National Museum.

We believe museum material from this Expedition can only be
considered to include syntypes of Steere’s new taxa if it can be shown
that the material was obtained from Steere; such material may however
include skins taken by Bourns, Moseley or Worcester. Curators should
be aware of this and adjust their collections accordingly.

COMMENTS ON RECOMMENDED DELETIONS

In this section, we recommend the deletion of ten island records of
nine species (see Table 2) due to a high probability of error or, in one
case, to lack of supporting evidence.

Ardea cinerea Linné, 1758: Steere (1890) recorded this species from
Guimaras but the skin that must have served as the basis for this record
(BMNH 1896.6.6.1360) is in fact Ardea sumatrana (see under New
Records). The sole record of A. cinerea for Guimaras must therefore be
deleted.

February 28, 1989 Dickinson, et al 5

Table 2. Recommended deletions of records solely dependant upon the
Steere Expedition.

aa
F ©
(7)
=) <t
Z : (a = An
Gq Z 22 90 4,
4 Os += ARQ & a
am Ore es 2 oe
Ardea cinerea x
Egretta intermedia xX
Dupetor flavicollis xX
Spizaetus philippensis x
Porzana paykullu x
Charadrius alexandrinus xX
Charadrius peronii
Calidris temminckii x
Anthus cervinus X xX

Egretta intermedia (Wagler, 1829): Steere (1890) listed Ardea intermedia
from Bohol and Samar. He also listed Ardea jugularis - which Bourns &
Worcester (1894) showed to be Egretta sacra (Gmelin, 1789) - from the
islands of Balabac, Bohol, Cebu and Marinduque. The BMNH register
Shows eight specimens of Demiegretta [sic] sacra from these four islands,
plus Samar. These eight specimens appear to represent both of Steere’s
listed species. Of these, five (BMNH 1896.6.6.1402, 1403, 1404, 1407 and
1408) are dark phase Egretta sacra. The three white ones (BMNH
1896.6.6.1401, 1405 and 1406) are not white-phase E. sacra but have been

6 NEMOURIA No. 32

shown to be Egretta eulophotes (Swinhoe, 1860) (see Amadon, 1951). In
summary E. sacra is represented from the four islands that Steere (1890)
listed for his Ardea jugularis, whilst Steere’s records of E. intermedia from
Bohol and Samar, which were subsequently used by McGregor (1909),
should be deleted. However, there are now two ringing recoveries of E.
intermedia from Samar (McClure, 1974).

Dupetor flavicollis (Latham, 1790): Steere (1890) listed this, as Ardetta
flavicollis, from Marinduque and Mindanao but did not include the related
species Ixobrychus sinensis (Gmelin, 1789) or I. eurhythmus (Swinhoe, 1873)
in his list. There are Steere Expedition specimens of I. sinensis (BMNH
1896.6.6.1393 and 1394) from Marinduque and Mindoro, and specimens
of I. eurhythmus from Mindanao (BMNH 1896.6.6.1399 and 1400) - one
of which was correctly re-identified by Ogilvie Grant, as mentioned in
McGregor (1909). It appears that Steere overlooked his Mindoro specimen
and took all these birds to be D. flavicollis. Therefore, Steere’s records of
D. flavicollis should be deleted, which leaves the species unrecorded for
Marinduque.

Spizaetus philippensis Gould, 1863: Steere (1890) listed this species
from Basilan, Masbate and Negros. The record from Basilan was apparently
based on a specimen (BMNH 1896.4.15.8) originally identified as S.
philippensis, since reidentified as Pernis celebensis steerit Sclater, 1919. The
Masbate record, if represented by a single skin, is probably based on a
Steere specimen in the Institut Scientifique d’Histoire Naturelle, Brussels
(ISB 9390), that is also believed to be P. c. steeritz. The Negros specimen
(BMNH 1896.4.15.40) was misidentified as well. In fact, Sclater (1919)
used this as the type specimen when he described P. c. steerii. Clearly
Steere (1890) listed S. philippensis in error and his records should be
deleted. However, there remain valid records of this species from Basilan
(Worcester & Bourns, 1898) and Negros (Tweeddale, 1878), but not from
Masbate.

Porzana paykulliu (Ljungh, 1813): Dickinson (1984) pointed out that
the bird from Basilan listed as this species by Steere (1890) is Rallina
fasciata.

Charadrius alexandrinus Linné, 1758: Steere (1890) listed this species, |
as Aegialites cantianus, from Bohol, Palawan and Panay. None of these |
three islands is represented by Steere skins in the BMNH; his only |
specimens of C. alexandrinus are from Negros and Siquijor. Steere’s “A.
cantianus” seems almost certain to have been C. mongolus for there are |
Steere skins of this from Bohol and Panay in the BMNH. McGregor |
(1909) listed other collectors of C. alexandrinus from Bohol and Palawan |

February 28, 1989 Dickinson, et al 7

but listed only the Steere Expedition as the source of a Panay record. As
the Steere record must be deleted there now no records of this species
from Panay.

Charadrius peronii Schlegel, 1865: Steere (1890) listed this species, as
Aegialites peronii, for Mindanao, Negros and Siquijor. McGregor (1909)
cited the Steere Expedition as the only collectors from Negros. The birds
before Steere seem likely to have been a composite collection of C.
alexandrinus - the skins mentioned above - and C. peronii. There are
indeed Steere skins of confirmed C. peroni from Mindanao in the BMNH,
and a Siquijor skin in the USNM taken on the Steere Expedition by
Bourns or Worcester; but no C. peroni skin has been traced from Negros.
Probably, Steere’s Negros record was based on a skin of C. alexandrinus.

Calidris temminckii (Leisler, 1812): Parkes (1986a) has shown that
Steere’s (1890) record from Negros is based on a specimen of C. minuta;
thus there is no valid record of C. temminckii from Negros.

Anthus cervinus (Pallas, 1811): Listed by Steere (1890) from Mindanao
and Basilan, but the Basilan record was not listed by McGregor (1909).
No skins in the BMNH substantiate these records from either island;
indeed it seems probable that Steere was confused between A. gustavi
Swinhoe, 1863, and A. cervinus. As there are no other records of A.
cervinus from either Basilan or Mindanao, it seems best to delete Steere’s
records until proof can be found.

COMMENTS ON RECORDS NEEDING SUBSTANTIATION

Because we have not located all of the specimens collected by the
Steere Expedition, we were not surprised that we could not find the
specimens to substantiate nine key records (see Table 3). However, the
number is remarkably small since the BMNH has obtained a representative
collection. We feel that each record will eventually be confirmed and thus
we do not recommend their deletion.

NEW OR FIRST ISLAND RECORDS

In all we have identified 30 new and/or first island records for 26
species that Steere and his colleagues either misidentified, or for one reason
or another, failed to report in their publications.

Ardea sumatrana Raffles, 1822: Guimaras: BMNH 1896.6.6.1360 taken
28 December 1887. Misidentified by Steere (1890) as A. cinerea (see above).

8 NEMOURIA No. 32

Table 3. Island records dependant upon Steere (1890) that have not been
substantiated so far.

ea)
eS 0
< 2 wm me
A -E Sm O
GQ 7 eS ieee
= 27 2 os.
5242439
Os = BS ae
Anas luzonica x
Falco peregrinus Xx
Rallus striatus xX
Ducula poliocephala x
Ninox philippensts xX
Pelargopsis capensis xX
Halcyon winchelli x
Motacilla cinerea x!
Anthus gustavi x

IThis record was not reported by Steere (1890) but was by McGregor (1909).

Ixobrychus sinensis (Gmelin, 1789): Marinduque: BMNH 1896.6.6.1393
taken 17 May 1888. Appears to have been treated as Dupetor flavicollis
by Steere (1890).

Pernis celebensis Wallace, 1868: Masbate: ISB 9390. As shown above,
Steere’s specimens of what he believed to be Spizaetus philippinensis appear
to have been P. c. steeriit. We have not examined the specimen from

February 28, 1989 Dickinson, et al 9

Masbate but the ISB register shows an annotation correcting Steere’s
identification to P. c. steerii. If correctly identified, this appears to be the
first record of this species from Masbate.

Haliaaetus leucogaster (Gmelin, 1788): Panay: USNM 161108 taken
in January 1888. Previously known only by sight records from islets near
Panay (Rabor, 1938; Alcala & Alviola, 1970).

Circus melanoleucos (Pennant, 1769): Panay: BMNH 1891.4.25.2 taken
in January 1888. Not mentioned by Steere (1890), this skin was purchased
from Moseley.

Accipiter soloensis (Horsfield, 1821): Guimaras: BMNH 1896.4.15.3
taken 25 January 1888, and omitted by Steere (1890).

Turnix suscitator (Gmelin, 1789): Guimaras: BMNH 1896.6.6.1229
taken 17 January 1888, and omitted by Steere (1890).

Rallina eurizonoides (Lafresnaye, 1845): Guimaras: BMNH 1896.6.6.1265
taken 6 January 1888. Marinduque: BMNH 1896.6.6.1266 taken 5 May
1888. Samar: BMNH 1896.6.6.1268 taken 2 April 1888. Siquijor: BMNH
1896.6.6.1267 taken 14 February 1888. All of these specimens were omitted
by Steere (1890), with the possible exception of the juvenile Siquijor bird,
which Steere may have listed as “‘Porzana erythrothorax.”

Amaurornis olivacea (Meyen, 1834): Marinduque: BMNH 1890.12.1.115
taken 11 May 1888; a Moseley specimen, not listed by Steere (1890).

Charadrius alexandrinus Linné, 1758: Negros: BMNH 1896.6.6.1313
taken 7 February 1888. This first specimen record of C. alexandrinus
dealbatus (Swinhoe, 1870), from Dumaguete, was probably listed by Steere
(1890) as Aegialites (= Charadrius) peronii (see above). McClure (1974)
reported a recent banding recovery of this species from Negros. DuPont
(1976) based his record of the race C. a. alexandrinus on this recovery.

Charadrius mongolus Pallas, 1776: Panay: BMNH 1896.6.6.1299 taken
1 January 1888. As mentioned earlier, Steere (1890) identified this specimen
as “‘Aegialites cantianus’’.

Gallinago megala Swinhoe, 1861: Samar: BMNH 1896.6.6.1341 taken
26 March 1888; omitted by Steere (1890).

Chlidonias hybridus (Pallas, 1811): Guimaras: BMNH 1896.6.6.1436
taken 6 January 1888; omitted by Steere (1890). Bourns & Worcester
(1894) claimed new records of C. hybridus for 15 islands, including
Guimaras, but only as a result of two errors; first, at the foot of page 30
only Luzon, Samar and Mindanao should have appeared-the islands listed

10 NEMOURIA No. 32

after the full stop that follows Mindanao are a duplication by the printer
of the islands listed correctly for Sterna bergi at the top of page 31, as
may be found by checking the table in Worcester & Bourns (1898);
secondly, that same table shows C. hybridus only for Luzon and Palawan,
so that the 1894 listing for Samar and Mindanao was presumably found
later to be based on misidentified specimens. We know of no other record
of this species from Guimaras.

Phapitreron leucotis (Temminck, 1823): Marinduque: BMNH
1896.6.6.1109 taken 15 May 1888; omitted by Steere (1890). Identified as
P. 1. leucotts.

Macropygia phasianella (Temminck, 1821): Malamaui: USNM 161089
taken 10 November 1887; probably listed by Steere (1890) under Basilan,
which Malamaui adjoins.

Cacomantis variolosus (Vigors & Horsfield, 1826): C. v. everetti
Hartert, 1825-Malamaui: USNM 161189 taken 9 November 1887. C. v.
sepulcralis (S. Miiller, 1843)-Panay: BMNH 1896.6.6.956 taken 14 January
1888. Steere (1890) did not list this species, but did list C. merulinus
Scopoli, 1786, from both Basilan (which could have comprised Basilan and
Malamaui) and Panay. It was only recently recognized that C. vartolosus
occurred sympatrically with C. merulinus almost throughout the Philippines.
Of the ten islands that Steere listed for C. merulinus, based on Steere
Expedition skins in the AMNH, the BMNH and the USNM, records from
Basilan, Cebu, Leyte, Luzon, Masbate, Palawan and Panay are authentic.
We have also found Steere Expedition skins of C. variolosus from Basilan,
Malamaui, Mindanao, Panay and Samar; those from Malamaui and Panay
appear to be first records. No Steere Expedition skin of either species from
Negros has been found.

Otus megalotis (Walden, 1875): Marinduque: OSUMZ (not numbered)
in May 1888 by E. L. Moseley from Boac, Marinduque. This first specimen
record has been identified by RSK and J. T. Marshall as a juvenile red
phase of O. m. megalotis. Marshall (1978) has reported on two more recent _
specimens (in the Philippine National Museum) of this species from —
Marinduque. |

Collocalia esculenta (Linné, 1758): Guimaras: BMNH 1896.6.6.786-87
taken 8 January 1888; omitted by Steere (1890).

Pericrocotus divaricatus (Raffles, 1822): Balabac: BMNH 1896.6.6.108 |
taken 29 September 1887; omitted by Steere (1890). |

February 28, 1989 Dickinson, et al 1]

Corvus macrorhynchos Wagler, 1827: Malamaui: OSUMZ 10537 taken
11 November 1887 by E. L. Moseley.

Copsychus saularis (Linné, 1758): Marinduque: BMNH 1896.6.6.276
taken 11 May 1888; BMNH 1896.6.6.277 taken 18 May 1888; omitted by
Steere (1890). The amount of buff on the underparts of females is said to
distinguish C. s. deuteronymus Parkes, 1963, of Luzon from C. s.
mindanensis (Boddaert, 1783) of the southern islands (duPont, 1971).
Judging from material in the BMNH this character varies clinally and the
Marinduque birds are intermediate.

Gerygone sulphurea Wallace, 1864: Basilan: BMNH 1896.6.6.129 and
130 taken 22 November 1887; omitted by Steere (1890). These two
specimens from Basilan are of the race G. s. rhizophorae Mearns, 1905,
which Parkes (1986b) has shown to be distinct.

Acrocephalus stentoreus (Ehrenberg, 1833): Mindanao: BMNH
1896.6.6.222 taken 25 October 1887. Presumably this is the specimen listed
for this date under A. arundinaceus by Steere (1890).

Acrocephalus [arundinaceus] orientalis (Temminck & Schlegel,
1847): Marinduque: BMNH 1896.6.6.223 taken 16 May 1888, wing length
86 mm, the wing formula confirming the identity. Steere (1890) listed
only Mindanao (the specimen discussed above) and Cebu. McGregor
(1909) cited both the Steere Expedition (which was not valid) and
Goodfellow as collectors of A. [a.] orientalis from Mindanao. There are
three skins in the BMNH taken by Goodfellow near Davao that support
his Mindanao records. Steere’s Cebu specimen has not been traced.
However, McGregor (1909) named himself and Meyer as other collectors
of A. [a.] orientalis from Cebu. We have located Meyer’s supporting
specimen in the BMNH.

Pachycephala cinerea (Blyth, 1847): Mindanao: BMNH 1896.6.6.345
taken 10 December 1887; omitted by Steere (1890). This specimen taken
at Ayala near Zamboanga, appears to match specimens of P. c. homeyeri
(Blasius, 1890) from the Sulu Archipelago.

Dicaeum bicolor (Bourns & Worcester, 1894): Guimaras: BMNH
1896.6.6.563 taken 30 December 1887; omitted by Steere (1890). D. bicolor
was described by Bourns & Worcester (1894) from their Menage Expedition
material, and yet the Steere Expedition had taken a male (BMNH

12 NEMOURIA No. 32

1896.6.6.562) in Negros and the above mentioned female (BMNH
1896.6.6.563) in Guimaras. The latter has upperparts more like D. 5.
inexpectatum (Hartert, 1895) and underparts like the nominate form and
thus closely matches the description of D. b. viridissimum Parkes, 1971, of
Negros.

SUBSTANTIATION OF UNUSUAL RECORDS
FROM THE EXPEDITION

In our review of the important island records from the Steere
Expedition, we identified 14 published records that were either questioned
or omitted by subsequent authors because the accounts lacked sufficient
supporting data. Here we substantiate these records by providing details
on the supporting specimens.

Sula sula (Linné, 1766): The synonymy of Sula spp. in the Philippine
literature is confused; for this reason the skin (BMNH 1896.6.6.1414) listed
by Steere (1890) as “Sula piscatrix’’ has been re-examined. It was collected
at Ayala, Mindanao on 10 October 1887, and is an immature Sula sula.

Megapodius freycinet Gaimard, 1823: Steere (1890) listed this species
from Masbate, but McGregor (1909) omitted the record. A specimen
(BMNH 1896.6.6.1225) taken 2 May 1888 substantiates the record.

Turnix suscitator (Gmelin, 1789): Although not listed by Steere (1890)
from Negros, McGregor (1909) credited the Expedition with a record from
there and mentioned eggs supposedly taken on 10 November [1887], when
in fact the Expedition was in Guimaras. M. P. Walters (in litt.) confirms
that the eggs, now in the BMNH, were in fact taken on 10 February 1888,
at which date the Expedition was indeed in Negros.

Porzana pusilla (Pallas, 1776): DuPont (1972) did not accept the
Marinduque record for this species. It was published, not by Steere, but
by Hachisuka (1931), and is confirmed by a specimen (BMNH 1896.6.6.1270)
collected on 12 May 1888.

Charadrius dubius Scopoli, 1786: McGregor (1909) credited the Steere

|

Expedition with a specimen from Panay, although this was not listed by |

Steere (1890). A skin (BMNH 1890.12.1.132) collected on 18 January |

q
i
i
|

taken in Palawan by the Steere Expedition between 12 and 22 September |

1888 by Moseley confirms this.

Charadrius veredus Gould, 1848: Not listed by Steere (1890). However,
three skins (BMNH 1896.6.6.1317 and 1318 and USNM 314793) were

February 28, 1989 Dickinson, et al 13

1887. The USNM skin no doubt provided the record in McGregor (1909),
credited to Bourns & Worcester; the date on the specimen label shows it
was collected by them during the Steere Expedition, not the Menage
Expedition as implied by the table in Worcester & Bourns (1898).

Numenius minutus Gould, 1841: Bourns & Worcester in McGregor
(1909) reported it taken in Marinduque in 1888 by Bourns. A specimen
(USNM 314815) taken on 15 May 1888 supports this.

Limosa limosa Linné, 1758: McGregor (1909) correctly credited the
Steere Expedition with a record of this species from Negros, in addition
to a record of Limosa lapponica, although Steere (1890) claimed only the
latter. Specimen BMNH 1890.12.1.124, purchased from Moseley and
registered as L. melanura, is L. limosa melanuroides Gould, 1846, while
BMNH 1896.6.6.1346 received from Steere and registered as L. novae-
zeelandiae is L. lapponica baueri Naumann, 1836.

Calidris tenuirostris (Horsfield, 1821): Omitted by Steere (1890), but
he, and also Bourns & Worcester, are given as collectors in Negros by
McGregor (1909). The record credited to Steere is a specimen (BMNH
1896.6.6.1338) collected on 4 February 1888. Moseley collected a further
specimen (BMNH 1890.12.1.127) on 6 February. A third specimen, once
given the Menage Expedition no. 4489, taken on 7 February 1888 is in
the Bell Museum (uncatalogued there), Minneapolis (per K.C. Parkes).

Tanygnathus sumatranus (Raffles, 1822): Listed only from Mindanao
by Steere (1890), but Bourns & Worcester (in McGregor, 1909) mentioned
that Moseley collected this species in Panay. The supporting specimen is
BMNH 1890.12.1.80, taken on 16 January 1888.

Chrysococcyx russatus Gould, 1868: Listed as C. basalis by Steere
(1890) and apparently overlooked by McGregor (1909), Parker (1981)
reidentified the specimen (BMNH 1896.6.6.963), collected in Basilan on
21 November 1887.

Ninox scutulata (Raffles, 1822) subsp.: Listed from Guimaras as Ninox
lugubris by Steere (1890) but not included by McGregor (1909). A specimen
(BMNH 1896.4.15.51) from Guimaras taken 25 January 1888 has not been
identified to subspecies.

Rhabdornis mystacalis (Temminck, 1825): DuPont (1971) omitted
Basilan from the range of R. m. minor Ogilvie-Grant, 1896, although
McGregor (1909) listed Basilan on the basis of a Celestino record. In
addition there is an unreported Steere Expedition specimen (USNM
161551) of R. m. minor collected in Basilan in November 1887.

14 NEMOURIA No. 32

Megalurus timoriensis Wallace, 1864: Although McGregor (1909)
listed this species from Masbate duPont (1971) omitted it. However
Moseley collected a specimen (BMNH 1890.12.1.29) at Palanog on 30
April 1888 during the Steere Expedition.

COMMENTS ON OTHER SPECIMENS COLLECTED
BY THE STEERE EXPEDITION

Below we comment on six species and two genera with doubtful
records or confused history.

Ixobrychus eurhythmus (Swinhoe, 1873): Not previously known from
Panay but a specimen (USNM 160965), supposedly taken on 18 January
1888, very probably came from Panay as the Expedition was there at that
time. Unfortunately, the label says ‘‘Palawan” and ‘“‘Jan. 18” [1888].
Collecting in Palawan seems to have ended about 26 September 1887, but
it could be the date rather than the island that is in error on the label.

Accipiter sp.: Of the various species of small Accipiter, Steere (1890) listed
only Accipiter soloensis (Horsfield, 1821) from Mindanao, but in fact he
probably did not have a specimen of this species from Mindanao. The
BMNH holds four skins of small Accipiter taken on the Steere Expedition,
two coming from Moseley and two from Steere. The two from Moseley
(BMNH_ 1890.12.1.8 from Guimaras and BMNH _ 1890.12.1.9 from
Mindanao), which were almost certainly not available to Steere (1890),
were registered as A. virgatus. The Guimaras specimen could not be found
but the Mindanao bird has been re-examined and is A. virgatus confusus
Hartert, 1910. Neither of Steere’s specimens is from Mindanao, which
may imply that a further specimen exists in another institution. One of
Steere’s, from Luzon (BMNH 1896.4.15.5), is a third specimen of A. v.
confusus and the other (BMNH 1896.4.15.3) has been listed earlier as the
first record of A. soloensis from Guimaras. Steere (1890) omitted these two
specimens or at least did not think that either was A. soloensis. He was
probably confused by the existing literature (see Hartert, 1910) and we
cannot be sure what species he took from Mindanao. However, there are
satisfactory records of A. soloensis from Mindanao (AMNH _ 532719
collected by Waterstradt, and AMNH 532720 collected by Goodfellow)
so that Steere’s (1890) record is not critical.

Porzana fusca (Linné, 1766): Steere (1890) listed P. fusca from Luzon

and Mindanao and P. erythrothorax - now considered the Chinese race of |
P. fusca - from Guimaras. McGregor (1909) apparently doubted this record |
because he omitted it. There is no skin now in the BMNH that throws |

February 28, 1989 Dickinson, et al 15

any light upon this unless it is the juvenile Rallina eurtzonoides (Lafresnaye,
1845) (BMNH 1896.6.6.1265) from Guimaras.

Fulica atra Linné, 1758: Not listed by Steere (1890). There is a specimen
(BMNH 1896.6.6.1442), purportedly from the Steere Expedition, dated
15 April when the Expedition was in Leyte. The words “Laguna” and
“old skin”? appear on the label of this and of Steere Expedition specimens
of Falco peregrinus Tunstall, 1771, (BMNH 1896.4.15.37) and Ninox
scutulata (Raffles, 1822) (BMNH 1896.4.15.54), all seem to have been
bought from a taxidermist. No record can be safely based on any of these
specimens.

Centropus bengalensis (Gmelin, 1788): Steere (1890) listed his Palawan
material as Centrococcyx affinis following Sharpe (1888), who reported that
Whitehead had taken C. affinis in Palawan; C. affinis was later synonymised
with C. bengalensis javanensis (Dumont, 1818). As discussed by Parkes
(1957, 1965) and Mees (1971), the birds of Palawan and the Sulu Islands
are C. b. javanensis and those of other islands C. b. philippinensis Mees,
1971 (of which the type was taken in Mindoro by the Steere Expedition).
Steere (1890) did not list his other material of “‘C. affinis’? but the BMNH
has Steere Expedition birds from Bohol, Cebu, Mindoro and Negros.

Collocalia sp.: Apart from C. troglodytes G. R. Gray & Mitchell, 1845,
over which no mistakes are likely to have been made, Steere (1890) listed
only C. francica (Gmelin, 1789) and did so for Cebu, Mindoro, Negros
and Panay. Hartert (1892) included Steere (1890) records of C. francica
in his synonymy of C. fuciphaga (Thunberg, 1812), probably based solely
on birds in the BMNH taken by Moseley. However, it has long been
known that the Steere Expedition took at least two other species of swiftlets.
In fact, McGregor (1909) credited it with the following; C. Jowz (Sharpe,
1879) from Palawan; C. fuciphaga from Cebu, Mindoro, Negros, Palawan
and Panay; C. germani Oustalet, 1878, from Cebu and Negros; and C.
troglodytes from Guimaras, Marinduque and Negros. The records of C.
lowi (see Ogilvie Grant, 1895) and C. troglodytes are correct, but there
seems to be no basis for the Cebu and Negros listings of C. germani -
upon which McGregor (1909) was probably misled by Oberholser (1906),
who wrote “‘... all Philippine records of both C. francica and C. inexpectata
refer of course to germani.”’

In a separate publication one of us (Dickinson, in press) will review
the larger species of Collocalia that occur in the Philippines; several taxa
will be treated under specific names that differ from duPont (1971). The
Steere Expedition material includes the following:

16 NEMOURIA No. 32

M@ Four generally dark forms:

Li a long winged, square tailed swiftlet with feathered tarsi: BMNH
1890.12.1.87 from Palawan. This has been shown by Ogilvie Grant (1895)
to be Jowi, a race of C. maxima Hartert, 1892, — a species omitted by
duPont (1971).

LI a long winged swiftlet with naked tarst: BANH 1890.12.1.86, 1896.6.6.798
and 799; USNM 161296 (once labelled 161299), 161297 and 161298 from
Palawan. These belong to the form palawanensis Stresemann, 1914. DuPont
(1971) treated this as a race of C. brevirostris.

CL} a medium sized swiftlet with feathered tarsi: BMNH 1890.12.1.88,
1896.6.6.794 and 795 from Negros; this has traditionally been called C.
mearnst Oberholser, 1912, although it was treated by duPont (1971) as a
synonym of the following form.

LI a medium sized swiftlet with naked tarsi: BMNH 1896.6.6.791, 792 and
797 from Cebu; BMNH 1896.6.6.796 from Mindoro; USNM 161295 from
Mindanao (the label shows the collecting date as November 1887 and the
register entry showing June 1888 is an error). These represent amelis
Oberholser, 1906; treated by duPont (1971) as a form of C. vanikorensis
(Quoy & Gaimard, 1830).

M@ Two forms with distinct plumage:

L} a medium sized swiftlet with a buffy rump patch: USNM 161299 from
Panay (once mislabelled 161296); this represents germani, considered a form
of Collocalia fuciphaga. This specimen was the basis for the record by
Bourns & Worcester (1894) but it is not a Menage Expedition specimen.
The record from Panay of Steere (1890)-which this pale rumped bird,
almost certainly not known to Steere, would not represent, is the only
Steere Collocalia francica record we have not been able to substantiate.

Ql a small swiftlet with white underparts: BMNH 1896.6.6.786 and 787 from
Guimaras, BMNH 1896.6.6.789 from Mindanao and BMNH 1896.6.6.788
from Negros all of which are forms of C. esculenta (Linné, 1758).

Parus semilarvatus (Salvadori, 1865): Parkes (1971) discussed the
attribution of an old Negros record to the Steere Expedition by McGregor
(1909), and has pointed out that Steere (1890) did not list it. No supporting
specimen has been found.

Phylloscopus olivaceus (Mosely, 1891): This species was described by —

Moseley (1891) as Abrornis olivacea based on a female (BMNH 1891.4.25.47) |

from Negros and a male (BMNH 1891.4.25.30) from Samar. However,
the female is in fact a specimen of Phylloscopus cebuensis (Dubois, 1900).
Rand & Rabor (1952) remarked upon this when restricting the type locality
to Samar-they also incidentally provided satisfactory records of P. oltvaceus

from Negros. We now consider it desirable to hereby formally designate —

/
i

February 28, 1989 Dickinson, et al 17

the presumed male P. olivaceus (BMNH 1891.4.25.30) taken on 12 April
1888 at Catbalogan, Samar as the lectotype so as to ensure citation of a
type specimen that is truly representative of the species.

ACKNOWLEDGMENTS

Our thanks are first due to those museum staff who have helped us
in tracking down Steere specimens and, sometimes, in identifying them:
Robert Storer and Stephen Goodman at the University of Michigan
Museum of Zoology (whom we particularly thank for copies of several of
Steere’s early articles); Ray Chandler and Elden Martin at Bowling Green
State University; Pierre de Villers at the ISB, Brussels; Mrs. F. M. Benson
and Miss Hilary Potter at the University of Cambridge; Ron Vasile,
Chicago Academy of Sciences; John Fitzpatrick and David Willard at the
Field Museum of Natural History; John N. Condit, Ohio State University
Museum of Zoology; Charles Chase III at the Denver Museum of Natural
History; Raymond Paynter at the Museum of Comparative Zoology; Gerlof
Mees at the Rijksmuseum van Natuurlijke Historie, Leiden; David F.
Parmelee at the Bell Museum of Natural History; Mary LeCroy at the
AMNH; Kenneth C. Parkes, Carnegie Museum of Natural History (to
whom we owe special thanks for assisting with data from Minneapolis and
reviewing early drafts of this paper); Jon C. Barlow, Royal Ontario
Museum; Ian Galbraith, Graham Cowles, Anne Vale and Michael Walters
at the BMNH; Russell Mumford at Purdue University; and Phillip Angle,
Ralph Browning, Storrs Olson, Charles Ross and Richard Zusi at the
USNM. We are also most grateful to Lord Cranbrook for his help over
Collocalia identification.

LITERATURE CITED

Alcala, A. C. & P. Alviola. 1970. Notes on the birds and mammals of Boracay,
Caluya, Carabao, Semirara and Sibay Islands, Philippines. Silliman Journal,
17: 444-454,

Amadon, D. 1951. Notes on Chinese Egret, Egretta eulophotes (Swinhoe). Philippine
Journal of Science, 80: 53-54.

Bourns, F. S. & D. C. Worcester. 1894. Preliminary notes on the birds and
mammals collected by the Menage Scientific Expedition to the Philippine

Islands. Occasional Papers of the Minnesota Academy of Natural Sciences,
1: 1-64.

Dickinson, E. C. 1984. Notes on Philippine birds 1. The status of Porzana

18 NEMOURIA No. 32

paykulli in the Philippines. Bulletin of the British Ornithologists’ Club, 104:
71-72.

— In press. A review of larger Philippine swiftlets of the genus Collocalia.
Forktail.

DuPont, J. E. 1971. Philippine Birds. Delaware Museum of Natural History
Monograph No. 2, Greenville, x + 480 pp.

— 1972. Notes on Philippine birds (No.3): Birds of Marinduque. Nemouria, 7:
1-14,

— 1976. Notes on Philippine Birds (No.4): Additions and corrections to
“Philippine Birds.” Nemouria, 17: 1-13.

Hachisuka, M. 1931. The birds of the Philippine Islands with notes on the
mammal fauna. Vol. 1, pt. 1, Witherby, 168 pp.

Hartert, E. 1892. Catalogue of the Picariae in the collection of the British Museum.
pp. 496-511.

— 1910. Birds of Hainan. Novitates Zoologicae, 17: 189-254.

Marshall, J. T., Jr. 1978. Systematics of smaller Asian night birds based on voice.
Ornithological Monographs, 25: 1-58, + 1 record.

McClure, H. E. 1974. Migration and survival of the birds of Asia. United States
Army Medical Component, SEATO Medical Project, Bangkok, 476 pp.
McGregor, R. C. 1909. A manual of Philippine birds. Bureau of Science, Manila.

No. 2. Part 1, pp. 1-412; Part 2, pp. 413-769.

Mees, G. F. 1971. The Philippine subspecies of Centropus bengalensis (Gmelin)
(Aves, Cuculidae). Zoologische Mededelingen (Leiden), 45 (18): 189-191.

Moseley, E. L. 1891. Descriptions of two new species of flycatchers from the island
of Negros, Philippines. Ibis, (6) 3: 46-47.

Oberholser, H. C. 1912. A revision of the forms of the Edible-nest Swiftlet
Collocalia fuciphaga (Thunberg). Proceedings of the United States National
Museum, 42: 11-20.

Ogilvie Grant, W. R. 1895. On the Birds of the Philippine Islands-Part V. The
Highlands of the Province of Lepanto, North Luzon. Ibis, (7) 1: 433-472.

Parker, S. A. 1981. Prolegomenon to further studies in the ‘“‘Chrysococcyx
malayanus’’? group (Aves. Cuculidae). Zoologische Verhandelingen (Leiden),
187: 1-56.

Parkes, K. C. 1957. Taxonomic notes on the Lesser Coucal, Centropus bengalensis.
Bulletin of the British Ornithologists’ Club, 77: 115-116.

— 1965. Character displacement in some Philippine cuckoos. Living Bird, 4:
89-98.

—— 1971. Taxonomic and distributional notes on Philippine birds. Nemouria, 4: —
1-67. |

— 1986a. Notes on Philippine birds, 9. Reidentification of a unique stint |

February 28, 1989 Dickinson, et al 19

specimen. Bulletin of the British Ornithologists’ Club, 106: 133-136.

— 1986b. Notes on Philippine birds, 10. On the validity of Gerygone sulphurea
rhizophorae Mearns. Bulletin of the British Ornithologists’ Club, 106:
149-152.

Rabor, D. S. 1938. The avifauna of the Gigante Islands. Philippine Journal of
Science, 66: 267-274.

Rand, A. L. & D. S. Rabor. 1952. Notes on Philippine birds. Natural History
Miscellanea (Chicago Academy of Sciences), 107: 1-5.

Sclater, W. L. 1919. [Descriptions of new hawks.] Bulletin of the British
Ornithologists’ Club, 40: 37-42.

Sharpe, R. B. 1877. On the birds collected by Professor J. B. Steere in the
Philippine Archipelago. Transaction of The Linnean Society of London, (6)
1: 307-355.

—— 1888. On a collection of birds from the island of Palawan. Ibis, (5) 6: 193-204.

Steere, J. B. 1888a. A month in Palawan. American Naturalist, 22: 142-145.

—— 1888b. Six weeks in southern Mindanao. American Naturalist, 22: 289-294.

—— 1888c. Observations made in the Central Philippines. American Naturalist,
22: 622-626, 779-784.

— 1889. A month in the Eastern Philippines. American Naturalist, 23: 102-107.

— 1890. A list of the birds and mammals collected by the Steere Expedition to
the Philippines, with localities, and with brief preliminary descriptions of
the supposed new species. Courier Office, Ann Arbor, Michigan, 30 pp.

— 189la. Ornithological results of an expedition to the Philippine Islands in
1887 and 1888. Ibis, (6) 3: 301-316.

— 1891b. A visit to the Philippine Islands of Masbate and Marinduque. American
Naturalist, 25: 731-734.

— 189lc. The island of Mindoro. American Naturalist, 25: 1041-1045.

— 1894. On the distribution of genera and species of non-migratory land-birds
in the Philippines. Ibis, (6) 6: 411-420 and Auk, 11: 231-240.

Tweeddale, Arthur, Marquis of. 1878. Contributions to the ornithology of the
Philippines. No. V. On the collection made by Mr. A. H. Everett in the
island of Negros. Proceedings of the Zoological Society of London, pp.
280-288.

Worcester, D. C. & F. S. Bourns. 1898. Contributions to Philippine ornithology.
Part I. A list of the birds known to inhabit the Philippine & Palawan Islands,
showing their distribution within the limits of the two groups. Proceedings
of the United States National Museum, 20 (1134): 549-566.

Reviewer of this paper: Dr. Kenneth C. Parkes, Carnegie Museum of Natural
History.

President
JOHN E. DU PONT

Managing Editor
RUDIGER BIELER

Consulting Editors
WILLIAM K. EMERSON, American Museum of Natural History, New York,
NY.
M. G. HARASEWYCH, National Museum of Natural History, Smithsonian
Institution, Washington, DC.
KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh,
PA.
RICHARD W. THORINGTON, JR., National Museum of Natural History,
Smithsonian Institution, Washington, DC.

Published by the

DELAWARE MUSEUM OF NATURAL HISTORY
4840 Kennett Pike, Wilmington,
Delaware 19807, U.S.A.

NEMOURIA

Occasional Papers of the Delaware Museum of Natural History

NUMBER 33 SEPTEMBER 20, 1989

NOTES ON THE MENAGE COLLECTION
OF PHILIPPINE BIRDS
1. REVISION OF PACHYCEPHALA CINEREA
(PACHYCEPHALIDAE) AND AN
OVERLOOKED SUBSPECIES OF
DICAEUM TRIGONOSTIGMA (DICAEIDAE)

Kenneth C. Parkes!

ABSTRACT. The Philippine whistlers currently considered conspecific with Pachycephala
cinerea are divided into three species, with only the Palawan race plateni being
retained in cinerea. The other two species are P. homeyeri of the southern and
central Philippines and P. albiventris of Luzon and Mindoro. Study of specimens
from the historic Menage Collection and later material results in the revival of
Hyloterpe major Bourns and Worcester as a valid subspecies of P. homeyeri endemic
to the island of Cebu, Philippines. The population of Dicaeum trigonostigma from
the island of Tablas, Philippines, suggested as probably distinct by Bourns and
Worcester, is described as a new subspecies. Lectotypes are designated for Hyloterpe
winchelli, Hyloterpe major, and Dicaeum intermedia, all of Bourns and Wortester
(1894),

INTRODUCTION

Two of the participants in the pioneering Steere Expedition to the
Philippines (1887-1888), Frank S. Bourns and Dean C. Worcester, were
convinced that much more ornithological collecting was needed in the
archipelago, “‘both in the discovery of new species and in the working out
of the exact distribution of species already known” (Bourns and Worcester,

‘Carnegie Museum of Natural History, 440 Forbes Serial Publication
Avenue, Pittsburgh, Pennsylvania 15213 U.S.A. ISSN 0085-3887

2 NEMOURIA No. 33

1894: 5). Through the generosity of Mr. Louis F. Menage of Minneapolis,
a businessman and member of the Minnesota Academy of Sciences, they
were able to return to the Philippines in the summer of 1890. The
specimens obtained during the following three years were donated to the
Minnesota Academy of Sciences, and the fruits of the expedition are
generally referred to as the “Menage Collection.” Almost 4500 bird
specimens (in addition to numerous mammals) were obtained in the
Philippines, and 81 more birds in Borneo at the end of the expedition.

Bourns and Worcester (1894) published “‘Preliminary Notes”? on the
birds and mammals collected by the Menage Expedition, in which they
described 36 new “species” of birds, most of which are now considered
subspecies and a few as synonyms. They had intended to publish a full
account of their expedition, with field notes, but were unable to find funds
to support such a publication. Their notes were made available to Richard
McGregor of the Philippine Bureau of Science, and are quoted extensively
in McGregor’s “Manual”? (McGregor, 1909).

The Minnesota Academy of Sciences transferred ownership of the
collection, at first to the Minneapolis Public Library where selected
mounted specimens were displayed for many years. Eventually the
collection, including the mounts, was deposited in the Minnesota Museum
of Natural History, University of Minnesota, now called the James Ford
Bell Museum of Natural History (for which the old acronym MMNH_ is
still used). Beginning in 1911, the majority of the Menage Collection was
transferred to the United States National Museum of Natural History
(USNM). Between 1958 and 1963, 539 Menage specimens were exchanged
to Carnegie Museum of Natural History (CM). Several dozen were
exchanged to other museums from MMNH, USNM or CM, and several
dozen more have been destroyed or have yet to be traced.

This is the first of several proposed papers based on my long-term
studies of the Menage Collection. Later papers will reconstruct the
unpublished itinerary of the Menage expedition, and will analyze the type
material of Bourns and Worcester’s new taxa. Bourns and Worcester (1894)
did not designate holotypes, and several of their new taxa were described
from long series. Most of these syntypes are now in the USNM, and were
discussed by Deignan (1961). Unfortunately Deignan made many errors
and unjustified assumptions in connection with the Menage types, which
I hope to clarify in my proposed paper on these type specimens.

In the present paper I call attention to two valid subspecies, one of
which, ‘“‘Hyloterpe major,” was described (as a species) by Bourns and
Worcester (1894:22) but synonymized by McGregor (1909: 603), Deignan
(1961: 471), Mayr (1967: 13), and duPont (1971: 351). In order to
determine the correct name to use for the species to which major belongs,

September 20, 1989 Parkes 3

I present a brief review of this group of whistlers. The other overlooked
subspecies, a race of the highly polytypic flowerpecker Dicaeum trigonos-
tigma, was suggested as possibly separable by Bourns and Worcester (1894:
19) and Deignan (1961: 505).

All measurements are in mm to the nearest 0.5 mm (Bourns and
Worcester published theirs in inches and tenths). I measured the wing
flattened on the ruler, and the bill along the culmen from its base at the
forehead. Altitudes taken from the literature are given in feet as published.

SYSTEMATICS
Pachycephala cinerea (Blyth, 1847) (Pachycephalidae)

In most of the current literature, (Mayr, 1967; duPont, 1971), all of
the Philippine whistlers except Pachycephala philippinensis (Walden, 1872)
are considered conspecific with P. cinerea (Blyth, 1847), which inhabits
southeast Asia from Calcutta to Indonesia. White and Bruce (1986) and
some other authors (references in White and Bruce) advocate the use of
Tephrodornis grisola Blyth, 1843, for this species; Mayr (1967) considered
grisola unidentifiable. I have not studied the nomenclatural arguments, and
for convenience I follow the nomenclature of the standard ‘“Peters”’
check-list (Mayr, 1967).

Delacour and Mayr (1946: 216) had earlier written of the Philippine
populations “Whether or not these 5 forms [albiventris, mindorensis,
crissalis, winchelli, homeyeri| are actually members of a single species, and
whether or not they are conspecific with cinerea (Malaysia) still remains
to be determined.” I have examined specimens of all of the Philippine
races, and have come to the conclusion that they are best treated as three
species.

The first of these is a relatively small, primarily grayish race, plateni
(Blasius, 1888), which is confined to the island of Palawan. It is a lowland
forest bird that is similar to the adjacent Bornean subspecies P. cinerea
secedens Stresemann, 1913, differing chiefly in its browner (less gray)
crown. It is the only Philippine race that I would continue to assign to
the species cinerea. This makes zoogeographic sense, as the birds of Palawan
are frequently more closely related to those of Borneo than to those of the
main Philippine archipelago (Delacour and Mayr, 1946).

The second species is characterized by distinctly rufous plumage
colors, absent in P. cinerea. Part of its range consists of quite small islands,
SO it cannot be said to be confined to highlands, but on Canlaon Volcano,
Negros, it occurs from 2500 feet to 7500 feet elevation (Ripley and Rabor,

& NEMOURIA No. 33

1956). Members of this reddish group inhabit the Sulu Archipelago and
the central Philippine islands of Cebu, Masbate, Negros, Panay, Sibuyan,
Tablas, and Ticao. The oldest available name for this group of subspecies
is homeyeri Blasius, 1890 (type locality Jolo Island, Sulu Archipelago).
Subspecific variation in Pachycephala homeyeri will be discussed below.

The third species is found in the mountains of the northern
Philippines, from 2500 feet (762 meters) upward, and is quite different in
appearance. It is greenish dorsally and gray below, with yellow under tail
coverts. There are three subspecies in this species, with the oldest name
being albiventris Ogilvie Grant, 1894, for the race of northern Luzon. This
name has six months priority over mindorensis Bourns and Worcester, 1894,
the name for the valid Mindoro race. The third subspecies, P. a. crissalis
Zimmer, 1918, appears to be confined to the vicinity of Mount Banahao,
Laguna Province, south-central Luzon, where the holotype and paratype
were collected at 3000 feet elevation.

The English name for P. cinerea (s.l.) in many handbooks is
‘““Mangrove Whistler.”? Although primarily found in mangroves in much
of its non-Philippine range, this species also inhabits lowland and foothill
forests. In most of the Philippines, however, the whistlers of this group
are chiefly birds of mountain forests. With the division of the former
Pachycephala cinerea into three species, I propose retaining the English
name Mangrove Whistler for P. cinerea in the restricted sense used here;
even though not wholly appropriate for some populations, it has neverthe-
less been so widely used that I do not venture to recommend any change.
White-bellied Whistler, as used by Delacour and Mayr (1946) and duPont
(1971) for the combined Philippine forms, I would retain for P. homeyert.
An appropriate new English name for P. albiventris, not previously used
in this very large genus, is Green-backed Whistler.

Bourns and Worcester (1894) described three species in the genus
Hyloterpe, now considered a synonym of Pachycephala. Of these, one (H.
mindorensis), is a subspecies of P. albiventris, and one (H. winchelli) of P.
homeyert. The third, Hyloterpe major, has been synonymized with winchelli
by McGregor (1909), Mayr (1967), and duPont (1971).

Bourns and Worcester (1894) attributed major to the Philippine islands
of Cebu, Tablas, and Sibuyan. They commented that this was a somewhat
peculiar distribution, as it was interrupted by the range they attributed to
winchelli, namely Negros, Panay, and Masbate. Furthermore, they called
attention to differences in both size and color between Cebu and
Tablas/Sibuyan specimens of major, but stated that they did “‘not think
[Tablas and Sibuyan populations] can be specifically separated.” This
represented a change of opinion for these authors, as both in the original
catalogue of the Menage Collection and on the specimen labels, the Tablas

September 20, 1989 Parkes 5
and Sibuyan specimens are identified as ““Hyloterpe minor,” a name they
never published, although in their catalogue they went so far as to designate
a male from Tablas (their no. 1454, now USNM 316148) and a female
from Sibuyan (their no. 1461, now USNM 316147) as “types” of ‘‘minor.”’

I have restudied the Menage Collection material of this species,
supplemented by additional specimens at the USNM. I find the name
major Bourns and Worcester to be valid for a subspecies of Pachycephala
homeyeri restricted to the island of Cebu alone. The universal failure to
recognize major is undoubtedly based on its authors’ allocation of the islands
of Tablas and Sibuyan to its range.

As originally stated by its authors, major of Cebu is substantially
larger than winchelli (see measurements below), and differs slightly in
color. Bourns and Worcester stated that major differs from winchelli in ‘‘the
white of the throat less sharply defined;’’ my notes taken at USNM state
that Cebu specimens are ‘‘more heavily washed with brownish across the
chest”’ than are winchelli from Negros, Panay, and Masbate.

Bourns and Worcester (1894: 21) gave the distribution of winchelli as
Panay, Masbate and Negros, and did not mention any variation within
their series. I find that specimens from Negros average slightly longer-tailed
than those from the other islands, but the overlap is extensive, and
measurements of the series from the three islands are combined for the
comparisons below. Color variation within winchelli is puzzling; as Ralph
Browning (pers. comm.) has also noted, the Menage specimens from
Negros are more reddish (less brownish) on the sides of the head than
those from other islands, but this difference is not apparent in Negros
specimens collected in the 1960’s.

Deignan (1961: 471) listed USNM 316149 from Negros as the “‘type”’
of winchelli, but his discussion of the Bourns and Worcester series was
based on several false premises that I intend to clarify in my future paper
on the Menage type material. Nevertheless, in view of the slight geographic
variation within winchelli it would be useful to restrict the type locality of
the subspecies. I therefore formally designate as the lectotype of Hyloterpe
winchellt Bourns and Worcester, 1894, the same specimen listed by Deignan
(1961), namely USNM 316149 (Menage 1440), adult male, collected by
D. C. Worcester and F. S. Bourns at Bais, Negros, Philippine Islands,
on 12 January 1891. I also formally designate as the lectotype of Hyloterpe
major Bourns and Worcester the “‘type”’ listed by Deignan, USNM 316145
(Menage 1450), adult male, collected by Worcester and Bourns at Toledo,
Cebu, Philippines, on 16 June 1892.

The status of the populations of Tablas and Sibuyan remains to be
discussed. These are the birds for which Bourns and Worcester used but
later abandoned the manuscript name “minor”. In their publication (1894:

6 NEMOURIA No. 33

22) they included these birds in their new “species” major, although
commenting on their smaller size. They gave only the measurement means
for their entire series of winchelli, and separate means for their Cebu and
Tablas series attributed to major. In remeasuring all of this material, I
find that sex for sex (males being larger), the wing lengths of Cebu
specimens do not even overlap with those from any of the other islands
under consideration (see table). There is also a gap in the tail measurements
of males, although in females there is a slight overlap. The only discrepancy
lies in the bill measurements. Contrary to Bourns and Worcester, the tail
and especially the wing measurements of Tablas and Sibuyan birds,
although averaging slightly larger than those of winchelli, more closely
match that race rather than Cebu major. The bills of the Tablas/Sibuyan
series, however, match those of major in length. The Tablas/Sibuyan
populations could conceivably be separated as an additional subspecies,
with the wing and tail measurements of winchelli and the bill measurement
of major. I prefer to consider those populations merely as the long-billed
extremes of winchelli rather than recognize a third subspecies, as they also
match winchelli in color. Combining these populations with winchelli rather
than major also corrects the apparent zoogeographic anomaly of a split
range for major mentioned by Bourns and Worcester.

Measurements: winchelli (Negros, Panay, Masbate; n=22) wing,
males 83-89 (85.6); females 79.5-86 (83.5). Tail, males 64.5-71.5 (67.8);
females 62-71.5 (66.6). Bill, males 17-18.5 (17.8); females 16.5-19 (17.6).

winchelli (Tablas, Sibuyan [’minor’’]; n=10) wing, males 86-89.5
(87.8); females 81.5-90 (85.1). Tail, males 70-72.5 (71.4); females 65.5-72.5
(68.2). Bill, males 19.5-21.5 (20.3); females 18.5-20 (19.0).

major (Cebu; n=8) wing, males 94.5-99 (96.5); females 90-94.5 (91.5).
Tail, males 72.5-78 (75.8); females 70.5-73 (72.0). Bill, males 20-21.5 (20.6);
females 19.5-20.5 (20.2).

Dicaeum trigonostigma (Scopoli, 1786) (Dicaeidae)

In their description of Dicaeum intermedia [sic], Bourns and Worcester
(1894: 19) gave the range as Romblon and Tablas. They implicitly restricted
the type locality to Romblon, however, in stating “‘It may ultimately prove
that the Tablas birds are distinct, the four specimens secured by us in
that island having a much heavier wash of yellow on the throat than the
Romblon birds.” The reference to “four specimens secured” is an error
even if intended to refer to adult males only, as the Menage Expedition
collected five such males. In all, they collected eleven specimens of this
species on Tablas, of which five are now in USNM, two in CM, one in

September 20, 1989 Parkes 7

MMNH, and three are unaccounted for.

Deignan (1961: 506) quoted Bourns and Worcester’s statement about
the Tablas birds, and formally restricted the type locality of Dicaeum
intermedia to Romblon. He listed only one of the syntypes from Romblon,
erroneously stating that it was ““Bourns and Worcester’s own lectotype.”’
According to the International Code of Zoological Nomenclature (Interna-
tional Commission on Zoological Nomenclature, 1985), Articles 72 (b) (vii)
and 73 (b) (i), “The mere citation of “Type’... in a list of types...
or on a label is not to be construed alone as evidence that a specimen is
or is fixed as any of the kinds of types referred to in this Chapter .. .
Syntypes may include specimens labelled ‘cotype’ or ‘type’. . .”’ There
are in fact four specimens from the original syntypical series of Dicaeum
intermedia in the USNM on which the word “type”? appears, two each
from Romblon and Tablas. The same notation appears, in the same
handwriting, with the entries for these four specimens in the Menage
Collection catalogue. However, the handwriting is not that of the original
labels and catalogue entries themselves, and appears to be that of W. A.
Bryan, who signed himself in the Menage Collection catalogue in 1907 as
“Curator Pac. Orn., Minn. Ac. Sci.”? Bourns and Worcester wrote “‘sp.
n.”’ on the labels of each of their specimens of their new taxa, some of
which bear red original labels and others white. Of the syntypes of
intermedia, three from Romblon and three from Tablas bear red original
labels. Although all of the series rank formally as syntypes, it can be
deduced that Bourns and Worcester gave special status to the red-labelled
specimens, perhaps intending them as “‘cotypes,” although this designation
was never published. The specimen listed by Deignan as the Bourns and
Worcester “‘lectotype” of Dicaeum intermedia, USNM 316172 (Menage
2699), bears a white label, whereas USNM 316182 (Menage 2698) bears
a red original label, and also bears the later entry as “type” in the same
(Bryan) handwriting as that on the alleged lectotype. It appears more
appropriate to designate formally as lectotype of Dicaeum intermedia one
of the specimens to which Bourns and Worcester had attached a red label,
so I therefore so designate USNM 316182 (Menage 2698), adult male,
collected on Romblon Island, Philippines, 6 September 1892 by the
Menage Expedition.

Bourns and Worcester and Deignan were correct in believing that the
specimens from Tablas are separable from Dicaeum trigonostigma interme-
dium of Romblon. In addition to his tentative suggestion in his USNM
type catalogue (Deignan, 1961: 506), Deignan left a handwritten note in
the tray containing the USNM specimens from Tablas, reading ‘‘Should
be named — like intermedium but throat washed with yellow (HGD).”
The Tablas subspecies is appropriately named as follows:

8 NEMOURIA No. 33
Dicaeum trigonostigma cnecolaemum, new subspecies

HOLOTYPE: USNM 315770, adult male, collected at ‘“Badajos”’
[=Badajoz], Tablas Island, Philippines, on 29 September 1892, by D. C.
Worcester and F. S. Bourns (Menage Expedition no. 2690 !/2).

DIAGNOSIS: Similar to D. t. intermedium of Romblon Island, but
adult males with throat washed with yellow instead of being pale neutral
gray, and breast slightly deeper orange-yellow; blue of dorsum slightly
less blackish, especially on the forehead. Immature males have chins
distinctly yellow rather than gray as in imtermedium, and the green of the
dorsum is slightly less grayish. No females from Romblon were available,
but females from Tablas are yellowish on the throat, and the subspecific
differences probably parallel those of the immature males. The two
subspecies do not differ in size.

RANGE: The island of Tablas, Philippines.

ETYMOLOGY: From the Greek, knekos, meaning pale yellow, and
laimos, meaning throat, the distinguishing character of the subspecies in
all known plumages.

SPECIMENS EXAMINED: D. t. cnecolaemum, adult males 6,
immature males 2, adult females 2, unsexed immature 1. D. t. intermedium,
adult males 5, immature males 2.

ACKNOWLEDGMENTS

The exchange whereby CM obtained Menage Collection specimens
was arranged by Dwain W. Warner and by Robert W. Dickerman, who
was then a graduate student at the University of Minnesota. David F.
Parmelee of the Bell Museum of Natural History generously allowed Dr.
Dickerman to hand-carry the Menage Collection catalogue from Minneapo-
lis for me, and was most hospitable while I was studying the Menage
specimens at the Bell Museum. Specimens were examined at the USNM
through the courtesy of the curatorial staff at that museum. The manuscript
was critically read by M. R. Browning, E. C. Dickinson, R. S. Kennedy,
C. A. Ross, and D. S. Wood.

LITERATURE CITED

Bourns, F. S., & D. C. Worcester. 1894. Preliminary notes on the birds and
mammals collected by the Menage Scientific Expedition to the Philippine
Islands. Occasional Papers of the Minnesota Academy of Natural Sciences,
1 (1): 1-64. |

September 20, 1989 Parkes 9

Deignan, H. G. 1961. Type specimens of birds in the United States National
Museum. Bulletin of the United States National Museum, 221: x + 718 pp.

Delacour, J.. & E. Mayr. 1946. Birds of the Philippines. Macmillan, New York,
xv + 309 pp.

DuPont, J. E. 1971. Philippine Birds. Delaware Museum of Natural History
Monograph No. 2, Greenville, x + 480 pp.

International Commission on Zoological Nomenclature. 1985. International Code
of Zoological Nomenclature, third edition. International Trust for Zoological
Nomenclature, London, xx + 338 pp.

Mayr, E. 1967. Subfamily Pachycephalinae. In: Check-list of Birds of the World,
vol. 12 (R. A. Paynter, Jr., ed.). Museum of Comparative Zoology,
Cambridge, Mass., x + 495 pp.

McGregor, R. C. 1909. A Manual of Philippine Birds. Bureau of Science, Manila,
x + 769 pp.

Ripley, S. D., & D. S. Rabor. 1956. Birds from Canlaon Volcano in the highlands
of Negros Island in the Philippines. Condor, 58: 283-291.

White, C. M. N., & M. D. Bruce. 1986. The birds of Wallacea. British
Ornithologists’ Union Check-list no. 7, London, 524 pp.

NOTE ADDED IN PROOF: Upon further investigation, I find that
Tephrodornis grisola Blyth, 1843, must indeed replace M/uscitrea] cinerea
Blyth, 1847, as the name for the Mangrove Whistler. Mayr and others
who have considered grisola unidentifiable were apparently not aware that
Blyth’s holotype is extant. See Mukherjee, 1970, Journal of the Bombay
Natural History Society, 67: 112-113.

9

7s

.—

ae

President
JOHN E. DU PONT

Managing Editor
RUDIGER BIELER

Consulting Editors
WILLIAM K. EMERSON, American Museum of Natural History, New York,
NY.
M. G. HARASEWYCH, National Museum of Natural History, Smithsonian
Institution, Washington, DC.
KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh,
PA.
RICHARD W. THORINGTON, JR., National Museum of Natural History,
Smithsonian Institution, Washington, DC.

Published by the

DELAWARE MUSEUM OF NATURAL HISTORY
4840 Kennett Pike, Wilmington,
Delaware 19807, U.S.A.

QL
lev /

t

NEMOURIA

Occasional Papers of the Delaware Museum of Natural History

NUMBER 34 DECEMBER 28, 1989

JOHANNES THIELE AND HIS CONTRIBUTIONS
TO ZOOLOGY.
PART 1. BIOGRAPHY AND BIBLIOGRAPHY

Riidiger Bieler! and Kenneth J. Boss?

ABSTRACT.. The first part of an evaluation of Johannes Thiele’s (1860-1935)
contributions to zoology provides a biographical sketch of his life and a complete
listing of his publications with special emphasis on the exact dates of publication.
During his professional career, most of which was spent at the Zoological Museum
in Berlin, Germany, Thiele produced 155 zoological publications. The majority of
his papers (129) dealt with the systematics and anatomy of Mollusca, but he made
numerous important contributions to the study of Crustacea (12) and Porifera (8).
Six publications treated miscellaneous zoological subjects.

INTRODUCTION AND BIOGRAPHY

One of the preeminent malacologists of the twentieth century, Johannes
Thiele, has been an inspiration to generations of students of the Phylum
Mollusca. Probably no other individual researcher has been more influen-
tial, and his Handbuch der systematischen Weichtierkunde has left an indelible
imprint on innumerable investigations in malacology. Not only did this
work treat thousands of generic-level names with individual diagnoses, but
it placed them in a newly established hierarchy of higher taxa. At the time,
this magnum opus was the only work which dealt extensively with anatomical
characteristics as well as conchological or hard-part features; it replaced
Fischer’s Manuel de Conchyliologie of fifty years before. With a detailed
acquaintance of most groups and a profound knowledge of the diverse
literature on the systematics and anatomy of mollusks, Thiele was

\Delaware Museum of Natural History Serial Publication
2Museum of Comparative Zoology, Harvard University, ISSN 0085-3887

Cambridge, Massachusetts 02138, U.S.A.

No. 34

NEMOURIA

December 28, 1989 Bieler & Boss S)

fortunately well prepared for his task. For over half a century, he published
critical studies on many diverse molluscan taxa from widely differing
geographical regions of the world, prompting Winckworth (1938) to remark:
“In no group, in no fauna was his knowledge superficial; steady work
prosecuted with that thoroughness in detail that is akin to genius made him
the great master of his subject.”

That one man could have intellectually embraced the entire diversity
of the Phylum Mollusca in such detail seemed so astonishing to us that
we resolved to review his career and research contributions and to compile
a comprehensive list of his publications and the new molluscan taxa, both
generic and specific, which he introduced. The decision to accomplish this
task was attended by a curious coincidence: we began searching for remarks
and obituaries on Thiele, came across Lamy’s Nécrologie (1938) on 5
August 1985, and read “Ses derniéres années furent consacrées 4
Pélaboration de son grand ouvrage Handbuch der systematischen Weichtierkunde
(1931-1935), qu’il put achever quelques mois avant sa mort survenue le 5
aout 1935.” Thus, on the fiftieth anniversary of Thiele’s death, we initiated
our investigation of his life and work.

In addition to Lamy’s obituary in the Journal de Conchyliologie (1938),
Winckworth (1938) published one in the Proceedings of the Malacological
Society of London which also provided a supplementary list to the
Festnummer in the Archiv fiir Molluskenkunde issued by Rensch (1930)
on the occasion of Thiele’s seventieth birthday. Rensch’s delineation of
Thiele’s malacological publications numbered 93 items while Winckworth
provided some additions which were overlooked and completed the list to
include papers published by Thiele from 1930 to his death in 1935. More
recently, Kiss (1985) contributed a short vignette in Hungarian on the life
of Thiele.

Emil Karl Hermann Johannes Thiele was born on 1 October 1960 in
Goldap (Goldapp), then in East Prussia, not far from the Masurian Lakes
and now part of Poland. As a young student, he showed an avid interest
in animals, particularly insects. His higher education was pursued in the
natural sciences, initially at Berlin and later at Heidelberg. Studying under
Professor Franz Eilhard Schulze, Director of the Zoological Institute in
Berlin and an expert on sponges, he produced his thesis in 1886 on the
nature of labial palps in bivalve mollusks. This consisted of a detailed
investigation of these structures and their probable function in many
families of lamellibranchs. Later in 1886, he spent six months at the
Zoological Institute in Naples. Also at about this time he met Miss Fanny
Wolle, whom he married in October 1888 and who, along with his daughter
and three sons, survived after his death.

In 1891, he accepted an appointment as an assistant in the invertebrate

4 NEMOURIA No. 34

section of the Dresden Museum where he was mainly occupied in arranging
the collection of mollusks. At about this time he commenced working on
the unfinished manuscripts of 1879 left by Franz Hermann Troschel, the
originator of Das Gebiss der Schnecken, and began his interest in the study
of Solenogastres which culminated in a number of subsequent studies [he
did not consider them to be mollusks and therefore did not treat them in
such works as the Handbuch]. By July 1895, Thiele had taken up residence
at Strassburg’s Zoological Institute where he was responsible for studying
and arranging the Japanese sponges collected by Ludwig Déderlein during
his tenure as Professor of Natural Sciences in Tokyo. In October 1896 he
moved on to Gédttingen’s Zoological Institute where he remained only
briefly since by the spring of 1898, he took a position as a Lecturer in the
Faculty of Entomology at the Zoological Institute of the Agricultural
College in Berlin. Thiele apparently never actually lectured here, because
he directly moved on to the Zoological Museum in Berlin in the spring
of 1899, where he remained for the rest of his professional career. At first
he was a scientific collaborator or helper (wissenschaftlicher H1lfsarbeiter),
then in 1900 a Research Assistant, and by 1903, Head or Keeper (Kustos)
of the Crustacea Section. In 1905, after the death of Eduard von Martens,
he took charge of the Mollusk Section. Reaching the mandatory age of
retirement, Bismarck’s Altersgrenze, he formally left his position as
Professor of Zoology in 1925 but actively continued research until his death
in 1935.

As can be seen from the following bibliography, Thiele sustained an
enviable outpouring of papers and monographs. To certain groups he paid
particular attention, contributing significant items on the Solenogastres and
the phylogeny of mollusks (1902b). He wrote a detailed revision of
Solenogastres in Das Tierreich (1913e), as well as a two-part monograph
of chitons (1909d,1).

For Martini-Chemnitz’s second edition of the Conchylien-Cabinet, he
pursued several different groups such as the Cocculinoidea (1909g),
Scissurellidae and Fissurellidae (1912a, 1913f, 1916b, 1919), and Limidae
(1918, 1920a). Taxonomic monographs on other individual families or
superfamilies included his revisions of the Trochacea (1924a), Columbelli-
dae (1924b), Assimineidae (1927a), and the Hydrobiidae and Melaniidae
(1928c). As for geographically circumscribed faunal reports, mention should
be made of his studies of the land and freshwater shells of Borneo (with
E. von Martens, 1908a), Brazil (1927c), central Africa (1911d), Abyssinia
(1933c), the West Indies (1910f), and southwest Australia (191le). He was
an outstanding contributor to several of the great German Imperial
expeditions, having studied both the gastropods and bivalves of the German
Deep-Sea or Valdivia Expedition; the first portion of this work was in

December 28, 1989 Bieler & Boss 5

collaboration with Eduard von Martens (1904a). A subsequent long and
extensive study was completed on the smaller and more obscure gastropods
(1925b), illustrated by Thiele’s own drawings. The bivalves were monogra-
phed in collaboration with Jaeckel (1931b). Additionally, he produced the
reports on most of the mollusks obtained by the German South Polar
Expedition (1908c, 1912b, 1920b).

In further preparation for the Handbuch der systematischen Weichtierkunde
and for his authoritative contributions to the Handworterbuch der Naturwis-
senschaften and to Ktikenthal’s Handbuch der Zoologie, Thiele compiled a
list of molluscan nomina conservanda, which, although not put before the
International Committee for authorization, was published by Apstein
(1915c). According to Winckworth (1938), he also prepared the list of
molluscan generic names for inclusion in F.E. Schulze & W. Kiikenthal’s
Nomenclator animalium generum et subgenerum.

In this first of a series of papers on Thiele and his contributions, we
provide a revised bibliography of his publications on mollusks and other
animal groups, adding several which were overlooked by both Rensch
(1930) and Winckworth (1938). We were aided in compiling our inventory
not only by our colleagues’ previous work but also by careful appraisals
of entries in the reviews of literature in the Archiv ftir Naturgeschichte,
Bibliographia Zoologica, Zentralblatt fiir Zoologie, Zoological Record, Zoologi-
scher Fahresbericht, and Zoologisches Centralblatt. We also consulted lists
of publications given by Buschmeyer (1938), Scheele and Natalis (1981),
Btirk and Jungbluth (1985), and Vosmaer (1928).

BIBLIOGRAPHY

Thiele’s bibliography comprises 155 papers, some of which are parts
of works published over a period of time. His journal publications appeared
almost exclusively in German serials, the majority in general biological or
zoological journals [e.g., Zeitschrift fiir wissenschaftliche Zoologie, Zoologischer
Anzeiger, Zoologisches Centralblatt (all published in Leipzig), and the Archiv
fiir Naturgeschichte (Berlin)]. The only exceptions were a contribution on
the chitons of the Antarctic Expedition of Dr. Charcot, which appeared (in
two versions) in French journals (1906e,f), a contribution to the reports
on the Albatross Expedition in the Bulletin of the Museum of Comparative
Zoology (1908b), and a paper on Solenogastres of the Russian Polar
Expedition in a St. Petersburg journal (1911f). All but the 1906 French
papers were published in German (an English translation of one of his
works on the branchial sense organs of Patellidae was subsequently
published in The Annals and Magazine of Natural History, 1893b).

6 NEMOURIA No. 34

Thiele’s publications are listed below in four sections, as papers on
Mollusca, Crustacea, Porifera, and miscellaneous zoological topics. Empha-
sis was placed on the malacological contributions, i.e., a paper is listed in
that section whenever Mollusca are involved, even though it may contain
data on other phyla. Examples are Thiele’s discussion on whether
chaetognaths are to be considered mollusks (1907b) and his publication
on the phyletic interrelationships of several unsegmented animal groups
(1922).

Throughout the listing a uniform spelling was used for the Nachrichtsblatt
der Deutschen Malakozoologischen Gesellschaft, which in the original was
frequently and inconsistantly modified between volumes, numbers and
off-prints.

As will be shown in later parts of this study, Thiele introduced a large
number of new taxonomic names. Not all of them are clearly designated
as such, and taxonomically important statements can be found in odd
places, such as indices of book publications (193lc) and footnotes of
seemingly unrelated works (1890b). A record for obscurity was set when
he proposed replacement names in footnotes of a literature review (1913a)
covering one of his earlier papers (1912b). In the following sections, an
asterisk (*) indicates publications containing new species- and genus-group
names.

In addition to standard citations (title, book or serial reference, place
and date of publication), English translations of the titles and additional
information (e.g., subsequent reviews) are provided. Special effort has been
made to determine the exact publication dates. In cases of journal
publications these were usually readily available. However, several book
publications and especially the taxomically important Valdivia Expedition
volumes were only dated by year. Since the publishing house Gustav
Fischer Verlag, where most of Thiele’s books and monographs were
published, was unable to provide more exact dates, we are supplying
ancillary information (such as the dates of receipt by major libraries) for
the earliest date “‘on which the work is demonstrated to be in existence
as a published work”? [ICZN Article 21(c)].

December 28, 1989 Bieler & Boss d

A. Publications on Mollusca

1886. Die Mundlappen der Lamellibranchiaten. Zeitschrift ftir wissen-
schaftliche Zoologie, 44: 239-272, pls. 17-18; Leipzig, 1 October
1886 [separately issued as Ph.D. dissertation (Inaugural-
Dissertation zur Erlangung der philosophischen Doktorwtirde
der Philosophischen Fakultéat der Friedrich-Wilhelms-
Universitat Berlin): 34 (+2) pp., pls. 17-18; Leipzig (Wilhelm
Engelmann); before 24 June 1886)]. [The labial palps of
lamellibranchs.] Reviews: Journal of the Royal Microscopical
Society, London, 1887: 200 (English); Zoologischer Jahresbericht
(Berlin) ftir 1886 (1888) (Mollusca): 5, 24 (German).

1887. Ein neues Sinnesorgan bei Lamellibranchiern. Vorldufige
Mitteilung. Zoologischer Anzeiger, 10(257): 413-414; Leipzig,
1 August 1887. [A new sense organ in lamellibranchs. Prelimi-
nary report.] Reviews: Journal of the Royal Microscopical
Society, London, 1887: 942 (English); Zoologischer Jahresbericht
(Berlin) ftir 1887 (1888) (Mollusca): 5, 17 (German).

1889. Die abdominalen Sinnesorgane der Lamellibranchier. Zeitschrift
fiir wissenschaftliche Zoologie, 48(1): 47-59, pl. 4; Leipzig, 5
April 1889. [The abdominal sense organs of lamellibranchs.|
Reviews: Journal of the Royal Microscopical Society, London,
1889: 374-375 (English); Zoologischer Jahresbericht (Berlin)
fiir 1889 (1891) (Mollusca): 8, 27 (German).

*1890a. Uber Sinnesorgane der Seitenlinie und das Nervensystem
von Mollusken. Zeitschrift fiir wissenschaftliche Zoologie,
49(3): 385-432, pls. 16-17; Leipzig, 14 February 1890. [On the
sense organs of the lateral line and the nervous system of
mollusks.] Review: Zoologischer Jahresbericht (Berlin) ftir 1890
(1892) (Mollusca): 7, 9 (German).

1890b. Die Stammesverwandtschaft der Mollusken. Ein Beitrag zur
Phylogenie der Tiere. Jenaische Zeitschrift fiir Naturwissen-
schaft, 25 (= new series 18) (3-4): 480-543; Jena, 30 May
1890. [The phyletic relationships of the mollusks. A contribution
to the phylogeny of animals.] Reviews: Journal of the Royal
Microscopical Society, London, 1891(6): 721-722 (English);
Zoologischer Jahresbericht (Berlin) fiir 1891 (1893) (Mollusca):
14, 20 (German).

8 NEMOURIA No. 34

1890c. Erwiederung. Zoologischer Anzeiger, 13(346): 559-561; Leipzig,
13 October 1890. [A reply (to Rawitz’s criticism).] Review:
Zoologischer Jahresbericht (Berlin) ftir 1890 (1892) (Mollusca):
7, 12 (German).

189la. Bericht tiber die wissenschaftlichen Leistungen im Gebiete
der Malakologie wahrend des Jahres 1888. I. Bericht
liber Allgemeines, Physiologie, Anatomie und Entwick-
lungsgeschichte. Archiv ftir Naturgeschichte, 55(2.1) (1889):
378-428; Berlin, October 1891. [Report on the scientific work
in the realm of malacology during the year 1888. I. Report on
general subjects, physiology, anatomy and embryology. ]

1891b. Das Integument der Chitonen. Biologisches Centralblatt, 11(23):
722-726; Leipzig, 15 December 1891. [The integument of
chitons.] Review: Zoologischer Jahresbericht (Berlin) fiir 1891
(1893) (Mollusca): 7, 20 (German).

*1891c. [Fortsetzung von / Continuation of:] Troschel, F. H.: Das Gebiss
der Schnecken zur Begriindung einer natiirlichen Classifi-
cation. Vol. 2(7) [Rhipidoglossa cont., Docoglossa]: 249-334,
pls. 25-28; Berlin (Nicolaische Verlags-Buchhandlung), 1891.
[The dentition of snails as a foundation for a natural classifica-
tion.]| Reviews: Zoologischer Jahresbericht (Berlin) fiir 1891
(Mollusca): 7, 37 (German); Nachrichtsblatt der Deutschen
Malakozoologischen Gesellschaft, 26(1-2) (1894): 25-27 (Ger-
man). [continued in 1893e]

1892a. Beitrage zur Kenntnis der Mollusken. I. Uber das Epipo-
dium. Zeitschrift fiir wissenschaftliche Zoologie, 53(4): 578-590,
pl. 23; Leipzig, 5 April 1892. [Contributions to the knowledge
of mollusks. I. On the epipodium.] Reviews: Journal of the
Royal Microscopical Society, London, 1892(3): 351 (English);
Zoologischer Jahresbericht (Berlin) fiir 1892 (1893) (Mollusca):
8, 13 (German).

1892b. Vortrag tiber Wurmmollusken. Sitzungsberichte und Abhand-
lungen der Naturwissenschaftlichen Gesellschaft Isis in Dres-
den, 1892 (Jan.-Jun.): 3-4; Dresden, 1892 (‘‘1893” on title
page; received by Museum of Comparative Zoology, Cambridge:
29 October 1892). [Discourse on vermiform mollusks. ]

1892c. Zur Phylogenie des Byssusapparats der Lamellibranchier.
Verhandlungen der Deutschen Zoologischen Gesellschaft auf
der zweiten Jahresversammlung zu Berlin, den 8. bis 10. Juni

December 28, 1989 Bieler & Boss 9

1892: 52-57; Leipzig, 1892. [On.the phylogeny of the byssal
apparatus of lamellibranchs.] Review: Zoologischer Jahresbericht
(Berlin), 1892 (Mollusca): 8, 31 (German).

1892d. Beitrage zur Kenntnis der Mollusken. II. Uber die Mollus-

1893a.

1893b

1893c.

1893d

kenschale. Zeitschrift fiir wissenschaftliche Zoologie, 55(2):
220-251, pl. 11, 1 text-fig.; Leipzig, 20 December 1892.
[Contributions to the knowledge of mollusks. II. On the
molluscan shell.] Reviews: Journal of the Royal Microscopical
Society, London, 1893(2): 164 (English); Zoologischer
Jahresbericht (Berlin) ftir 1892 (1893) (Mollusca): 8, 13 (Ger-
man).

Uber die Kiemensinnesorgane der Patelliden. Zoologischer

Anzeiger, 16(412): 49-50; Leipzig, 13 February 1893. [On the
branchial sense-organs of the Patellidae.] Reviews: Journal of
the Royal Microscopical Society, London, 1893(5): 618 (En-
glish); Zoologischer Jahresbericht (Berlin) ftir 1893 (1894)
(Mollusca): 7, 36 (German). [Republished in English, 1893b]

. On the branchial sense-organs of the Patellidae. The Annals

and Magazine of Natural History, (6)11(65): 413-414; London,
May 1893. [English translation of 1893a]

Ueber das Kriechen der Schnecken. Sitzungsberichte und

Abhandlungen der Naturwissenschaftlichen Gesellschaft Isis in
Dresden, 1892 (Jul.- Dec.): 72-75; Dresden, 1893 (received by
Museum of Comparative Zoology, Cambridge: 8 June 1893).
[On the crawling of snails.] Review: Zoologischer Jahresbericht
(Berlin) fiir 1893 (1894) (Mollusca): 7, 29 (German).

. Bericht tiber die wissenschaftlichen Leistungen im Gebiete

der Malakologie wahrend des Jahres 1889. I. Bericht
liber Allgemeines, Physiologie, Anatomie u. Entwicklung.
Archiv fiir Naturgeschichte, 56(2.1) (1890): 381-424; Berlin,
June 1893. [Report on the scientific work in the realm of
malacology during the year 1889. I. Report on general subjects,
physiology, anatomy and embryology. ]

*1893e. [Fortsetzung von / Continuation of:] Troschel, F. H.: Das Gebiss

der Schecken zur Begriindung einer natiirlichen Classifica-
tion. Vol. 2(8) [Docogiossa cont., Lepidoglossa]: 336-409, pls.
29-32; Berlin (Nicolaische Verlags-Buchhandlung), 1893. [The
dentition of snails as a foundation for a natural classification. ]
Reviews: Zoologischer Jahresbericht (Berlin) fiir 1893 (1894)

10 NEMOURIA No. 34

(Mollusca): 7, 36 (German); Nachrichtsblatt der Deutschen
Malakozoologischen Gesellschaft, 26(1-2)(1894): 25-27 (Ger-
man). [continued from 1891c]

*1894a. Ueber die Zungen einiger Landdeckelschnecken. Nachrichtsblatt
der Deutschen Malakolozoogischen Gesellschaft, 26(1-2): 23-25;
Frankfurt am Main, January 1894. [On the radulae of a few
operculate land snails. ]

*1894b. Beitrage zur vergleichenden Anatomie der Amphineuren.
I. Uber einige Neapeler Solenogastres. Zeitschrift fiir
wissenschaftliche Zoologie, 58(2): 222-302, pls. 12-16, 3 text-
figs.; Leipzig, 7 August 1894. [Contributions to the comparative
anatomy of amphineurans. I. On a few Solenogastres from
Naples.] Reviews: Journal of the Royal Microscopical Society,
London for 1895: 45 (English); Zoologischer Jahresbericht
(Berlin) fiir 1894 (1895) (Mollusca): 7, 13 (German).

1895a. Zur Phylogenie der Gastropoden. Biologisches Centralblatt,
15(6): 220-236; Leipzig, 15 March 1895. [On the phylogeny of
gastropods. |

*1895b. Hemitrichia guimarasensis n. sp. Nachrichtsblatt der Deutschen
Malakozoologischen Gesellschaft, 27(7-8): 131-132; Frankfurt
am Main, August 1895.

1895c. Bericht tiber die wissenschaftlichen Leistungen im Gebiete
der Malakologie wahrend des Jahres 1890. I. Bericht
liber Allgemeines, Physiologie, Anatomie und Entwick-
lung. Archiv fiir Naturgeschichte, 57(2.1) (1891): 300-324;
Berlin, October 1895. [Report on the the scientific work in the
realm of malacology during the year 1890. I. Report on general
subjects, physiology, anatomy and embryology.]

1895d. Ueber die Verwandtschaftsbeziehungen der Amphineuren.
Biologisches Centralblatt, 15(24): 859-869; Leipzig, 15 De-
cember 1895. [On the phyletic relationships of amphineurans.]
Review: Zoologischer Jahresbericht (Berlin) ftir 1895 (1896)
(Mollusca): 5, 12 (German).

1897a. Beitrage zur Kenntnis der Mollusken. III. Uber Hautdriisen |
und ihre Derivate. Zeitschrift fiir wissenschaftliche Zoologie,
62(4): 632-670, pls. 31-32; Leipzig, 11 May 1897. [Contributions |
to the knowledge of the mollusks. III. On the skin glands and |
their derivatives.] Reviews: Journal of the Royal Microscopical |

December 28, 1989 Bieler & Boss ih

Society, London, 1897: 368 (English); Zoologischer Jahresbericht
(Berlin) fiir 1897 (1898) (Mollusca): 5, 10 (German).

*1897b. Zwei australische Solenogastres. Zoologischer Anzeiger, 20(542):
398-400; Leipzig, 11 October 1897. [Two Australian Sole-
nogastres.] Reviews: Journal of the Royal Microscopical Society,
London, 1897: 527 (English); Zoologischer Jahresbericht (Ber-
lin) fiir 1897 (1898) (Mollusca): 5, 16 (German).

*1900a. Proneomenia thulensis nov. spec. Fauna Arctica. Eine Zusam-
menstellung der arktischen Tierformen, mit besonderer
Berticksichtigung des Spitzbergen-Gebietes auf Grund der
Ergebnisse der Deutschen Expedition in das Noérdliche Eismeer
im Jahre 1898, 1(1): 109-116, pl. 5; Jena, January 1900. Review:
Zoologischer Jahresbericht (Berlin) fiir 1900 (1901) (Mollusca):
6, 13 (German).

*1900b. Verzeichnis der von Herrn Dr. A. Voeltzkow gesammelten
marinen und litoralen Mollusken. Abhandlungen
herausgegeben von der Senckenbergischen Naturforschenden
Gesellschaft, 26(2) [Wissenschaftliche Ergebnisse der Reisen
in Madagaskar und Ostafrika in den Jahren 1889-95 von Dr.
A. VoeltzKow, 2(2)]: 241-252, 6 text-figs.; Frankfurt am Main,
1900. [List of marine and littoral mollusks collected by Dr. A.
Voeltzkow. |

190la. Ueber die Ausbildung der Korperform der Gastropoden.
Archiv fiir Naturgeschichte, 67, Beiheft (Festschrift ftir Eduard
von Martens): 9-22, 4 textfigs; Berlin, 18 April 1901. [On the
formation of body shape of gastropods.] Reviews: Journal of
the Royal Microscopical Society, London, 1901(5): 520-521
(English); Zoologischer Jahresbericht (Berlin) fiir 1901 (1902)
(Mollusca): 6, 36 (German).

1901b. Ueber die phyletische Enstehung und die Formentwicklung
der Moliuskenschale. Biologisches Centralblatt, 21(9): 274-
277; Leipzig, 1 May 1901. [On the phyletic origin and the
development of form of the molluscan shell.] Review: Zoologi-
scher Jahresbericht (Berlin) fiir 1901 (1902): 6, 7 (German).

1901c. Bemerkungen zur Systematik der Landdeckelschnecken.
Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft,
33(5-6): 89-93; Frankfurt am Main, 11 June 1901. [Remarks
on the systematics of operculate land snails.]

12 NEMOURIA No. 34

1902a. Zur Colomfrage. Zoologischer Anzeiger, 25(661): 82-84; Leipzig,
13 January 1902. [On the problem of the coelom.] Review:
Zoologischer Jahresbericht (Berlin) fiir 1902 (1903) (Mollusca):
6, 14 (German).

*1902b. Die systematische Stellung der Solenogastren und die
Phylogenie der Mollusken. Zeitschrift fiir wissenschaftliche
Zoologie, 72(2-3): 249-466, pls. 18-27, 21 text-figs.; Leipzig,
16 September 1902. [The systematic position of the solenogas-
ters and the phylogeny of mollusks.] Review: Zoologischer
Jahresbericht (Berlin) fiir 1902 (1903) (Mollusca): 6, 8 (Ger-
man).

*1902c. Proneomenia valdiviae n. sp. Wissenschaftliche Ergebnisse der
deutschen Tiefsee-Expedition auf dem Dampfer “Valdivia’’
1898-1899, 3(5): 167-174, pl. 23; Jena (Gustav Fischer), 1902
(March or later, p. 174 states “im Marz’’). Review: Zoologischer
Jahresbericht (Berlin) fiir 1903 (1904) (Mollusca): 5, 12 (Ger-
man).

*1903. Proneomenia amboinensis n. sp. [In:] Semon, R.: Zoologische
Forschungsreisen in Australien und dem Malayischen Archipel,
5(VI). Denkschriften der medicinisch-naturwissenschaftlichen
Gesellschaft zu Jena, 8: 735-737, pl. 66, figs. 5-9; Jena, 1903.
Review: Zoologischer Jahresbericht (Berlin) fiir 1903 (1904)
(Mollusca): 5, 11 (German).

*1904a. [Artbeschreibungen in: Martens, E. von:] Systematisch-
geographischer Teil. [In:] Martens, E. von & J. Thiele: Die
beschalten Gastropoden der deutschen Tiefsee-Expedition 1898-
1899. Wissenschaftliche Ergebnisse der deutschen Tiefsee-
Expedition auf dem Dampfer ‘Valdivia’ 1898-1899, 7(A):
1-146, pls. 1-5, 1 text-fig.; Jena, 1903 [title page states “1903”’’,
but ‘According to J. Thiele (letter, April 14, 1905), this work
was published during the second half of January, 1904; he
received his copy on January 30, 1904” (Weaver & DuPont,
1970: 353)]. [(Descriptions of species in: Martens, E. von:)
Systematic and geographic part . . . The shelled gastropods of
the German Deep-Sea Expedition of 1898-1899.] Review:
Zoologischer Jahresbericht (Berlin) fiir 1903 (1904) (Mollusca):
5, 24 (German).

December 28, 1989 Bieler & Boss 13

1904b. Anatomisch-systematische Untersuchungen einiger Gastro-
poden. [In:] Martens, E. von & J. Thiele: Die beschalten
Gastropoden der deutschen Tiefsee-Expedition 1898-1899. Wissen-
schaftliche Ergebnisse der deutschen Tiefsee-Expedition auf
dem Dampfer ‘Valdivia’? 1898-1899, 7(B): 147-179 [1-33], pls.
6-9 [1-4]; Jena, 1904 (title page states “‘1903’’; see above).
[Anatomical and systematic investigations of a few gastropods.
The shelled gastropods of the German Deep-Sea Expedition
of 1898-1899.]

*1906a. Bemerkung tiber die Gattung Photinula. Nachrichtsblatt der
Deutschen Malakozoologischen Gesellschaft, 38(1): 12-15; Frank-
furt am Main, 25 January 1906. [Remark on genus Photinula.]

*1906b. Zwei neue Macrodontes-Arten. Nachrichtsblatt der Deutschen
Malakozoologischen Gesellschaft, 38(2): 69-71, 2 text-figs.;
Frankfurt am Main, 2 April 1906. [Two new species of
Macrodontes. |

*1906c. Archaeomenia prisca n. g., n. sp. Wissenschaftliche Ergebnisse
der deutschen Tiefsee-Expedition auf dem Dampfer “‘Valdivia”’
1898-1899, 9(2): 315-324 [1-10], pl. 28; Jena (after 14 March
1906, date of manuscript submission). Review: Zoologischer
Jahresbericht (Berlin) fiir 1906 (1907) (Mollusca): 6, 7 (Ger-
man).

*1906d. Ueber die Chitonen der deutschen Tiefsee-Expedition.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition
auf dem Dampfer “Valdivia” 1898-1899, 9(2): 325-336 [11-22],
pl. 29; Jena (after 31 May 1906, date of manuscript submission).
[On the chitons of the German Deep-Sea Expedition.] Review:
Zoologischer Jahresbericht (Berlin) fiir 1906 (1907) (Mollusca):
6, 8 (German).

*1906e. Amphineures. 3 pp. [In:] Expédition Antarctique Frangaise
(1903-1905) commandée par le Dr. Jean Charcot. Sciences
naturelles: documents scientifiques. Paris (Masson & Cie),
December 1906. [Text identical with 1906f]

1906f. Note sur les Chitons de l’Expédition Antarctique du Dr.
Charcot. Bulletin Muséum d’Histoire naturelle Paris, 12 (1906
no. 7): 549-550, 1906. [Note on the chitons of the Antarctic
Expedition of Dr. Charcot.] [Text identical with 1906e]

14 NEMOURIA No. 34

1907a. Polyplacophora und Solenogastres fiir 1894-1905. Archiv fiir
Naturgeschichte, 68(2.3)(1902): 1-16; Berlin, September 1907.
[(Literature on) Polyplacophora and Solenogastres from 1894
to 1905.]

1907b. Sind die Chaetognathen als Mollusken aufzufassen? Zoologi-
scher Anzeiger, 32(14): 428-430; Leipzig, 10 December 1907.
[Are chaetognaths to be considered as mollusks?] Review:
Zoologischer Jahresbericht (Berlin) ftir 1907 (Vermes): 15, 54
(German).

*1908a. [Herausgabe von und Artbeschreibungen in:] Martens, E. von:
Beschreibung einiger im dstlichen Borneo von Dr. Mar-
tin Schmidt gesammelten Land- and Siifs$wasser-
Conchylien. Mitteilungen aus dem Zoologischen Museum in
Berlin, 4(1): 249-291, pls. 5-6, July 1908. [(Editing of and
species descriptions in:) Martens, E. von: Description of a few
land and freshwater shells collected in eastern Borneo by Dr.
Martin Schmidt.]

1908b. Ueber die Anatomie und systematische Stellung von Bathy-
sciadium, Lepetella, and Addisonia. Reports on the scien-
tific results of the Expedition to the Eastern Tropical Pacific,
in charge of Alexander Agassiz, by the U.S. Commission
Steamer ‘‘Albatross,” from October, 1904, to March, 1905,
Lieut. Commander L. M. Garrett, U.S.N., commanding, XV.
Bulletin of the Museum of Comparative Zoology at Harvard
College, 52(5): 81-89, pls. 1-2; Cambridge, October 1908. [On
the anatomy and systematic position of Bathysciadium, Lepetella,
and Addisonia.| Review: Zoologischer Jahresbericht (Berlin) ftir
1908 (1909) (Mollusca): 6, 12 (German).

*1908c. Die antarktischen und subantarktischen Chitonen. Deutsche
Stidpolar-Expedition 1901-1903, 10 (Zool., 2)(1): 7-23, pl. 1;
Berlin (Georg Reimer), October 1908. [The antarctic and
subantarctic chitons.]

1909a. Einige Bemerkungen iiber deutsche Siisswassermollusken
und ihre Namen. Nachrichtsblatt der Deutschen Malakozoologi-
schen Gesellschaft, 41(1): 25-34, pl. 1; Frankfurt am Main,
20 January 1909. [A few remarks on German freshwater
mollusks and their names.]

1909b. Ueber einige neuseelandische Chitonen. Nachrichtsblatt der
Deutschen Malakozoologischen Gesellschaft, 41(3): 131-132;

December 28, 1989 Bieler & Boss 15

Frankfurt am Main, 20 July 1909. [On a few chitons from New
Zealand. |

1909c. Zum Namen Unio. [In:] Kobelt, W. (ed.): Beitrége zur Kenntniss
der mitteleuropdischen Najadeen, 3. Beilage zum Nachrichtsblatt
der Deutschen Malakozoologischen Gesellschaft, 41: 48; Frank-
furt am Main, July 1909. [On the (generic) name Unio...
Contributions to the knowledge of Central European naiades.|

*1909d. Revision des Systems der Chitonen. I. Teil. Zoologica.
Original-Abhandlungen aus dem Gesamtgebiete der Zoologie,
22(56/1): 1-70, pls. 1-6, 5 text-figs.; Stuttgart, 1909 (received
by the Academy of Natural Sciences of Philadelphia: 19 July
1909). [Revision of the systematics of chitons. Part 1.]|

1909e. Solenogastres fiir 1906. Archiv fiir Naturgeschichte, 73(2.3)
(1907): 1 p.; Berlin, September 1909. [(Literature on) Solenogas-
tres in 1906.]

1909f. Polyplacophora fiir 1906. Archiv ftir Naturgeschichte, 73(2.3)
(1907): 2 pp.; Berlin, September 1909. [(Literature on) Polyplaco-
phora in 1906.]

*1909g. Cocculinoidea und die Gattungen Phenacolepas und Titi-
scania. Systematisches Conchylien-Cabinet von Martini und
Chemnitz (Ktister, H.C. & W. Kobelt, eds.), 2/11a(539): 1-48,
pls. 1-6; Ntirnberg (Bauer & Raspe), 1909 (dates imprinted on
individual sheets: 10 September-13 October). [The Cocculinoi-
dea and the genera Phenacolepas and Titiscania.]

1909h. Uber einige “Realiiden”. Archiv fiir Naturgeschichte, 75(1.3):
387-390, 4 text-figs.; Berlin, December 1909. [On several
“‘realiids’’. ]

*1909i1. Revision des Systems der Chitonen. II. Teil. Zoologica.
Original-Abhandlungen aus dem Gesamtgebiete der Zoologie,
22(56/2): 71-132, pls. 7-10; Stuttgart, 1909 (‘‘1910” on title
page; listings on subsequent journal covers show it as of 1909).
[Revision of the systematics of chitons. Part 2.]

1909}. Mollusca, Weichtiere. [In:] Brauer, A. (ed.): Die Siisswasserfauna
Deutschlands. Eine Exkursionsfauna, 19. Pp. 1-46 (+ register:
195-199), 112 text-figs.; Jena (Gustav Fischer), 1909; reprinted
by Verlag von J. Cramer, Weinheim, 1961. [Mollusca, . . . The
freshwater fauna of Germany. A field guide.]

16 NEMOURIA No. 34

1910a. Polyplacophora fiir 1907. Archiv fiir Naturgeschichte, 74(2.3)
(1908): 1 p.; Berlin, January 1910. [(Literature on) Polyplaco-
phora in 1907.]

1910b. Uber die Auffassung der Leibeshéhle von Mollusken und
Anneliden. Zoologischer Anzeiger, 35(22): 682-695, 2 text-figs.;
Leipzig, 24 May 1910. [On the concept of the body cavities of
mollusks and annelids.] Review: Zoologischer Jahresbericht
(Berlin) fiir 1910 (1911) (Mollusca): 6, 7 (German).

*1910c. Eine arabische Ennea und Bemerkungen itiber andere
Arten. Sitzungsberichte der Gesellschaft Naturforschender
Freunde zu Berlin, 6: 280-284, pl. 10; Berlin, June 1910. [An
Arabic Ennea and remarks on other species. ]

1910d. Solenogastres fiir 1908. Archiv ftir Naturgeschichte, 75(2.3)
(1909): 1 p.; Berlin, September 1910. [(Literature on) Solenogas-
tres in 1908.]

1910e. Polyplacophora ftir 1908. Archiv fiir Naturgeschichte, 75(2.3)
(1909): 1 p.; Berlin, September 1910. [(Literature on) Polyplaco-
phora in 1908.]

*1910f. Molluskenfauna Westindiens. Zoologische Jahrbiicher, Supple-
ment 11: 109-132, pl. 9; Jena, 21 November 1910. [Molluscan
fauna of the West Indies.]

*1910g. Polyplacophora, Gastropoda marina, Bivalvia. [In:] Schultze,
L.: Zoologische und anthropologische Ergebnisse einer
Forschungsreise im westlichen und zentralen Stidafrika ausgefiihrt
in den Jahren 1903-1905 mit Unterstiitzung der Kgl. Preussi-

schen Akademie der Wissenschaften zu Berlin, 4 (Systematik ©
und Tiergeographie) (1). Denkschriften der medicinisch- |
naturwissenschaftlichen Gesellschaft zu Jena, 16: 269-270, 1 |

text-fig.; Jena (Gustav Fischer), 1910.

1910h. Uber die Anatomie von Hydrocena cattaroensis Pf. Abhand-
lungen der Senckenbergischen Naturforschenden Gesellschaft, |
32 (Festschrift Kobelt): 349-358, pl. 25, 2 text-figs.; Frankfurt |
am Main, 1910. [On the anatomy of Hydrocena cattaroensis Pf.] |
Review: Zoologischer Jahresbericht (Berlin) fir 1910 (1911) |

(Mollusca): 6, 14 (German).

191la. [Heligerone Strand]. Nachrichtsblatt der Deutschen Malakozoolo- |
gischen Gesellschaft, 43(2): 104; Frankfurt am Main, 4 May |

1911.

\

!
i
,

|
|

F

;

December 28, 1989 Bieler & Boss 17

191lb. Die Solenogastres der Russischen Polar-Expedition 1900-
1903. Mémoires de l’Académie Impériale des Sciences de
St.-Pétersbourg, (8)29(5); Résultats scientifiques de l’Expédition
Polaire Russe en 1900-1903 sous la direction du Baron E. Toll,
Sect. E, Zool., 2(5): 1-4, 1 pl.; St. Petersburg, July 1911. [The
Solenogastres of the Russian Polar Expedition of 1900-1903.]

191lc. Polyplacophora fiir 1909. Archiv ftir Naturgeschichte, 76(6/1):
111-116; Berlin, November 1911. [(Literature on) Polyplaco-
phora in 1909.]

*1911d. Mollusken der Deutschen Zentralafrika-Expedition. Wissen-
schaftliche Ergebnisse der deutschen Zentral-Afrika-Expedition
1907-1908 unter Fiihrung Adolf Friedrichs, Herzog zu Meck-
lenburg, 3 (Zool.): 175-214, pls. 4-6; Leipzig (Klinkhardt &
Biermann), 1911. [Mollusks of the German Central African
Expedition. ]

*191le. Polyplacophora. [In:] Michaelsen, W. & R. Hartmeyer (eds.):
Die Fauna Stidwest-Australiens. Ergebnisse der Hamburger
stidwest-australischen Forschungsreise 1905, 3(2): 397-406, 1
pl.; Jena (Gustav Fischer), 1911. [Polyplacophora ... The
fauna of southwest Australia. ]

*1912a. Familia Scissurellidae. Systematisches Conchylien-Cabinet von
Martini und Chemnitz (Ktister, H.C. & W. Kobelt, eds.),
2/4a(558): 1-36, pls. 1-4; Ntirnberg (Bauer & Raspe), 1912
(dates imprinted on individual sheets: 5 April-23 May).

*1912b. Die antarktischen Schnecken und Muscheln. Deutsche Stidpolar-
Expedition 1901-1903, 13 (Zool., 5) (2): 183-285, pls. 11-19,
18 text-figs.; Berlin (Georg Reimer), August 1912. [The
antarctic snails and bivalves. ]

1912c. Amphineura. Handwéorterbuch der Naturwissenschaften, 1: 347-
354, 12 text-figs.; Jena (Gustav Fischer), 1912.

1912d. Lamellibranchiata. Handworterbuch der Naturwissenschaften, 6:
6-21, 19 text-figs.; Jena (Gustav Fischer), 1912.

1912e. Mollusca. Handworterbuch der Naturwissenschaften, 6: 1045-1049,
4 text-figs.; Jena (Gustav Fischer), 1912.

18 NEMOURIA No. 34

*1913a. [Literature review of 1912b:] Thiele, J., Die antarktischen
Schnecken und Muscheln. Zentralblatt fiir Zoologie, allge-
meine und experimentelle Biologie, 2(3): 86-87; Leipzig and
Berlin, 10 January 1913.

1913b. Dr. Naef und das “Cdlom” der Mollusken. Zoologischer
Anzeiger, 41(6): 251-252; Leipzig, 31 January 1913. [Dr. Naef
and the “coelom”’ of mollusks.]

*1913c. Ein neuer Solenogaster von Spitzbergen. Sitzungsberichte der
Gesellschaft Naturforschender Freunde zu Berlin, 1913(2):
160-162, 2 text-figs.; Berlin, February 1913. [A new solenogaster
from Spitzbergen.]

*1913d. Antarktische Solenogastren. Deutsche Siidpolar-Expedition
1901-1903, 14 (Zool., 6)(1): 35-65, pls. 4-8, 2 text-figs.; Berlin
(Georg Reimer), May 1913. [Antarctic solenogasters. ]

*1913e. Mollusca. Solenogastres. [In:] Schulze, F.E. (ed.): Das Tier-
reich. Eine Zusammenstellung und Kennzeichnung der rezenten
Tierformen, 38: x + 57 pp., 28 text-figs.; Berlin (R. Friedlander
& Sohn), June 1913; reprinted by Verlag von J. Cramer,
Weinheim, 1966.

*1913f. Familia Fissurellidae. Systematisches Conchylien-Cabinet von
Martini und Chemnitz (Ktister, H.C. & W. Kobelt, eds.),
2/4a(568): 37-68, pls. 5-8; Ntirnberg (Bauer & Raspe), 1913
(dates imprinted on individual sheets: 11-30 September). [con-
tinued in 1915b, 1916b and 1919]

1913g. Scaphopoda. Handwé6rterbuch der Naturwissenschaften, 8: 832-
835, 3 text-figs.; Jena (Gustav Fischer), 1913.

1914. Siidafrikanische Schnecken. Deutsche Stidpolar-Expedition 1901-
1903, 16 (Zool., 8)(1): 97-100, 2 text-figs.; Berlin (Georg
Reimer), November 1914. [South African snails.]

*1915a. Eulima capensis n. sp. [In:] Clark, A. H.: Die Crinoiden der
Antarktis. Deutsche Stidpolar-Expedition 1901-1903, 16 (Zool.,
8)(2): 165, 1 text-fig.; Berlin (Georg Reimer), April 1915.

*1915b. Familia Fissurellidae. Systematisches Conchylien-Cabinet von
Martini und Chemnitz (Ktister, H.C. & W. Kobelt, eds.),
2/4a(577): 69-104, pls. 9-12; Niirnberg (Bauer & Raspe), 1915
(dates imprinted on individual sheets: 3-19 May). [continued
from 1913f; continued in 1916b and 1919]

December 28, 1989 Bieler & Boss 19

1915c. Molluscoidea. Mollusca. [In:] Apstein, C.: Nomina conservanda.
Unter Mitwirkung zahlreicher Spezialisten. Sitzungsberichte
der Gesellschaft Naturforschender Freunde zu Berlin, 1915(5):
180-184; Berlin, May 1915.

*1916a. Bemerkungen iiber die Systematik der achtarmigen Cepha-
lopoden. Zoologischer Anzeiger, 48(1): 3-4; Leipzig, 19 Septem-
ber 1916. [Remarks on the systematics of the eight-armed
cephalopods. |

*1916b. Familia Fissurellidae. Systematisches Conchylien-Cabinet von
Martini und Chemnitz (Ktister, H.C. & W. Kobelt, eds.),
2/4a(578): 105-144, pls. 13-16; Ntirnberg (Bauer & Raspe),
1916 (“1917” on title page; dates imprinted on individual sheets:
28 October - 30 November 1916). [continued from 1913f and
1915b; continued in 1919]

*1917a. Bemerkungen iiber das “Tierreich” und den Nomenclator
Generum Animalium. Nachrichtsblatt der Deutschen Malako-
zoologischen Gesellschaft, 49(1): 19-24; Frankfurt am Main,
2 January 1917. [Remarks on the “Animal Kingdom” and the
Nomenclator Generum Animalium.]

*1917b. Marine und litorale Mollusken von Madagascar, den
Comoren und Ostafrika (Sammlung Voeltzkow 1903-05).
[In:] Voeltzkow, A.: Reise in Ostafrika in den Jahren 1903-1905.
Wissenschaftliche Ergebnisse. Band III. Systematische Arbeiten,
5: 561-564; Stuttgart (E. Schweizerbart’sche Verlagsbuchhand-
lung), 1917. [Marine and littoral mollusks of Madagascar, the
Comores and East Africa (collection Voeltzkow 1903-05).]

*1918. Familia Limidae. Systematisches Conchylien-Cabinet von Martini
und Chemnitz (Ktister, H.C., W. Kobelt & F. Haas, eds.),
7/2a(579): 1-24, pls. 1-5; Niirnberg (Bauer & Raspe), 1918
(dates imprinted on individual sheets: 10-19 July). [continued
in 1920a]

*1919. Familia Fissurellidae. Systematisches Conchylien-Cabinet von
Martini und Chemnitz (Kiister, H. C., W. Kobelt & F. Haas,
eds.), 2/4a(580): 145-168 [ends in the middle of a sentence],
pls. 17-20; Ntirnberg (Bauer & Raspe), 1919 (dates imprinted
on individual sheets: 18-24 May). [continued from 1913f, 1915b,
and 1916b]

20 NEMOURIA No. 34

*1920a. Familia Limidae. Systematisches Conchylien-Cabinet von Mar-
tini und Chemnitz (Kiister, H.C., W. Kobelt & F. Haas, eds.),
7/2a(583): 25-66, pls. 6-10; Nurnberg (Bauer & Raspe), 1920
(dates imprinted on individual sheets: 1 September 1919-19
June 1920). [continued from 1918]

*1920b. Die Cephalopoden der deutschen Siidpolar-Expedition
1901-1903. Deutsche Stidpolar-Expedition 1901-1903, 16 (Zool.,
8)(4): 433-465, pls. 52-55; Berlin and Leipzig (Vereinigung
wissenschaftlicher Verleger), December 1920. [The cephalopods
of the German South Polar Expedition of 1901-1903.]

1921a. Ueber die Radula und systematische Stellung von Nautilinus
clymene (Shuttl.). Archiv fiir Molluskenkunde, 53(1/2): 111-
112; Frankfurt am Main, 12 April 1921. [On the radula and
the systematic position of Nautilinus clymene (Shuttl.).]

*1921b. Zur Systematik der Mollusken. Archiv ftir Molluskenkunde,
53(3): 140-163, pl. 4; Frankfurt am Main, 12 July 1921. [On
the systematics of mollusks. ]

1922. Uber die Verwandtschaftsbeziehungen einiger ungegliederter
Tiergruppen. Zoologischer Anzeiger, 54(5/6): 140-144; Leipzig,
14 February 1922. [On the phyletic interrelationships of several
unsegmented animal groups.]

*1923. Uber die Gattung Philobrya und das sogenannte Buccalner-
vensystem von Muscheln. Zoologischer Anzeiger, 55(11/13):
287-292; Leipzig, 16 January 1923. [On the genus Philobrya
and the so-called buccal nervous system of bivalves.]

*1924a. Revision des Systems der Trochacea. Mitteilungen aus dem
Zoologischen Museum in Berlin, 11(1): 49-72, 1 pl.; Berlin,
February 1924 (frequently cited as of 1921, even by Thiele;
manuscript was submitted in 1921). [Revision of the systematics
of the Trochacea.]

*1924b. Ueber die Systematik der Columbelliden. Archiv fiir Mollus-
kenkunde, 56(5): 200-210, pl. 9; Frankfurt am Main, 15
October 1924. [On the systematics of columbellids.]

1925a. Ueber Diala leithit Edg. Smith. Archiv ftir Molluskenkunde,
57(3): 111-112; Frankfurt am Main, 7 May 1925. [On Duala
leithit Edg. Smith.]

December 28, 1989 Bieler & Boss Zi

*1925b. Gastropoda der Deutschen Tiefsee-Expedition. II. Teil.

1925c.

Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition
auf dem Dampfer “Valdivia”? 1898-1899, 17(2): 35-382 [1-348],
pls. 13-46 [1-34], 31 text-figs.; Jena (Gustav Fischer), 1925
(introduction signed “‘Februar 1925”; received by Zoologisches
Museum in Berlin: 10 November). [Gastropoda of the German
Deep-Sea Expedition. Part 2.]

Solenogastres. [In:] Ktikenthal, W. & T. Krumbach (eds.):

Handbuch der Zoologie. Eine Naturgeschichte der Stémme des
Tierreiches, 5(1): 1-14, text-figs. 1-7; Berlin and Leipzig (Walter
de Gruyter & Co.), 1 November 1925.

1925d. Mollusca = Weichtiere. [In:] Kikenthal, W. & T. Krumbach

1926a.

1926b

(eds.): Handbuch der Zoologie. Eine Naturgeschichte der
Stéamme des Tierreiches, 5(1): 15-96, text-figs. 8-109; Berlin
and Leipzig (Walter de Gruyter & Co.), 1 November 1925.
[continued in 1926a,b]

Mollusca = Weichtiere. [In:] Kiikenthal, W. & T. Krumbach

(eds.): Handbuch der Zoologie. Eine Naturgeschichte der
Stémme des Tierreiches, 5(2): 97-176, text-figs. 110-269; Berlin
and Leipzig (Walter de Gruyter), 20 February 1926. [continued
from 1925d; continued in 1926b]

. Mollusca = Weichtiere. [In:] Kiikenthal, W. & T. Krumbach

(eds.): Handbuch der Zoologie. Eine Naturgeschichte der
Stamme des Tierreiches, 5(3): 177-256, text-figs. 270-340; Berlin
and Leipzig (Walter de Gruyter), 30 June 1926. [continued from
1925d and 1926a]

*1927a. Uber die Schneckenfamilie Assimineidae. Zoologische

Jahrbiicher, Abteilung fiir Systematik, Okologie und Geographie
der Tiere, 53(1/3): 113-146, pl. 1, 13 text-figs.; Jena, 17
February 1927. [On the gastropod family Assimineidae.]

1927b. Ueber die Gattung Ceratodiscus. Archiv fiir Molluskenkunde,

59(2): 155-157, pl. 9; Frankfurt am Main, 1 March 1927. [On
the genus Ceratodiscus.]

22 NEMOURIA No. 34

*1927c. Uber einige brasilianische Landschnecken. Abhandlungen
herausgegeben von der Senckenbergischen Naturforschenden
Gesellschaft, 40(3) (Ergebnisse einer zoologischen Forschungs-
reise in Brasilien 1913-1914 von Prof. Dr. E. Breslau, 1. Teil):
305-329, pl. 26, 7 text-figs.; Frankfurt am Main, 30 December
1927. [On a few Brazilian land snails. ]

*1928a. Mollusken vom Bismarck-Archipel, von Neu-Guinea und
Nachbar-Inseln. Zoologische Jahrbticher, Abteilung ftir Systema-
tik, Okologie und Geographie der Tiere, 55(1-3): 119-146, pl.
5, 1 text-fig.; Jena, 25 April 1928. [Mollusks from the Bismarck
Archipelago, New Guinea, and neighboring islands. |

*1928b. Arktische Loricaten, Gastropoden, Scaphopoden und Bi-
valven. Fauna Arctica. Eine Zusammenstellung der arktischen
Tierformen, mit besonderer Beriicksichtigung des Spitzbergen-
Gebietes auf Grund der Ergebnisse der Deutschen Expedition
in das N6rdliche Eismeer im Jahre 1898, 5(2): 561-632, pl. 10;
Jena (Gustav Fischer), 3 August 1928. [Arctic loricates, gastro-
pods, scaphopods and bivalves. ]

*1928c. Revision des Systems der Hydrobiiden und Melaniiden.
Zoologische Jahrbiicher, Abteilung fiir Systematik, Okologie
und Zoogeographie der Tiere, 55(5-6): 351-402, pl. 8, 63
text-figs.; Jena, 12 September 1928. [Revision of the systematics
of hydrobiids and melaniids].

1928d. Uber ptenoglosse Schnecken. Zeitschrift fiir wissenschaftliche
Zoologie, 132 (Korschelt-Festband): 73-94, 11 text-figs.; Leipzig,
September 1928. [On ptenoglossate snails].

*1929. Handbuch der systematischen Weichtierkunde. Vol. 1(1)
[Loricata; Gastropoda: Prosobranchia]: 1-376, 470 text-figs.;
Jena (Gustav Fischer) (‘‘1931” on title page of hard-cover ~
edition; published after 4 September 1929 teste Thiele, 1931c, —
p. 739; received by Zoologisches Museum in Berlin: 21 October
1929). Reprinted by A. Asher & Co., Amsterdam, 1963.
[Handbook of systematic malacology. English language transla- |
tion by Bieler, R. & P. M. Mikkelsen (eds.), Smithsonian
Institution Libraries & National Science Foundation, Amerind |
Publishing Co., New Delhi, in press.] [continued in 193lc, _
1934 and 1935]

December 28, 1989 Bieler & Boss 23

*1930. Gastropoda und Bivalvia. [In:] Michaelsen, W. & R. Hartmeyer
(eds.): Die Fauna Stidwest-Australiens. Ergebnisse der Ham-
burger stidwest-australischen Forschungsreise 1905, 5(8): 561-
596, pl. 4, 1 text-fig.; Jena (Gustav Fischer), 1930. [Gastropoda
and Bivalvia . . . The fauna of southwest Australia].

*193la. Ueber einige hauptsachlich afrikanische Landschnecken.
Sitzungsberichte der Gesellschaft Naturforschender Freunde
zu Berlin, 1930: 392-403, 1 pl., 7 text-figs.; Berlin, 1 March
1931. [On a few, principally African land snails.]

*1931b. Thiele, J. & S. Jaeckel: Muscheln der Deutschen Tiefsee-
Expedition. Wissenschaftliche Ergebnisse der deutschen Tiefsee-
Expedition auf dem Dampfer “Valdivia” 1898-1899, 21(1):
159-268 [1-110], pls. 6-10 [1-5]; Jena (Gustav Fischer), 1931
(received by Zoologisches Museum in Berlin: 14 July 1931).
[Bivalves of the German Deep-Sea Expedition. ]

*193lc. Handbuch der systematischen Weichtierkunde. Vol. 1(2)
[Gastropoda: Opisthobranchia and Pulmonata; additions; index
for parts 1 and 2]: 377-778, 313 text-figs.; Jena (Gustav Fischer),
1931 (received by Zoologisches Museum in Berlin: 31 October
1931). Reprinted by A. Asher & Co, Amsterdam, 1963.
[Handbook of systematic malacology.] [continued from 1929;
continued in 1934 and 1935]

1931d. Bivalvia. Handworterbuch der Naturwissenschaften. Zweite Auflage
[2nd ed.], 1: 996-1010, 18 text-figs.; Jena (Gustav Fischer),
1931 (received by Zoologisches Museum in Berlin: 15 January
1932).

*1932a. Das Gonaxis-Problem. Archiv fiir Molluskenkunde, 64(1):
11-12; Frankfurt am Main, 1 January 1932. [The Gonaxis
problem. |

1932b. Loricata. Handwéorterbuch der Naturwissenschaften. Zweite Auflage
[2nd ed.], 6: 550-552, 3 text-figs.; Jena (Gustav Fischer), 1932
(received by Zoologisches Museum in Berlin: 15 January 1932).

1932c. Die Solenogastres des arktischen Gebietes. Fauna Arctica.
Eine Zusammenstellung der arktischen Tierformen, mit beson-
derer Berticksichtigung des Spitzbergen-Gebietes auf Grund
der Deutschen Expedition in das No6rdliche Eismeer im Jahre
1898, 6(4): 379-382; Jena (Gustav Fischer), 18 October 1932.
[The Solenogastres of the arctic region. ]

24 NEMOURIA No. 34

1932d. Mollusca. Handworterbuch der Naturwissenschaften, Zweite Auflage
[2nd ed.], 6: 1079-1083, 2 textfigs; Jena (Gustav Fischer), 1932.

1933a. Scaphopoda. Handwéorterbuch der Naturwissenschaften. Zweite
Auflage [2nd ed.], 8: 997-999, 2 text-figs.; Jena (Gustav
Fischer), 1933 (before November, listed as “ausgegeben bis
Oktober”’ in Vol. 9).

1933b. Solenogastres. Handwérterbuch der Naturwissenschaften. Zweite
Auflage [2nd ed.], 9: 144-146, 2 text-figs.; Jena (Gustav
Fischer), 1933 (received by Zoologisches Museum in Berlin:
15 November 1933).

*1933c. Die von Oskar Neumann in Abessinien gesammelten und
einige andere afrikanische Landschnecken. Sitzungsberichte
der Gesellschaft Naturforschender Freunde zu Berlin, 1933(4-7):
280-323, pls. 1-2, 13 text-figs.; Berlin, 30 December 1933. [On
a few African land snails, including those collected by Oskar
Neumann in Abyssinia. ]

*1934. Handbuch der systematischen Weichtierkunde. Vol. 2(3)
[Scaphopoda; Bivalvia; Cephalopoda; additions and corrections
for parts 1 and 2; index for part 3]: 779-1022, 110 text-figs.;
Jena (Gustav Fischer), 1934 (“‘1935”’ on title page of hard-cover
edition; received by Zoologisches Museum in Berlin: 19 January
1934). Reprinted by A. Asher & Co., Amsterdam, 1963.
[Handbook of systematic malacology.] [continued from 1929
and 1931c; continued in 1935]

*1935. Handbuch der systematischen Weichtierkunde. Vol. 2(4)
[general part; corrections for parts 1-3]: i-iv, 1023-1154, 4
text-figs., pp. i-vi for vol. 1 (1929-1931), unnumbered divider
pages for parts 1-4; Jena (Gustav Fischer), 1935 (received by
Zoologisches Museum in Berlin: 27 March 1935). Reprinted
by A. Asher & Co., Amsterdam, 1963. [Handbook of systematic
malacology. Partial English language translation of pp. 1099-
1110 by Jacobson, 1953.] [continued from 1929, 193lc, and
1934]

December 28, 1989 Bieler & Boss 25
B. Publications on Crustacea

*1900a. Diagnosen neuer Arguliden-Arten. Zoologischer Anzeiger,
23(606): 46-48; Leipzig, 22 January 1900. [Diagnoses of new
argulid species.] Review: Journal of the Royal Microscopical
Society, London for 1900: 201 (English).

*1900b. Ueber einige Phyllopoden aus Deutsch Ost-Afrika. Zoologi-
sche Jahrbiicher, Abtheilung fiir Systematik, Geographie und
Biologie der Thiere, 13(6): 563-578, pls. 37-38; Jena, 10 October
1900. [On a few phyllopods from German East-Africa.]

*1904a. Ueber eine von Herrn O. Neumann gefundene Phyllopoden-
Art. Zoologische Jahrbticher, Abtheilung ftir Systematik, Geogra-
phie und Biologie der Thiere, 20(4): 371-374, pl. 13; Jena, 23
June 1904. [On a phyllopod species found by Mr. O. Neumann.]

*1904b. Beitrage zur Morphologie der Arguliden. Mitteilungen aus
dem Zoologischen Museum in Berlin, 2(4): 1-51, pls. 6-9;
Berlin, 22 December 1904. [Contributions to the morphology
of argulids.]

*1904c. Die Leptostraken. Wissenschaftliche Ergebnisse der deutschen
Tiefsee-Expedition auf dem Dampfer ‘Valdivia’? 1898-1899,
8(1): 1-26, pls. 1-4; Jena (Gustav Fischer), 1904. [The leptostra-
cans.] Review: Zoologischer Jahresbericht (Berlin) ftir 1905
(Arthropoda): 19, 35 (German).

*1905a. Uber die Leptostraken der Deutschen Siidpolar-Expedition
1901-1903. Deutsche Siidpolar-Expedition 1901-1903, 9 (Zool.
1)(1): 59-68, pl. 2, 1 map; Berlin (Georg Reimer), June 1905.
[On the leptostracans of the German South Polar Expedition
of 1901-1903.]

1905b. Betrachtungen iiber die Phylogenie der Crustaceenbeine.
Zeitschrift fiir wissenschaftliche Zoologie, 82: 445-471, pls.
26-27, 1 text-fig.; Leipzig, 10 November 1905. [Considerations
on the phylogeny of the legs of crustaceans.] Review: Zoologi-
scher Jahresbericht (Berlin) fiir 1905 (1906) (Arthropoda): 19,
24 (German).

26 NEMOURIA No. 34

1905c. Uber einige stielaugige Krebse von Messina. Zoologische
Jahrbticher, Supplement 8 (Festschrift M6bius): 443-474, pls.
14-16; Jena, 1905. [On a few stalk-eyed crabs from Messina.]
Review: Zoologischer Jahresbericht (Berlin) fiir 1905 (Arthro-
poda): 19, 36, 38 (German).

*1907a. Einige neue Phyllopoden-Arten des Berliner Museums.
Sitzungsberichte der Gesellschaft Naturforschender Freunde
zu Berlin, 1907(9): 288-297, pls. 1-2; Berlin, November 1907.
[A few new phyllopod species in the Berlin Museum. ]

1907b. Crustacea. IV. - Leptostraca. National Antarctic Expedition.
1901-1904. Natural History. Vol. II, Zoology and Botany
(Invertebrata: Marine Algae, Musci). 2 pp., 2 figs.; London
(British Museum [Natural History]), 1907.

1926. Leptostraca. [In:] Grimpe, G. & E. Wagler. Die Tierwelt der
Nord- und Ostsee. Lieferung VII, Teil X-g,, pp. 1-8, figs.
1-6; Leipzig (Akademische Verlagsgesellschaft), 1926 (“1927”
on title page, p. 8 stating: “X.g,;: Oktober 1926’’).

1927a. 1. und einzige Ordnung der ‘“Reihe Phyllocarida” der
Crustacea Malacostraca. 6. Ordnung der Crustacea:
Leptostraca. [In:] Ktikenthal, W. & T. Krumbach. Handbuch
der Zoologie. 3(1:5): 567-592, figs. 537-591; Berlin and Leipzig
(Walter de Gruyter), 10 June 1927. [First and only order of
the “series Phyllocarida’”’ of the Crustacea Malacostraca. 6th
order of the Crustacea: Leptostraca.]

C. Publications on Porifera

*1898. Studien iiber pazifische Spongien. Zoologica. Original-
Abhandlungen aus dem Gesammtgebiete der Zoologie, 24(1): —
1-72, pls. 1-8, 1 text-fig.; Stuttgart, 1898 (received by Museum
of Comparative Zoology, Cambridge: 27 June). [Studies on |
Pacific sponges.] [continued in 1899b]

1899a. Ueber Crambe crambe (O. Schmidt). Archiv fiir Naturges-
chichte, 65(1.1): 87-94, pl. 7; Berlin, April 1899. [On Crambe —
crambe (O. Schmidt).] Reviews: Journal of the Royal Micro-
scopical Society, London, for 1899: 399 (English); Zoologischer
Jahresbericht (Berlin) fiir 1899 (Porifera): 2, 7 (German).

December 28, 1989 Bieler & Boss Pa

*1899b. Studien iiber pazifische Spongien. Zoologica. Original-
Abhandlungen aus dem Gesammtgebiete der Zoologie, 24(2):
1-33, pls. 1-5; Stuttgart, 1899. [Studies on Pacific Ocean
sponges.] Review: Zoologischer Jahresbericht (Berlin) flr 1899
(1900) (Porifera): 2, 7 (German). [continued from 1898]

*1900. Kieselschwi4mme von Ternate. I. [In:] Ktikenthal, W.: Er-
gebnisse einer zoologischen Forschungsreise in den Molukken
und Borneo. Zweiter Teil: Wissenschaftliche Reiseergebnisse.
Band III. Abhandlungen herausgegeben von der Senckenber-
gischen Naturforschenden Gesellschaft, 25(1): 17-80, pls. 2-3,
1 text-fig.; Frankfurt am Main, 1900. [Siliceous sponges of
Ternate. Part 1.] Review: Zoologischer Jahresbericht (Berlin)
fiir 1900 (1901) (Porifera): 2, 6 (German).

*1903a. Beschreibung einiger unzureichend bekannten monaxonen
Spongien. Archiv fiir Naturgeschichte, 69(1.3): 375-398, pl.
21; Berlin, August 1903. [Description of a few insufficiently
known monaxonal sponges.] Review: Zoologischer Jahresbericht
(Berlin) fiir 1903 (1904) (Porifera): 1, 5 (German).

*1903b. Kieselschwamme von Ternate. II. [In:] Ktikenthal, W.:
Ergebnisse einer zoologischen Forschungsreise in den Molukken
und Borneo. Abhandlungen herausgegeben von der Senckenbergi-
schen Naturforschenden Gesellschaft, 25(4): 931-968, pl. 28;
Frankfurt am Main, 1903. [Siliceous sponges of Ternate. Part
2.]. Review: Zoologischer Jahresbericht (Berlin) fiir 1903 (1904)
(Porifera): 1, 5 (German).

*1905. Die Kiesel- und Hornschwaéamme der Sammlung Plate.
Zoologische Jahrbticher, Supplement 6, Vol. 3(3) (Fauna Chilen-
sis. Abhandlungen zur Kenntniss der Zoologie Chiles nach den
Sammlungen von Dr. L. Plate): 407-496, pls. 27-33; Jena, 16
June 1905. [The silicious and corneous sponges of the Plate
collection.] Review: Zoologischer Jahresbericht (Berlin) fiir
1905 (1906) (Porifera): 1, 6 (German).

1921. Uber das System der Demospongien. Zoologischer Anzeiger,
53(1/2): 28-30; Leipzig, 3 June 1921. [On the systematics of
demospongeans. |

28 NEMOURIA No. 34
D. Miscellaneous zoological publications

1887. Der Haftapparat der Batrachierlarven. Zeitschrift fiir wissen-
schaftliche Zoologie, 46(1): 67-79, pl. 10, 1 text-fig.; Leipzig,
25 November 1887. [The adhesive apparatus of batrachian
larvae.] Review: Zoologischer Jahresbericht (Berlin) fiir 1887
(1888) (Vertebrata): 31, 87 (French).

1893. Die primitivsten Metazoen. Sitzungsberichte und Abhandlungen
der Naturwissenschaftlichen Gesellschaft Isis in Dresden, 1892
(Jul.-Dec.): 54-57; Dresden, 1893 (received by the Museum of
Comparative Zoology, Cambridge: 8 June 1893). [The most
primitive metazoans. |

1905. [Diskussion in:] Ziegler, H.E.: Das Ectoderm der Plathelminthes.
Verhandlungen der Deutschen Zoologischen Gesellschaft auf
der ftinfzehnten Jahreshauptversammlung zu Breslau, den 13.
bis 16. Juni 1905, 15: 35-41, 4 text-figs.; Leipzig, 1905
[Discussion by Korschelt, Schulze, Spengel, Woltereck and
Thiele, pp. 41-42. The ectoderm of Platyhelminthes. ]

1909. Brachiopoda fiir 1906. Archiv fiir Naturgeschichte, 73(2.3) (1907):
2 pp.; Berlin, September 1909. [(Literature on) Brachiopoda in
1906.]

1910a. Brachiopoda fiir 1907. Archiv fiir Naturgeschichte, 74(2.3) (1908):
1 p.; Berlin, January 1910. [(Literature on) Brachiopoda in
1907.]

1910b. Brachiopoden fiir 1908. Archiv fiir Naturgeschichte, 75(2.3)
(1909): 1 p.; Berlin, September 1910. [(Literature on) brachi-
opods in 1908.]

December 28, 1989 Bieler & Boss 29

SECONDARY BIBLIOGRAPHY

Biirk, R. & J. H. Jungbluth. 1985. 140 Jahre Molluskenkunde im deutschsprachigen
Raum, 1844-1984. Ungarisches Naturwissenschaftliches Museum, Budapest,
348 pp.

Buschmeyer, H. 1938. 70 Jahre Molluskenkunde (1869-1938). Senckenbergbuch
13. Senckenbergische Naturforschende Gesellschaft, Frankfurt am Main, 108
Pp.

Clench, W. J. 1952. Dates of publication of Johannes Thiele: Handbuch der
Systematischen Weichtierkunde, Jena, Germany. The Nautilus, 66(1): 33.

[ICZN] International Commission on Zoological Nomenclature. 1985. International
code of zoological nomenclature. Third edition. International Trust for
Zoological Nomenclature, etc., London, xx + 338 pp.

Jacobson, M. K. 1953. The phyletic development of the Pulmonata after Johannes
Thiele. Translated from the German. Circulares del Museo y Biblioteca de
Malacologia de la Habana, Dic. 1953: 973-984.

Johnson, R. I. 1968. Martini and Chemnitz (Kuester’s edition) Systematisches
Conchylien-Cabinet, 1837-1920, a complete collation. Journal of the Society
for the Bibliography of Natural History, 4(7): 363-367.

Kiss, E. 1985. Malakologiai Evfordulok. Malakologische Jubiléen, 1985. Soosiana,
Budapest, 13: 35-38.

Lamy, E. 1938. Nécrologie. J. Thiele (1860-1935). Journal de Conchyliologie, 82(3):
259-260.

Rensch, B. 1930. Johannes Thiele zum 70. Geburtstage. Archiv fiir Mollus-
Kenkunde, 62(6) [Festnummer zum 70. Geburtstag von Johannes Thiele]:
201-209, pl. 11.

Robertson, R. 1957. Publication dates of Troschel’s ‘‘Das Gebiss der Schnecken’’.
The Nautilus, 70(4): 136-138.

Scheele, M. & G. Natalis. 1981. Biologie-Dokumentation. Bibliographie der
deutschen biologischen Zeitschriftenliteratur 1796-1965. Vol. 17. K. G. Saur,
Miinchen, New York, London, Paris.

Smith, E. A. & H. W. England. 1937. Martini und Chemnitz (Kuester’s edition)
Systematisches Conchylien-Cabinet, 1837-1918. Journal of the Society for
the Bibliography of Natural History, 1(4): 89-99.

Vosmaer, G. C. J. (Bidder, G. P. & C. S. Vosmaer-RGell, eds.). 1928. Bibliography
of Sponges, 1551-1913. University Press, Cambridge, xii + 234 pp.

Weaver, C. S. & J. E. duPont. 1970. The living volutes. A monograph of the
Recent Volutidae of the world. Delaware Museum of Natural History
Monograph Series No. 1, Greenville, xv + 375 pp. incl. 79 pls.

Winckworth, R. 1938. Johannes Thiele, 1860-1935. Proceedings of the Malacological
Society of London, 23(1): 9-11.

30 NEMOURIA No. 34
ACKNOWLEDGMENTS

We are grateful to the following for providing copies of Thiele’s
publications and/or information concerning his life and work: Mr. Bernd
von Breitenbuch (Gustav Fischer Verlag, Stuttgart), Dr. Ronald Janssen,
Mr. Alan Kabat, Dr. Rudolf Kilias, Ms. Paula M. Mikkelsen, Dr. Robert
Robertson, Dr. Gary Rosenberg, Dr. Klaus Ritzler. Mr. L. Laszlo Meszoly
kindly translated Kiss’ (1985) paper from the Hungarian for us.

Founding Editor Managing Editor
JOHN E. DU PONT RUDIGER BIELER

Consulting Editors

WILLIAM K. EMERSON, American Museum of Natural History, New York,
NY.

M. G. HARASEWYCH, National Museum of Natural History, Smithsonian
Institution, Washington, DC.

KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh,
PA.
RICHARD W. THORINGTON, JR., National Museum of Natural History,
Smithsonian Institution, Washington, DC. |

Published by the

DELAWARE MUSEUM OF NATURAL HISTORY
4840 Kennett Pike, Wilmington,
Delaware 19807, U.S.A.

NEMOURIA

Occasional Papers of the Delaware Museum of Natural History

NUMBER 35 April 30, 1990

MORPHOLOGY AND HOMOLOGOUS FEATURES
IN THE MALE PALPAL ORGAN
IN PISAURIDAE AND
OTHER SPIDER FAMILIES,
WITH NOTES ON THE TAXONOMY OF PISAURIDAE
(ARACHNIDA: ARANEAE)

Petra Sierwald!

“The question as to whether the genitalic characters on which this
subfamily [Otiothopinae; Araneae, Palpimanidae] is based are primitive
or derived cannot be answered with certainty. Although it is often
comparatively easy to make this decision regarding somatic characters
in spiders, the opposite is generally true for genitalic characters. Our
understanding of the evolution of spider genitalia is extremely limited,
because of the astounding diversity of structures... ”

N. I. Platnick (1975: 2)

ABSTRACT. A detailed morphology of the male copulatory organs, the genital bulb,
in Architis, Charminus, Dolomedes, Paradossenus, Pisaura, Pisaurina, Staberius,
Thaumasia and Trechalea is presented. The homology of several palpal elements
within the Pisauridae is established; possible homology with those in various other
spider families is discussed. Homology concepts concerning the median apophysis,
the conductor, and the spiral embolus are discussed. The character states are
evaluated and polarized. The groundplan of the pisaurid genital bulb is compared
with the bulb in Mesothelae. An update on the systematics of the family Pisauridae
is presented and systematic implications from the analysis of the male copulatory
organs are discussed. Two large monophyletic genus-groups, a mainly African
Pisaura genus-group including about 20 genera, and a Trechalea genus-group
comprising eleven South American genera, are recognized. Paradossenus taczanowskii
Caporiacco, 1948, is a junior synonym of P. nigricans F. O. Pickard-Cambridge,
1903. The validity of two nominal genera, Architis and Staberius, is discussed. The
genera are likely to be synonymous. The structure of the egg sacs of Syntrechalea,
Dyrines and Hesydrus are described for the first time.

Research Associate Serial Publication
Delaware Museum of Natural History ISSN 085-3887

2 NEMOURIA No. 35

Table of Contents

ADStract » 6006 see ew ae ew bes be oat, Bee ]
Introduction ... . 0.6. . fei Ge 626 A Sh we us 2
Materials and Methods .... . 2. i.2 ss. 4 2 A ee ee 4
Systematics of the Family Pisauridae, An Update. 2:2 eee 8
Review: Homology in Palpal Structures
(1) Hypotheses on the Evolution of the Male Genital Bulb... .12
(2) Literature Overview = =... 2 ss «i awe eee 13
(3) Ontogenetic Evidence®... . 34 i. «2.25 oe ee 15
Results
(1) Morphology of Palpal Elements... .”"».... “2 16
(2) Groundplan of the Pisaurid Palpal Organ ..~ ') =) 3 32.2 18
(3) Palp Morphology in American Nursery-Web Pisauridae . . . .22
(4) Palp Morphology in Other Nursery-Web Pisauridae...... o)
(5) Palp Morphology in the South American
Tvrechalea Genus-Group .2 2 «22.2 4s 2 ee ee 33
Discussion
(1) Homology of Palpal Elements within
Pisauridae and Evaluation of Character States.......... 35
(2) Systematic Implications : ... 2.2.22... 4 eee 40
(3) Homology of Pisaurid Palpal Elements
with Those of Other Families. < «. <9 ps tees eee 42
SUMMAaLy 2.6 4 a Sade Bh dhe was 6 GA Soe ee 48
Acknowledgments ... 2.5 5 4 5 %s #4 Se eerste 51
Literature Cited . ss .55 6 63s es ew eee % Ce eee 52
Endnotes s.2 « 9.2 Wh Ae See So ee Ne 59
INTRODUCTION

The copulatory organs in male spiders are located at the distal segment
of their pedipalps. They are secondary organs serving the transfer of sperm.
They are not connected to the testes but have to be charged prior to
copulation. The male copulatory organs form a highly complex morphologi-
cal structure, the genital bulb. It functions hydraulically and is composed
of inflatable membranes (hematodochae) and larger sclerites (subtegulum
and tegulum). Additionally, projections and processes, often called apophy-
ses, are attached to various parts of the palp. Inside the genital bulb lies
the sperm duct that serves as a storage place for sperm. The sperm duct
opens at the tip of the embolus, an intromittant organ. During copulation,
the embolus is partly or totally inserted in the female epigynum to transfer
sperm.

April 30, 1990 Petra Sierwald 3

The male genital bulbs are diverse in structure and in most cases
species-typical. They have been used for species identification since the
study of male sexual organs started with the works by Westring (1861),
Blackwall (1843) and Menge (1843, 1866-1879). An important work on
details of their morphology was published by Comstock (1910). He studied
unexpanded and expanded bulbs of different spider groups and introduced
a nomenclature of hematodochae, sclerites and apophyses that is still used
today. There are numerous taxonomic monographs that illustrate male
(and female) copulatory organs. But accurate labelling of those figures is
rare. Often, Comstock’s nomenclature is applied uncritically, with the
result that homology of sclerites is inevitably implied but seldom discussed.
Since homology of many elements in the male genital bulbs has never
been clearly established, the use of these organs for phylogenetic reconstruc-
tion is hampered.

The theme of this paper arose from a study on the systematics of the
spider family Pisauridae. The systematics of the Pisauridae is unsatisfacto-
rily resolved. The sister group is unknown, the superfamilial position is
in dispute, and currently used subdivisions within the family are not
monophyletic. In this study, characters of the male copulatory organs will
be analyzed to clarify systematic problems of Pisauridae.

in this endeavor, there are two main obstacles. First, the structures
of male genital bulbs of Pisauridae are poorly known. Roewer’s (1955)
drawings and labelling are not accurate enough to reveal the morphology
of the genital bulb in African pisaurid genera he revised and are useless
for systematic and phylogenetic conclusions. Blandin’s (1974-1979) work
on African and Carico’s (1972-1981) work on American Pisauridae provide
better insight into the morphology of palps of particular genera. However,
male organs were rarely studied in their expanded form. Therefore, the
serial relationships of palpal sclerites and apophyses are mostly unknown.
Secondly, morphological features found in genital bulbs have to be
evaluated regarding the question of homology. Only homologous characters
with their character states furnish data for a cladistic analysis. The main
objective of the present study is the definition of homologous structures
and homologous sectors of the male genital bulb within Pisauridae and to
discuss some aspects of homology of palpal structures in Araneae in general.

Although the male copulatory organs in spiders show, as Eberhard
(1985) discussed, rapid and divergent evolution, the morphological ground-
plan of palps appears to be unexpectedly old. In the third part of the
DISCUSSION some rather provocative (and speculative) hypotheses
concerning the homology of embolus, median apophysis and conductor are
presented. Homology concepts are discussed in the light of other such
hypotheses put forward by Kraus (1978), Haupt (1983), Raven (1985) and
Coddington [in press].

4 NEMOURIA No. 35

MATERIALS AND METHODS

Genital bulbs of the following species were studied (locality and
collection included):

Architis cymatilis Carico, 1981. d preserved with expanded palp [Peru:
Madre de Dios, Zona Reservada Pakitza (11°58’S, 71°18’W); Museo
de Historia Natural, Universidad Nacional Mayor de San Marcos
(MHNSM), Lima, Peru].

Architis tenuis Simon, 1898 [Brazil: Mato Grosso, Vila Vera, Oct. 1973;
American Museum of Natural History (AMNH), New York. Colombia:
Mata, Puerto Lopez; Museum of Comparative Zoology (MCZ),
Cambridge, USA].

Charminus camerunensis Thorell, 1899 [Cameron: Kitta; syntypes; Naturhis-
toriska Riksmuseet (NHR), Stockholm].

Dyrines striatipes (Simon, 1898) [Peru: Madre de Dios, Zona Reservada
de Manu, Puesto de Vigil. Pakitza, quebrada El Bano (11°58’S,
71°18°W) 5.X.1987; National Museum of Natural History (USNM),
Washington, DC. 2 with egg sac; Peru: Amazonas, alto Rio Comaina,
Puesto de Vigilancia, 850-1150m; MHNSM].

Dolomedes scriptus Hentz, 1845 [USA: Virginia, Lynchburg, April 1985;
USNM].

D. tenebrosus Hentz, 1843 [USA: Washington, DC; USA: Virginia,
Lynchburg, April/May 1985; USNM].

D. triton (Walckenaer, 1837) [USA: Virginia, Lynchburg; USNM].

Enna O. Pickard-Cambridge, 1897, unidentified species [Bolivia: USNM.
Peru: Piura, Rio Mangas, 1500m; MHNSM].

Hesydrus Simon, 1898, unidentified species [Peru: Amazon, Cordillere del
Condor, alto Rio Comaina, Puzz, Falso Paynisha, November 1987;
MHNSM. & with egg sac; Venezuela: Muséum Nationale d’Histoire
Naturelle, Paris (MNHN) Boc 2042, Boc 2043, Boc 2044].

Paradossenus nigricans F. O. Pickard-Cambridge, 1903 [Guyana: Two
Mouths, Essequibo River; Museo Zoologico della Specola (MZS),
Firenze, Italy. Guyana: Tibicuri-Cuyaha, Demerara River; Capori-
acco’s specimens, type of P. tascanowskit Caporiacco, 1948; MZS].

Pisaura mirabilis (Clerck, 1758) [Austria: Tirol, Innsbruck, June 1964;
USNM].

April 30, 1990 Petra Sierwald 5

Pisaurina mira (Walckenaer, 1837) [USA: Virginia, Lynchburg; USNM.
USA: New York, Bear Mt., May 1947; AMNH].

Staberius spinipes (Taczanowski, 1873) [Peru: Pucallpa, Nov. 1946; AMNH].

Syntrechalea F. O. Pickard-Cambridge, 1902, unidentified species. 2 with
egg sac [Peru: Cusco, Atacaya; MHNSM].

Syntrechalea porschi Reimoser, 1939 [Costa Rica: Limén, Hamburg Farm;
Naturhistorisches Museum, Wien, Austria (NHMW)].

Thaumasia cf. uncata F. O. Pickard-Cambridge, 1901 [South America,
country unknown, “San Esteban’; MNHN Boc. 2047b, No. 11750.
Mexico: Veracruz (95°07’W, 18°36’N); MCZ]. Carico (1981: 150) states
that Thaumasia argenteonotatus (Simon, 1898) is a senior synonym of
Thaumasia uncata.

Thaumasia cf. velox Simon, 1898 [Colombia(?): ‘Col. Alpine’; MNHN,
Boc. 2047b, No. 19190. Ecuador: Morona-Santiago, Suctia, 1000m,
June/July 1979, N. Engler; MCZ].

Tinus peregrinus (Bishop, 1924) [Mexico: Nuevo Leon, Linares, July 1956;
AMNH].

Thalasstus spinosissmus (Karsch, 1879) [South Africa: Natal, Pietermaritzburg;
USNM].

Trechalea Thorell, 1870, unidentified species [Venezuela: Miranda, Guatopo,
National Park Santa Cruzita, 450m, J. Coddington; USNM].

Expansion of genital bulbs

Palps were submerged overnight in a weak watery solution of
potassium hydroxide (KOH), in which they expanded to varying degrees.
They were then transferred to distilled water, where further inflation
occurred. The structure of unexpanded bulbs was studied by submerging
them in Hoyer’s mounting medium for the time of observation. They
became partly translucent. The drawings were made with aid of a dissecting
microscope with drawing tube.

Homology

Classical criteria were applied to reach decisions on the homologous
status of features: (a) criterion of position, (b) criterion of special,
morphological similarity, and (c) criterion of concordance with other
characters (Remane, 1952; Wiley, 1981: 130-138; Patterson, 1982). To allow
comparison of the positions of apophyses on the tegulum in different
genera, the terms proximal and distal are used as defined below in order
to accommodate criterion (a).

6 NEMOURIA No. 35

Terms

Differing from common usage, the terms proximal and distal in the
tegulum here refer to the position of a particular part in relation to the
trajectory of the sperm duct. Following the course of the sperm duct,
starting at the fundus, apophyses inserting on the tegulum near the fundus
are considered proximally located. Apophyses inserting closer to the
ejaculatory duct are referred to as being distal. The terms dorsal and ventral
refer to the position of a particular part within the depth of the genital
bulb. Parts nearest to the dorsal surface of the cymbium are located
dorsally; ventrally located parts are mostly visible in the unexpanded bulb
in ventral view (as they are figured here).

In descriptions of pisaurid palps, the terms median apophysis and
conductor are used to name tegular apophyses that are proposed to be
homologous within (at least) all Pisauridae. The term terminal apophysis
is used for any apophysis of the apical division; thus homology among
different pisaurid genera is not implied.

The term homologous refers to the actual structures (= character,
e.g., conductor) that are proposed to be based on an identical evolutionary
event. The particular structure may appear in several character states.
Similar character states of a character in several taxa may be based on the
same evolutionary event, thus the character state in question is considered
homologous and representing a synapomorphy. Apparently similar charac-
ter states of one character based on different evolutionary events have to
be considered homoplasous, and thus do not indicate phylogenetic
relationship (see Patterson, 1982: 25).

Text conventions

All statements of positions of sclerites refer to the unexpanded left
bulb in ventral view, unless indicated otherwise.

All references to figures presented in this paper are made using the
abbreviation “Fig.’’; references to figures in other publications are
abbreviated as “‘fig.”” Terms used by other authors to describe parts of the
male copulatory organs are added in parentheses to facilitate finding the
cited text.

Abbreviations (including terms of authors)

ad I, II, III bulbous sections of the apical division in Pisaurina

an anelli of the subtegulum

bmt basal membraneous tube of apical division

bl broad lamella at sclerite III in Liphistius sensu Kraus, 1978 (Haupt,
1983: Basalkante des Embolus)

(s conductor (Menge, 1866: Spermophorum; Kraus, 1978, and Raven,
1985: sclerite III [in part])

April 30, 1990 Petra Sierwald 7

cy
da

cymbium (= Schiffchen, by many German authors)

distal apophysis in the Prsaura genus-group

dorsal branch of distal apophysis in Pisaura and Charminus

distal tegular projection

distal sclerotized tube of apical division

embolus (Menge, 1866: Eindringer; Harm, 1931, 1934: Stylus; Kraus,
1978, and Raven, 1985: sclerite III [in part])

fundus of sperm duct

fulcrum

hyaloid lamella sensu Kraus, 1978, at the embolus of Liphistius
lateral subterminal apophysis in Dolomedes

median apophysis (Haupt, 1983: Kontrategulum)

median hematodocha (Osterloh, 1922: Haematodochula)

petiolus (Osterloh, 1922: Chitinretraktor)

palea in Lycosidae

pars fundi of the subtegulum (Loerbrocks, 1984: 386; Chamberlin,
1904: lunate plate; Osterloh, 1922: vallum chitinosum; Lamoral, 1973:
ental boss of subtegulum)

prolateral (ental)

pars pendula of the embolus (Comstock, 1910: 183)

proximal tegular projection in Thaumasia and Tinus

retrolateral (= ectal)

retrolateral tibial apophysis

sclerite III in Liphistius

saddle (in Dolomedes)

sail-shaped apophysis in Pisaurina

sperm duct (Osterloh, 1922: Spermakanal; Bertkau, 1875: Spermo-
phor; Wagner, 1888: Receptaculum seminis)

subtegulum (Wagner, 1888: s.Teg.; Osterloh, 1922: Basalschale,
Patina basalis; Harm, 1931, 1934: Basalschale; Gassmann, 1925:
Samenkapsel [in part]; Kraus, 1978, and Raven, 1985: sclerite I)
subtegular apophysis

tegulum (Osterloh, 1922; Harm, 1931, 1934: Spermakanalkapsel;
Gassmann, 1925: Samenkapsel [in part]; Kraus, 1978, and Raven,
1985: sclerite IT)

terminal apophysis

truncus of the embolus (Comstock, 1910: 183; Osterloh, 1922:
Chitinfaden)

ventral branch of distal apophysis in Pisaura and Charminus

ventral tibial apophysis

8 NEMOURIA No. 35
SYSTEMATICS OF THE FAMILY PISAURIDAE, AN UPDATE

The nursery-web, a web built by the female for the freshly hatched
spiderlings, is assumed to be a characteristic feature of members of the
family Pisauridae. Pisaurid females carry the spherical egg sac in their
chelicerae! until the spiderlings are due to hatch. They then produce an
irregular web and place the egg sac in its center. To date, a nursery-web
has been reported from nine of the 55 pisaurid genera belonging to all
three subfamilies sensu Simon, Pisaurinae, Dolomedinae and Thalassiinae,
see Table I (Afropisaura Blandin, 1976 [see Blandin, 1976 and 1979b: 82];
Dolomedes Latreille, 1804, and Pisaura Simon, 1885 [see Gerhardt &
Kastner, 1938]; Thalassius, Simon, 1885 [see Sierwald, 1989a]; Pisaurina
Simon, 1898 [see Carico, 1972: 303]; Tinus F. O. Pickard-Cambridge, 1901
[see Carico, 1976: 301]; Megadolomedes Davies & Raven, 1980 [see Davies
& Raven, 1980: 139]; Architis Simon, 1898 [see Nentwig, 1985: 301] and
Ancylometes Bertkau, 1880 [see Merrett, 1988: 200]). So far, the nursery-
web is the only synapomorphy proposed for the family Pisauridae. Thus,
to assign a genus to the Pisauridae requires either the discovery of the
nursery-web in such a genus or the demonstration of morphological
synapomorphies relating such a genus to at least one of the nine genera
listed above.

Carico (1986: 305) suggested that ‘“‘a large group” of South American
pisaurid genera, including the genus Trechalea, constitute a distinct
monophyletic group, but gave no further details on the group’s possible
apomorphic characters. Simon described the family Trechaleidae (1890:
82; consisting of the genera Trechalea and Dendrolycosa Doleschall 1859);
later he (1898: 278) listed Trechalea under Pisauridae. If these South
American genera can be recognized as a distinct group, the family name
Trechaleidae could be used (Carico’s [1986: 305] statement that the name
Trechaleidae is not available because it is a nomen oblitum is not
corroborated by the third edition of the International Code of Zoological
Nomenclature [ICZN, 1985: 260]). The spiders of this group are poorly
known; all genera are currently unrevised (revisions of these genera are in
preparation, teste Carico & Minch, 1981: 154).

The females of Trechalea carry the egg sac attached to the spinnerets. |
The egg sac is hemispherical and possesses a seam. The upper-surface is —
vaulted and consists of thick silk. The bottom layer is flat and the silk is —
thin, the eggs are visible through that bottom layer (Simon, 1898: 281; |
Berkum, 1982: 120). The scar of the attachment to the spinnerets is visible |
on the upper-surface of the egg sac even in preserved specimens. The |
same type of egg sac and mode of attachment was found in preserved |
specimens of Hesydrus (Venezuela, MNHN), Dyrines striatipes (Peru;

April 30, 1990 Petra Sierwald 9

MHNSM), and Syntrechalea (Peru; MHNSM) [pers. observ.]. Apparently,
Trechalea does not produce a nursery-web, but the spiderlings remain on
the egg sac and opisthosoma of the female after hatching (Carico et al.,
1985: 292, fig. 5; and pers. observ., Panama, Barro Colorado Island). It
is proposed here that the structure of the egg sac represents a synapomor-
phy for these South American “pisaurid” genera. They all may lack a
nursery-web. Unless synapomorphies can be found linking the Trechalea
genus-group and any nursery-web pisaurid, both groups may not even be
closely related?.

The majority of the remaining pisaurid genera (circa 25) occur in
Africa. Some genera (circa 5, e.g., Perenethis L. Koch, 1878, Thalassius
Simon, 1885, Euprosthenops, Pocock, 1897) contain African and Asian
species. A few genera are known from Asia and the palearctic region, with
some species also in Africa (e.g., Eurychoera Thorell, 1897, Pisaura Simon,
1885, Polybaea Thorell, 1895, Eucamptopus Pocock, 1900, and Hygropoda
Thorell, 1894). The family is represented in the Australian region (e.g.,
Megadolomedes Davies & Raven, 1980, Inola Davies, 1982, Anoteropis L.
Koch, 1878). The more than 100 nominal species of Dolomedes Latreille,
1804, are distributed worldwide. Five genera (see below) of nursery-web
Pisauridae (and Dolomedes) occur in the Americas.

Monophyletic subunits have not been defined for Pisauridae. Simon’s
(1898) subdivisions of Pisauridae (Pisaurinae, Dolomedinae, Thalassiinae)
are still in use; Sierwald (1987) showed that his Thalassiinae is not a
monophyletic clade. This study identifies one monophyletic unit within
the nursery-web pisaurids, the Pisaura genus-group, based on apomorphies
in the copulatory organs.

Thus far, the monophyly of Pisauridae is unsupported by morphologi-
cal synapomorphies. The sister group is unknown. Several families have
been proposed as relatives of Pisauridae: Dondale (1986: 328) used
Pisauridae as a sister group to Lycosidae; Brady (1964: 436) suggested a
close relationship to Oxyopidae. Lehtinen (1967) assigned the pisaurid
genera to three different families, Dolomedidae (= Thaumasiinae auct.),
Pisauridae sensu stricto (Pisaurinae auct.), and Thalassiinae in the Ctenidae.
He placed Dolomedidae and Pisauridae in two different superfamilies.

Lehtinen (1967: 372) based the placement of Dolomedidae (in
Lycosoidea, see Table 1) and of Pisauridae (in Pisauroidea; 1967: 379) on
the presence or absence of a secondary conductor. Lehtinen’s definitions
of primary and secondary conductor (1967: 412) are ambiguous. While he
defines a primary conductor ‘‘as a tegular process of variable shape and
Structure that acts as the primary support for the distal part of the
embolus”, his definition of the secondary conductor lacks details about its

Structure and place of attachment within the genital bulb, thus making it

10 NEMOURIA No. 35

impossible to identify a primary or secondary conductor. As Lehtinen’s
phenogram shows (page 285, fig. 6), he did not consider the conductor of
Dolomedes and Pisaura as homologues but as analogous structures. The
Opposite position is taken in this paper as discussed later and the placement
of nursery-web pisaurid genera in three families and two superfamilies is
rejected.

“In cases where the primary conductor of the group concerned has
lost its function, the analogous structure is called the secondary conductor
(some Amaurobioidea [sensu Lehtinen] and Gnaphosoidea as well as all
Lycosoidea), unless it is a sclerite or part of one that is generally named
in another way” (Lehtinen, 1967: 412). He states (1967: 412) that “‘it is
usually impossible to show absolute homology between different superfa-
milies and thus the term conductor refers to homological structures only
in restricted groups.” The diversity of apophyses, either reductions in
certain genera or the appearance of apophyses unique to a subunit, may
be great within certain families, as it will be shown in this study. Detailed
data on the morphology of palpal apophyses are extremely limited and, at
this point, do not allow predictions concerning homology within larger
groups.

Homann (1971: 263) included Pisauridae, Ctenidae and Rhoicininae
at the level of subfamilies in the Lycosidae. So far, Homann’s definition
of the Lycosoidea (Table 1) based on the grate-shaped tapetum of the
secondary eyes, is the only suggestion for the placement of Pisauridae that
is founded on a clear synapomorphy. Therefore, the palpal structures of
other Lycosoidea are of special importance, since the sister group of
Pisauridae is most likely to be found within Lycosoidea.

April 30, 1990 Petra Sierwald 1]

Table 1: Comparison of the classifications of Pisauridae and pisaurid
subunits by Simon, Lehtinen and Levi.

Simon, 1898 Lehtinen, 1967 Levi, 1982
(based on Homann, 1971)

Lycosoidea Lycosoidea
Cycloctenidae
Selenopidae
Lycosidae — Lycosidae
Zoridae — Zoropsidae
Pisauridae
Thalassiinae — Ctenidae incl. — Ctenidae
Thalassiinae,
Acanthocteninae —— Acanthoctenidae
Dolomedinae — Dolomedidae — Pisauridae sensu lato,
(= Thaumasiinae incl. Thalassiinae,
of authors) Dolomedinae, Rhoici-
ninae, Pisaurinae
Pisauroidea
Pisaurinae — Pisauridae (sensu
stricto)
Oxyopidae — Oxyopidae
Senoculidae — Senoculidae
Homalonychidae
Toxopidae (isolated Toxopidae
Psechridae and Psechridae
derived)
Amaurobioidea
Amaurobiidae
Stiphidiinae — Stiphidiidae
Rhoicininae

(and Miturgidae, Amaurobiidae,
Liocranidae, Agelenidae, Dictynidae,
and Hahniidae)

Forster & Wilton (1973: 15-17) listed Lycosidae, Dolomedidae and
Pisauridae as having ‘‘probably been derived from Amaurobioid stock.”
Their Amaurobioidea include Amaurobiidae, Agelenidae, Stiphidiidae,
Amphinectidae, Neolanidae, Psechridae, Ctenidae, and Cycloctenidae.

12 NEMOURIA No. 35
REVIEW: HOMOLOGY IN PALPAL STRUCTURES

(1) Hypotheses on the Evolution of the Male Genital Bulb

Wagner (1886, cited from Osterloh, 1922: 329) presented the following
hypothesis. A cavity in the metatarsus of the pedipalp was used originally
for the transfer of sperm. The metatarsus differentiated later in phylogeny
to form the genital bulb. The basal hematodocha originates from the
membrane of the joint between the metatarsus and tarsus; the cavity of
the tarsal joint is the predecessor of the alveolus. Later (1888: 65), Wagner
rejected this hypothesis and suggested that an accidental projection of the
tarsus preceded the genital bulb.

Considering the structure of the intima of the sperm duct, Comstock
(1910: 163) concluded that the sperm duct was an invagination of the
body wall ‘‘like a trachea,” surrounded by a specialized tip of the tarsus,
the genital bulb.

Barrows (1925) considered the genital bulb as a homologue of the
palpal claw and its teeth to be the predecessors of accessory structures
(sclerites and apophyses) of the genital bulb. Harm (1931, 1934), Kraus
(1978, 1984) and others followed Barrows. Kraus (1984: 374, figs. 1la-f)
presented a hypothetical transformation series from a simple tarsal claw
to a tripartite hydraulic bulb. Harm (1934: 133) stated that the embolus
is a homologue of the tarsal claw. Raven (1985: 15) argued that the
plesiomorphic state of the embolus is fully sclerotized (versus flexible and
soft), since “‘the embolus was presumed to have been derived from the
totally sclerotized tarsal claw’? [see DISCUSSION under (3)].

The ontogenetic evidence (see below) reveals that the genital bulb is
formed by the claw fundament at the distal end of the tarsus. Coddington
[in press] stated that the “hypothesis that the genital bulb is a literal,
transformed homologue of the palpal claw is rather decisively refuted’’,
since rudimentary palpal claws are often contemporaneous with the
developing genital bulb.

Until recently, the simple pyriform genital bulbs of Mygalomorphae
and ‘“‘haplogyne”’ spiders were considered primitive for all spiders, meaning
phylogenetically old (e.g., Wagner, 1888: 65; Osterloh, 1922: 329; Gerhardt
& Kastner, 1938: 530; Wiehle, 1960: 458; Barnes, 1980: 632). The more
complex bulbs of entelegyne spiders were considered derived. As Gertsch
(in Platnick & Gertsch, 1976) and Kraus (1978) pointed out, this hypothesis
was challenged by the fact that primitive, segmented spiders (Mesothelae,
Liphistiidae) possess a complex, clearly tripartite genital bulb. The
Mesothelae (family Liphistiidae) are believed to be very similar to the
earliest known Araneae from the Carboniferous period (300 million years
ago; Foelix, 1987: 261). Today, the pyriform and simple bulbs of

April 30, 1990 Petra Sierwald 13

Mygalomorphae and “haplogyne’’ spiders are regarded as secondarily
simplified and therefore derived (Platnick & Gertsch, 1976; Kraus, 1978;
Schult, 1983a, 1983b; Haupt, 1983; Raven, 1985; Coddington [in press]).

(2) Literature Overview

Early Studies. — The study of the structure of the male palpal organs
began with the works by Blackwall (1843), Westring (1861), and Menge
(1843, 1866-1879). Menge (1843: 35, pl. III figs. 13-27) distinguished three
divisions and apophyses within the genital bulb. Bertkau (1875) produced
the first histological sections of palps (Segestria) and described for the first
time pores in the wall of the sperm duct. Hasselt (1889) analyzed the fine
structure of the hematodocha. He found that the hematodocha does not
consist of musculature as Menge believed. Wagner (1888: 65) gave
schematic drawings of different palp types, and drawings of expanded
genital bulbs. He used (1) the shape of the cymbium to divide the Araneae
into four subgroups, and (2) the shape and the processes of the tegulum
to define families.

Comstock (1910), who studied Filistata, Hypochilus, Loxosceles,
Pachygnatha, Araneus, Linyphia, and Dolomedes, produced the most
influential work on spider palps. His extensive nomenclature is still in use
today. Since he developed the more elaborate terms for members of the
Araneoidea, the use of his terms in other, unrelated groups led to serious
misapplications (e.g., the median apophysis sensu Baum [1972, figs. 62-64]
in Oecobiidae is definitely no homologue of the median apophysis sensu
Comstock in Dolomedes or Araneus [Comstock, 1910: figs. 22, 23)).
Comstock (1910: 161) was concerned about homology as his introduction
showed, “we find different terms applied to homologous parts...”
Obviously, he intended to apply identical terms to homologous structures.
But he himself used some terms inconsistently (e.g., in Dolomedes [fig. 22]
and Araneus [figs. 19, 21], the conductor clearly inserts at the tegulum,
while in Linyphia [fig. 10], it is part of the apical division).

Osterloh’s (1922) study on palps of Linyphia, Agelena, Lycosa, and
Meta included histological sections as well. His drawings of the expanded
palps are excellent. He assumed that the genital bulb would consist of
homologous parts and tried to identify them within the species he studied.
He recognized (1922: 352) the homology of subtegulum and tegulum based
on morphological similarity and position within the genital bulb. He also
proposed homologies of processes (= retinacula), e.g., homology of terminal
-apophyses in Lycosa and Linyphia (1922: 356, stema-retinaculum), but
those proposals were not convincing. He described in detail the function
| of both male and female copulatory organs during copula.
| The function of the genital bulb is still poorly understood. The modes

14 NEMOURIA No. 35

of sperm intake and ejaculation are not known with certainty (Harm, 1931:
645; Osterloh, 1922: 415; Wiehle, 1967: 480; Cooke, 1966; Lamoral, 1973:
638-644). Few studies have dealt with the role of particular parts of the
genital bulb during copula (Heimer, 1982; Levi, 1961; Loerbrocks, 1983,
1984; Schult & Sellenschlo, 1983; Sierwald & Coddington, 1988; Gering,
1953).

Recent Studies on the Homology of Palpal Sclerites. — In numerous
taxonomic monographs, the male (as well as the female) copulatory organs
are well figured, but in the form of finger-print patterns for identification
purposes only. Detailed morphological analyses that include discussions
on homology of elements and labelling of elements are often lacking. Such
drawings may furnish only limited data for future comparative and
analytical studies. Several authors (e.g., Baum, 1972: figs. 49, 62-64;
Gering, 1953: figs. 4, 6; Grasshoff, 1968: fig. 33; Haupt, 1983: figs. la-e;
Heimer, 1986: fig. 1; Loerbrocks, 1984: figs. 8, 12; Merrett, 1963: figs.
1-3; Saaristo, 1971: figs. 1; Millidge, 1977: figs. 1-8; Schult, 1983a: fig.
16) produced schematic or generalized drawings of palps of the groups
under study. Such drawings are especially valuable, since they present the
authors concept of homology within the group studied. Additionally, such
drawings illustrate more clearly the serial relationships of sclerites and
apophyses. The notion that simple bulbs are primitive and complex bulbs
are derived often caused the listings of plesiomorphic characters as
characteristics (1.e., apomorphic) of a particular group (e.g., Millidge, 1980:
98).

Statements on homology are sometimes vague, making it difficult for
subsequent workers to evaluate the presented data and conclusions (e.g.,
Haupt [1983: 277]: ‘‘a conductor can be found in Heptathelidae as well
as in entelegynes ... [citation translated].’’ Judging from the context, Haupt
intended to imply homology).

[In order to solve systematic problems in Linyphiidae, several attempts
have been made using genitalic characters (Blauvelt, 1936; Merrett, 1963;
Saaristo, 1971; Miullidge, 1977; for discussion see Coddington [in press]).
Saaristo (1971) illuminated an important autapomorphy of the linyphiid
palp: the suprategulum, a distal prolongation of the tegular ring, which
had been named median apophysis by previous authors. Saaristo showed
that the suprategulum surely is not a homologue of the araneid median ~
apophysis. All authors emphasized the problems and uncertainties of —
homologizing apophyses of the apical division between linyphiid genera.
But especially the complex structure of the apical division (and of apophyses
of the suprategulum) present the most reliable generic characters in ©
Linyphiidae. Thus, the problems of homologizing hamper the use of these |
characters for phylogenetic reconstruction. ] |

April 30, 1990 Petra Sierwald 15

With the general acceptance of the hypothesis that the tripartite genital
bulb is plesiomorphic and that pyriform bulbs are derived, Haupt (1983,
figs. la-e) produced a scheme in which he homologized sclerites and
processes of genital bulbs in Mesothelae (Liphistius, Heptathela), Mygalo-
morphae (Atypus, Amblyocarenum) and Araneomorphae (Nuctenea, Ara-
neidae). His proposed homologies were refuted by Raven (1985: 14-17)
[see DISCUSSION under (3)].

Recently, the question of homology of palpal elements has received
more attention. Griswold [in press], in his taxonomic revision of Phyxelidi-
nae (Amaurobiidae), discussed the homology of palpal elements (and female
copulatory organs) for the subfamily in detail and based taxonomic
decisions on synapomorphies found in copulatory organs. Coddington [in
press] presented an extensive study focusing on the homology of palpal
structures, dealing with orb-weaving families (Orbiculariae) and their
possible outgroups. For the first time, the homologies of median apophysis,
conductor, and radix within the orb-weavers have been analyzed. He
showed that these terms were often applied to structures which are not
homologous even among orb-weavers but are autapomorphies for certain
groups. He reviewed the available data on the ontogeny of the male spider
palp and discussed their implications on homology and phylogeny for
Orbiculariae. Relevant results of the ontogenetic evidence are summarized
here (see below).

(3) Ontogenetic Evidence

The available ontogenetic evidence does not reflect the general pattern
of palp ontogeny, since only very few species in disparate families have
been studied (Salticus [Salticidae] by Wagner [1886]; Agelena similis
[Agelenidae] by Szombathy [1915]; Steatoda borealis [Theridiidae], Phidip-
pus audax [Salticidae], and Lycosa nidicola [Lycosidae] by Barrows [1925];
Lepthyphantes nebulosa [Linyphiidae] by Gassmann [1925]; Segestria bavarica
[Segestriidae] and Evarcha marcgravi [Salticidae] by Harm [1931, 1934];
Latrodectus curacaviensis [Theridiidae] by Bhatnagar & Rempel [1962];
Lycosa chaperi [Lycosidae] by Sadana [1971]).

The claw fundament, responsible for the secretion of the dorsal claw
extensor and the ventral flexor tendon, produces the male genital bulb.
At a very early stage, the cell mass of the claw fundament is already divided
into a dorsal and ventral lobe. The ventral lobe will form basal, middle,
and apical division of the palp, including the sperm duct. The ventral lobe
divides two times, first early in the development to separate the apical
division from the still undivided basal and middle division of the bulb.
The separation of basal and middle division (= subtegulum and tegulum)
is a late event. The apophyses of the apical division develop late in

16 NEMOURIA No. 35

ontogeny.

The dorsal lobe of the claw fundament bifurcates very late in ontogeny
(studied in Latrodectus only) to form two tegular apophyses (the median
apophysis and the conductor in Latrodectus). Important in the context of
this study is the status of the theridiid median apophysis. As Levi (1961:
3) already suspected, the tegular apophyses in different theridiid genera
may not be homologues. Coddington [in press] concluded that those
“‘theridiid median apophyses containing loops of the sperm duct’? may
just be projections or outgrowths of the tegular wall [called theridiid tegular
apophysis (TTA) by Coddington]. But in certain theridiid genera (e.g.,
Archaearanea, Theridion, and Latrodectus), the sperm duct does not pass
through the median apophysis. It is most parsimonious to assume that the
tegular apophysis in Latrodectus is a homologue of the median apophysis
as it is found in different groups of Orbiculariae (see Coddington [in press])
and in Pisauridae. But its homology is not established. Therefore, the
ontogenetic evidence concerning the origin of the median apophysis and
the conductor from the dorsal lobe of the claw fundament in Latrodectus
allows at present no conclusions for other spider groups, not even for other
Theridiidae [see DISCUSSION under (3)].

The ontogenetic evidence suggests that a tripartite palp is primitive
and therefore subtegulum and tegulum are homologous within Araneae.
The complex palpal structures in Mesothelae and in Atypus among the
Mygalomorphae are concordant with this hypothesis.

RESULTS

(1) Morphology of Palpal Elements

The genital bulb of male spiders lies in a cavity (alveolus) of the
cymbium, the modified tarsus of the pedipalp. Projections or outgrowths
may occur at the patella, tibia (tibial apophyses) or cymbium (paracym-
bium). The genital bulb itself consists of three divisions, the basal, middle
and apical (= embolic) divisions. The basal division contains a usually
large, inflatable, membraneous tube, the basal hematodocha, and an often
funnel-shaped sclerite, the subtegulum. Parts of the wall of the basal
hematodocha may be sclerotized, forming the petiolus. The middle division
consists of the median hematodocha and the sclerotized, often ring-like
tegulum. The median hematodocha, connecting subtegulum and tegulum,
may be large or small. The upper-surface of the tegular ring may be
membraneous and sometimes inflatable. It can also be partly or totally
sclerotized and more or less continuous with the tegular ring. In such
cases, the tegulum appears plate-like or like an upside-down bowl.

Habitually, all structures distal to the tegulum are assigned to the

April 30, 1990 Petra Sierwald 17

apical division (= embolic division; Osterloh [1922], Gassmann [1925]:
Stema). The apical division may contain as little as the embolus alone
(e.g., Paradossenus, Fig. 47; Heriaeus [Thomisidae, Loerbrocks, 1983: fig.
6]; Evarcha [Salticidae, Harm, 1934: figs. 1-5]; Loxosceles [Loxoscelidae,
Comstock, 1910: fig. 4]). In certain families, it is very elaborate (e.g., in
Linyphiidae and Araneidae), consisting of sclerotized tubes and membrane-
ous tubes (often labelled distal hematodocha, Comstock, 1910: 177, figs.
13, 14; Osterloh, 1922: Stemablase), bearing the embolus and none to
several processes of various size (often labelled terminal apophyses). The
apical division in Pisauridae is moderately complex.

Distal Hematodocha. — The term distal hematodocha has been used
for various membraneous, more or less inflatable parts within the embolic
division, being either connected to the embolus or to terminal apophyses.
Therefore, the term is strictly descriptive and only ambiguously defined.
It is not used in the present paper.

Embolus. — The term embolus is used for that part of the genital
bulb where the sperm duct opens. Comstock (1910: 182, fig. 25) described
three different types of emboli, based on their shape and supposed function:
the spiral embolus (occurring in many spider families, including Pisauri-
dae), the coniform embolus (e.g., in Araneidae) and a lamelliform embolus
(e.g., in Linyphiidae). He described only the spiral embolus in detail,
distinguishing different structures (a sclerotized truncus and a membrane-
ous pars pendula). Wiehle’s study (1960) on different types of emboli is
devoted to the function and does not give morphological details of different
emboli.

At this point, it is important to note that the emboli in different
families are not necessarily fully homologous [see DISCUSSION, (3),
under Embolus].

Sperm Duct. — In most spiders, the sperm duct follows a smooth
spiral from its blind end (fundus) in the subtegulum to the end of the
tegulum, where it becomes very thin (ejaculatory duct). It opens at or
near the tip of the embolus. More complex trajectories of sperm ducts
with several switchbacks occur in a few disparate groups and are surely
of non-homologous origin (and are apomorphies, whenever and wherever
they are found).

Processes of the Genital Bulb

Morphological Structure. — Besides the two large, basic sclerites,
subtegulum and tegulum, several sclerotized processes have been described
for male palps, occurring in the middle division of the bulb as tegular
processes, and in the apical division*. These processes have often been
named apophyses, e.g., median and terminal apophyses (labelled Retinacu-

18 NEMOURIA No. 35

lum? by several German authors). The homology of such processes
between and within different spider groups is the most ambiguous, yet
crucial issue for phylogenetic reconstructions. While labelling processes
many incorrect homologies have been implied.

Different structural types of processes can be distinguished. A
process can either be formed as a distinct sclerite, separated from other
sclerites by a flexible and somewhat inflatable membrane, or may be a
mere projection or outgrowth of a sclerite and directly connected to it.
These projections may be large and strongly sclerotized, as is the conductor
in Tinus (Figs. 11, 13), or occur as a low hump or prominence, as the
conductor in Architis (Fig. 42). Outgrowths of the tegulum may (or may
not) contain loops of the sperm duct (e.g., in several genera of Theridiidae,
Levi, 1961: figs. 5, 8, 9; labelled median apophysis). Such morphological
differences, as criterion of special similarity together with the criterion of
position, can be applied to recognize homologous processes. Such morpho-
logical qualities of processes are rarely considered in taxonomic mono-
graphs, descriptions and drawings.

Median Apophysis. — A median apophysis has been labelled in the
palps of many spider families (e.g., Pisauridae, Araneidae, Theridiosomat-
idae, Oececobiidae [including Urocteidae], Lycosidae, Theridiidae, Age-
lenidae, Anyphaenidae, Gnaphosidae, Ctenidae, Sparassidae). The term is
used almost always for a tegular apophysis. The homology of these
structures is, for many groups, unstudied.

Conductor. — A conductor has always been defined by its proposed
function: to protect and support the tip of the embolus (Comstock, 1910:
181; Wiehle, 1960: 460; Lamoral, 1973: 616). To my knowledge, there are
no data available to support this functional assumption. It is, however,
noteworthy that in many spider groups at least the tip of the embolus is
in close proximity to either the tip of a tegular or an apical apophysis.
Since such apophyses are usually labelled ‘“‘conductor’’, this term has been
used frequently for surely non-homologous apophyses, some of them
inserting at the tegulum (e.g., Araneidae, Pisauridae), some of them being
part of the apical division (e.g., Linyphiidae).

Terminal Apophyses. — Homologies of terminal apophyses have not
been established so far. Merrett (1963) and Saaristo (1971) discussed the
problems for Linyphiidae (see above: Literature Overview, Recent Studies).
Osterloh’s attempt (1922) did not yield convincing results (see above:
Literature Overview, Early Studies).

(2) Groundplan of the Pisaurid Palpal Organ
The Dolomedes bulb (Fig. 2) serves as a preliminary basis of the
groundplan of the pisaurid palp. It is not necessarily thought to be

April 30, 1990 Petra Sierwald 19

primitive or plesiomorphic among Pisauridae. But, the Dolomedes palp
is rich in structure and apparently not secondarily simplified. For this
study, tegulum and embolic divisions are broken into discrete sectors with
clear, recognizable limits and characteristic features. While analyzing bulbs
of other pisaurid genera, I attempt to identify the equivalent and
homologous sectors as defined in the Dolomedes bulb. Criteria to identify
homologous elements are (a) their position within the bulb, (b) similarities

Tibial Apophysis. — The majority of Pisauridae possess an apophysis
on the tibia of the male palp°. Usually, the tibial apophysis arises
retrolaterally, at the distal end of the tibia. Its shape may be simple or
very complex, but it is species-typical in all Pisauridae studied to date
(e.g., Figs. 20, 21; Thaumasia).

Basal Division of Genital Bulb. — All Pisauridae studied possess a
well-developed petiolus (Figs. 2, 13, 28; p). The petiolus is a strongly
sclerotized and thickened region within the wall of the basal hematodochal
membrane on the prolateral-ventral side. Its proximal tip is rounded and
fits into a complementary notch in the alveolus edge (Fig. 2). Its distal tip
is connected to the proximal end of the subtegulum via a tough membrane.

The subtegulum (Figs. 2, 8, 10, 11, 16, 22, 27, 31; st) is very similar
in all Pisauridae studied. It forms a funnel-shaped cup. The ventral-
prolateral part of the subtegulum is enlarged and often visible in the
unexpanded bulb. Under this conspicuous boss (= lunate plate sensu
Chamberlin, 1904: 174; Comstock, 1910: 176; pars fundi sensu Loerbrocks,
1984: 391; pf) lies the fundus (Figs. 2, 11, 37-47; f) of the sperm duct.
The prolateral side of the subtegulum bears the anelli (an). The anelli are
thickened sclerotized ridges. Between the anelli, the subtegular wall is
split. During inflation of the bulb, the clefts are spread (Figs. 2, 13, 32).
The margin of the subtegulum is a thick sclerotized rim that serves as a
region of attachment for the basal and median hematodochae. The basal
hematodocha is usually large. The median hematodocha is present in most
genera, but sometimes only weakly developed.

Middle Division of Genital Bulb. — The tegulum (Figs. 2, 5, 14,
37-47; t) [and the apical division of the genital bulb] differs considerably
among pisaurid genera and is described in detail for each genus. The
tegulum is a tubular, ring-like or plate-like sclerite. The upper-surface (or
distal surface) of the tegulum is membraneous and inflatable in some
pisaurid genera. In other pisaurid genera, it is partly sclerotized (Fig. 37,
saddle) and partly membraneous. In some Pisauridae, the distal end of the
open tegular ring is enlarged to form a distinct projection (Figs. 2, 5, 16,

22, 27, 35, 49; distal tegular projection, dtp).

In many genera, the apical division is attached below the tip of the
- distal tegular projection by a membraneous tube. The distal end of the

20 NEMOURIA No. 35

Figure 1. Schematic drawing of the male genital bulb of Liphistius batuensts
(drawing made after descriptions and SEM-photographs by Kraus, 1978: 237, figs.
3, 4). Figure 2. Schematic drawing of the left expanded genital bulb of Dolomedes.
Figures 3, 4. Dolomedes tenebrosus. 3. Expanded left genital bulb, ventral-prolateral
view. 4. Apical division of left expanded genital bulb, retrolateral view. Scale bars:
Fig. 3, 1 mm; Fig. 4, 0.5 mm.

April 30, 1990 Petra Sierwald 21

tegulum is defined here by the distal margin of the tegular sclerite. The
membraneous tube is therefore considered as the proximal part of the
apical division. There are two tegular apophyses, differing both morphologi-
cally and in their relative position on the tegulum (see below under Tegular
Apophyses).

Trajectory of Sperm Duct. — The sperm duct runs clockwise with a
spiral trajectory within the subtegulum and tegulum. The fundus is located
ventrally in the subtegulum, beneath the pars fundi. The sperm duct enters
the dorsal part of the tegular ring (Figs. 37-47), follows the dorsal part
to the retrolateral corner of the tegulum and continues its trajectory in the
ventral part of the tegular ring. The sperm duct extends nearly to the tip
of the distal tegular projection, turns and enters the apical division, thus
forming a loop (Figs. 2, 17, 23, 30, 44). The sperm duct narrows in this
region and becomes the ejaculatory duct. The loop of the sperm duct
identifies the distal tegular projection.

Tegular Apophyses. — An apophysis at the retrolateral corner of the
tegulum is traditionally (Comstock, Blandin, Carico) called conductor (c).
The conductor inserts directly on the tegulum and appears to be a mere
extension (e.g., in Pzsaura, Figs. 7, 45) of the tegular wall. In Pisauridae,
the conductor is often genus-specific, but never species-typical as the
median apophysis frequently is.

Approximately halfway along the ventral part of the tegular ring, there
is a second apophysis, traditionally called median apophysis (Figs. 2, 3,
5, 7-11, 16, 22, 27, 31, 34, 48, 49; ma)®. Positionally, the insertion of the
median apophysis on the tegulum is always closer to the apical division
than the insertion of the conductor. The median apophysis is attached to
the tegulum via an inflatable membrane. During expansion of the genital
bulb, this membrane is inflated and moves the median apophysis out of
its resting position. This different mode of attachment distinguishes the
median apophysis clearly from the conductor, which, as it is defined here,
is immovably attached to and continuous with the tegular wall. Often, the
tip of the median apophysis is distinctly sclerotized. The shape of the tip
may be genus-specific (as in Thaumasia, Pisaura), species-group-specific
(as in the tenebrosus- and fimbriatus-groups of Dolomedes; Carico, 1973:
figs. 25-33) or species-typical (as in Thalassius; Sierwald, 1987: figs. 1, 2,
54, 55, 58-68, 126, 127).

The terms median apophysis and conductor, as defined here, denote
homologous structures in all pisaurid genera.

Apical Division. — The apical division consists of a basal membrane-
ous tube (bmt) and a distal sclerotized tube (dst). The ejaculatory duct
extends through the basal membraneous and the distal sclerotized tubes.

_ Proximally, the basal membraneous tube is connected directly to the distal

22 NEMOURIA No. 35

tegular projection. Distally, it is connected to the sclerotized tube. The
distal end of the sclerotized tube bears the embolus. None to several
apophyses may be attached to both the basal membraneous [e.g., the distal
apophysis (da) in Pisaura; Fig. 7] and distal sclerotized tubes [e.g., the
fulcrum (fu) in Dolomedes; Fig. 6].

Embolus. — In most Pisauridae studied to date, the embolus can be
described as the spiral type sensu Comstock (1910: 181, 183). It consists
of a strongly sclerotized, gutter-like sclerite on its convex side (truncus of
the embolus, tr; Figs. 2, 14, 15). The truncus is basally broad and
scoop-shaped. It narrows distally. The concave side is covered by a
membrane (pars pendula, pp; Figs. 15, 17, 27, 30), thus forming a tube.
The pars pendula is large and inflatable in certain genera (members of the
Pisaura genus-group).

(3) Palp Morphology in American Nursery-Web Pisauridae

Carico (1973, 1972, 1976, 1981) provided good illustrations of the
male copulatory organs in the American species of Dolomedes, Pisaurina,
Tinus, Architis, and Staberius. The results obtained through the present
study suggest new and different interpretations of certain parts of the
organs. The genus Thaumasia, distributed in Central and South America,
is unrevised (revision in preparation, Carico, pers. communic.). Therefore,
the use of Thaumasia species-names has to be regarded as provisional.
Their copulatory organs are still poorly known.

Dolomedes Latreille, 1804 (Figs. 2-6, 37)

The tibial apophysis is large. The median hematodocha is present.
The tegulum is an open spiral (Fig. 37), not a closed ring, with a single
loop. The distal end of the tegular ring somewhat overlaps the proximal
end and forms a distinct distal tegular projection. The outer margins of
the tegular ring are rounded and U-shaped in cross section with the closed
portion being peripherally located. The upper-surface of the tegular ring
is partly sclerotized forming a saddle (Fig. 37, sa). The conductor is thin,
membraneous and lamelliform (Figs. 3, 5). The tip of the median apophysis
is sclerotized and flat. The apical division of the bulb (Figs. 3-6) contains
the basal membraneous tube (bmt) and the distal sclerotized tube (dst).
In the tenebrosus-group, the latter is very large and clearly visible in the
unexpanded palp (Fig. 5); it is somewhat smaller in the fimbriatus-group.

The distal end of the sclerotized tube (Figs. 4-6) bears a sclerotized
lamelliform apophysis (= lateral subterminal apophysis, Comstock, 1910:
181; Ista). Juxtaposed to this apophysis, another terminal apophysis called
fulcrum (Comstock, 1910: 181; fu) and the embolus arise (Figs. 4-6). In

April 30, 1990 Petra Sierwald 25

Figures 5-8. SEM-photographs, unexpanded left genital bulb. 5. Dolomedes
tenebrosus. 6. Dolomedes tenebrosus, apical division. 7. Pisaura mirabilis. 8. Thalassius
rubromaculatus. Scale bars: 0.25 mm.

24 NEMOURIA No. 35

Figures 9, 10. Thalassius spinosissimus, unexpanded left genital palp. 9. ventral
view. 10. retrolateral view. Scale bars: 0.25 mm.

the tenebrosus-group, the distal margin of the fulcrum is tightly wrapped
around the embolus. In the fimbriatus-group, the embolus follows a furrow
of the convex side of the fulcrum.

In Dolomedes, the embolus appears to consist of the truncus alone, the
pars pendula is not well developed.

Tinus F. O. Pickard-Cambridge, 1901 (Figs. 11-15, 38)

The conspicuous tibial apophysis in Timus arises in a more dorsal
position and bends retrolaterally. The median hematodocha is well
developed. The tegulum bears three processes (Fig. 11). Following the
sperm duct from the fundus in the subtegulum into the dorsal portion of
the tegulum, one encounters a large process of the tegulum on the prolateral
side of the bulb (proximal tegular projection, ptp; Figs. 11, 12, 14, 38).
This process is stiff and sclerotized. Next to the large conductor on the
retrolateral side is the median apophysis in the ventral sector of the tegular
ring. The median apophysis is rather short, flat, rounded at the tip and
shaped like a spatula. Only its tip is slightly sclerotized.

April 30, 1990 Petra Sierwald 25

After passing the median apophysis, the sperm duct enters a large
membraneous sac (= tegulum sensu Carico, 1976: figs. 6, 7). The sperm
duct is very slender in this section, thus forming the ejaculatory duct. It
follows the outer margin of the sac. In the unexpanded bulb, this sac is
coiled. The number of loops is species-typical (Fig. 15; Carico, 1976: figs.
8-17). In the left, unexpanded palp (ventral view), the tegular sac is twisted
ascending-counterclockwise (Fig. 38). Distally, the sac is closed by a small,
strongly sclerotized tubercle (Fig. 15). Additionally, the tip bears a
sclerotized rounded projection. The long embolus arises here. Its proximal
part shows clearly that it consists of a peripheral sclerite on the convex
side, the truncus, and a membraneous part on the concave side, the pars
pendula. The remaining part of the embolus is long, whip-like and forms
three loops. It is coiled ascending-clockwise (left unexpanded palp in ventral
view; Figs. 15, 38).

Thaumasia Perty, 1833 (Figs. 16-21, 41)

In the genus Thaumasia, the tibial apophyses of different species are
large and complex. They appear to be species-typical (Fig. 20: Thaumasia
cf. uncata, and Fig. 21: Th. cf. velox). A median hematodocha was not
observed during inflation of the bulb. [This may be due to incomplete
inflation, since old preserved material was used. The basal hematodocha
also did not expand well.]

The tegulum (Fig. 41) is rather similar in structure and position to
the one in Tizmus. Immediately upon the entry of the sperm duct into the
tegulum on the prolateral side, the tegular wall bears a low broad hump
(Fig. 41). This hump is not visible in the unexpanded bulb, since it lies
behind the large distal tegular projection. In Thaumasia cf. velox this
hump is scoop-shaped and its form is similar to the proximal tegular
apophysis (ptp) in Tznus in the same position, hence considered homolo-
gous. The tegulum bears three conspicuous processes that are visible in
the unexpanded bulb (Figs. 16, 17). At the retrolateral side, there is a
long, slender apophysis that is slightly sclerotized. Its tip is swollen,
sclerotized, and tilted. Because of its position and its type of connection
to the tegular ring, it is considered to be the conductor. The conductors
in Tinus and Thaumasia are similarly large, but in Tinus strongly sclerotized
and of different shape. The median apophysis is short, broad, flat and
similar to the one found in Tinus. In Thaumasia however, it is bifurcated
(in cf. uncata and cf. velox). The third process on the retrolateral side of
the bulb is long, shaped like a duckbill and sclerotized. The sperm duct
extends to its tip and undergoes a large loop within it. On its dorsal side,
the sperm duct narrows and runs down to enter the embolic division.

No. 35

NEMOURIA

26

April 30, 1990 Petra Sierwald 27

From its position and the trajectory of the sperm duct this process is clearly
the distal tegular process, homologous to that found in Dolomedes.

The apical division consists mainly of the embolus. The embolus arises
at the basal part of the distal tegular projection and branches off it in a
nearly right angle. The embolus is connected to the distal tegular projection
by a membrane. Within this membrane lie two small sclerites (Fig. 18;
sc). Both are connected to the basal section of the embolus, the retrolateral
sclerite is attached via a membraneous bridge, and the prolateral sclerite
is directly attached to the base of the embolus. The proximal section of
the embolus (Figs. 16, 17) shows clearly the truncus at the convex side
and the pars pendula at the concave side. The base of the truncus is
enlarged; its base fits into a notch in the proximal tegular projection (ptp).

Architis Simon, 1898, and Staberius Simon, 1898 (Figs. 22-26, 39, 42)

The morphology of the genital bulbs in both genera is very similar.
Whereas Architis currently contains seven species, Staberius is monotypic
(Carico, 1981). Therefore, synapomorphies for Architis (excluding Staberius)
cannot be defined at this point. Characters listed here for Staberius may
be autapomorphies for a single species and do not justify a separate genus.

In Architis, the tibial apophyses are complex and species-typical. In
Staberius spinipes, the tibia bears a large hammer-shaped ventral apophysis
in addition to the retrolateral apophysis (see Carico, 1981: figs. 34, 35).
The ventral tibial apophysis is connected to the tibia via a flexible
membrane. The retrolateral apophysis in Staberius is a conspicuous process
pointing laterally.

In the unexpanded bulb (Fig. 22; Carico, 1981: figs. 12-23, 34) the
tegulum and its distal projection are the prominent structures. The tegular
ring is rather small; its distal projection is large. There are two small
apophyses on the tegulum, barely visible in the unexpanded bulb (Figs.
22, 39, 42). The conductor, juxtaposed to the distal tegular projection, is
a small hump-shaped tubercle. The median apophysis is short and
membraneous. Only its tip is slightly sclerotized. The sperm duct does
not progress straight through the tegular ring, but performs two curves.
The sperm duct extends to the tip of the distal tegular projection and
performs a loop. At that point, it starts to narrow becoming the ejaculatory
duct (Figs. 25, 26). The ejaculatory duct enters a large, less sclerotized
hump that is mesally attached to the distal tegular projection. The

Figures 11-15. Left palp of Tinus peregrinus. 11. Unexpanded bulb, ventral view.
12. Unexpanded bulb, prolateral view. 13. Expanded bulb, retrolateral view. 14.
Expanded bulb, prolateral view. 15. Apical division. Scale bars: Figs. 11-14, 1
mm; Fig. 15, 0.5 mm.

28 NEMOURIA No. 35

Figures 16-21. Left palp of Thaumasia. 16. T. cf. velox, unexpanded, ventral view.
17. T. cf. uncata, unexpanded, ventral view. 18. T. cf. uncata, genital bulb, partly
expanded, prolateral view, sc = sclerites. 19. T. cf. uncata, expanded, retrolateral
view. Figures 20, 21. Tibial apophyses. 20. T. cf. uncata. 21. T. cf. velox. Scale
bars: Figs. 16, 17, 19-21, 1 mm; Fig. 18, 0.5 mm.

April 30, 1990 Petra Sierwald 29

sclerotized embolus branches off dorsally from this hump (Figs. 39, 42).
The base of the embolus is broad, consisting of the truncus and pars
pendula.

Pisaurina Simon, 1898 (Figs. 27-29, 44)

In this genus, the tibial apophysis is small. At the distal tip of the
tibia in Pisaurina mira, there is a broad, flat, sclerotized structure. It can
be considered a ventral tibial apophysis (Fig. 27, vta). The petiolus is very
broad (Fig. 28). The tegulum is plate-like (Fig. 44) and lies flat in the
alveolus, thus its position is very different from all other American
Pisauridae. The tubular outer rim of the tegulum containing the sperm
duct is strongly sclerotized, the upper-surface is sclerotized to a lesser
degree, but not membraneous. On its retrolateral side the tegulum produces
a hump with a spur (Figs. 27, 44; dtp). The sperm duct extends to the
distal tip of this hump and performs a loop turning back.

The apical division of Pisaurina consists of three sclerotized bulbous
sections (ad I, II, III). The prolateral side of these three sections is
membraneous; the membrane visible at the proximal end of the genital
bulb (Fig. 27, bmt) is part of it. At the distal end of the second bulbous
section the sperm duct narrows to become the ejaculatory duct. Attached
to the distal region of the third bulbous section is a membrane, regarded
here as the pars pendula of the embolus. The pars pendula is clearly
separated by a sclerotized section from the bmt-membrane. The embolus
is extremely long, thin and whip-like. Over much of its course, it appears
to consist of the ejaculatory duct alone. The broader, basal part of the
truncus appears to be fused to the third bulbous section of the apical
division.

There are two conspicuous apophyses, one of which can be identified
as the median apophysis. The latter possesses a bifurcated sclerotized tip
(similar to Thaumasia) and a membraneous base. It is attached to the
membraneous central area of the plate-like tegulum. Its tip rests in the
spur of the distal tegular hump.

The second apophysis, labelled here sail-shaped apophysis (ssa; labelled
conductor by Carico, 1972), is large, flat and sail-shaped, ending in a long
spur. Its base is membraneous, with a sclerotized strap attached to the
second bulbous section of the apical division. Its outer margin is sclerotized

and lamellar. It is wrapped around the thin embolus. In the undisturbed
bulb, the embolus extends to the tip of the sail-shaped apophysis.
- Apparently, this feature caused its misidentification as a conductor. But
_heither its position nor its mode of attachment to the tegulum and to the

second bulbous section of the apical divisions warrants this homology-

30 NEMOURIA No. 35

i

rs b
PEEP,

SRA TD

So

ue)

Figures 22, 23. Architis tenuis. 22. Unexpanded right bulb, ventral view. 23.
Expanded left palp. Figures 24-26. Staberius spinipes. 24. Expanded left palp,
retrolateral view. 25. Tegulum, prolateral view. 26. Distal tegular projection and
apical division, retrolateral-dorsal view. Scale bars: Figs. 22, 25, 26, 0.5 mm; Figs.
23, 24, 1 mm.

April 30, 1990 Petra Sierwald 31

Figures 27-29. Pisaurina mira, left palp. 27. Unexpanded, ventral view. 28.
Expanded, retrolateral view. 29. Expanded, prolateral view. Figure 30. Pisaura
mirabilis, left expanded palp, ventral view. Scale bars: 1 mm.

hypothesis. The tip of the sail-shaped apophysis lies parallel to the upper
branch of the tip of the median apophysis.

(4) Palp Morphology in Other Nursery-Web Pisauridae

Pisaura Simon, 1885 (Figs. 7, 30, 45)

The tibial apophysis is thick and tubular, bearing brushes of hair in
some species. The upper-surface of the tegulum is mostly sclerotized, thus
the tegulum is bowl-like, with the sperm duct following the outer margins
of the bowl. The upper-surface of the tegulum surrounding the distal

SZ NEMOURIA No. 35

tegular projection is membraneous, strongly inflatable, and continues to
become the basal membraneous tube (Fig. 30). The most prominent
features of the Pisaura bulb are three conspicuous apophyses (Fig. 7; c,
ma, da). The conductor at the retrolateral side of the genital bulb is large
and sclerotized. The next apophysis is smaller, with a hook-shaped tip.
Because of its position and membraneous attachment to the tegulum, it is
considered to be the median apophysis.

The distal apophysis, situated distad of the median apophysis (Figs.
7, 45) is large, consisting of a dorsal branch and a ventral branch (Fig.
45, db, vb). The dorsal and ventral branch are fused distally and form a
distinct hook. The ventral branch anchors the distal apophysis in the
tegulum with a membraneous connection. The dorsal branch is attached
to the membraneous upper-surface of the tegulum and extends towards
the distal sclerotized tube of the apical division. From the ventral branch
a sclerotized clasp, also attached to the membraneous upper-surface of the
tegulum around the distal tegular projection, extends to the basal
membraneous tube of the apical division. Therefore, the distal apophysis
is connected to both the middle and apical division of the genital bulb.

The apical division (Fig. 30) consists of a rather large basal
membraneous tube, the distal, sclerotized tube and a spiral embolus with
a large, flag-like pars pendula. The truncus of the embolus is fused to the
distal end of the sclerotized tube. Within the wall of the pars pendula lies
a thin, T-shaped sclerite.

Blandin (1976) described the genus Afropisaura for African species
formerly assigned to Pisaura. He based his decision on differences in
features of male and female copulatory organs (Blandin, 1976: 928, figs.
17, 18). The species remaining in Pisaura comprise four to six species
besides mirabilis, occurring in the palearctic region (see Brignoli, 1984).
The genus Pisaura is in need of revision.

Approximately 20, mainly African genera possess the same basic bulb
configuration as Pisaura (Fig. 45): a bowl-like tegulum, a large conductor,
a slender median apophysis with a pointed, curved tip, and a large distal
apophysis comprised of two branches. In certain genera, such as Charminus
Thorell, 1899 (Figs. 49-51), the conductor is morphologically more
complex. It consists of a sclerotized convex retrolateral part. This
sclerotized part is connected directly to the tegular ring. The inner, concave
side is an inflatable membrane, and actually a continuation of the
upper-surface of the tegulum. During inflation, this inner part of the
conductor, and also the basal membraneous sections of the median
apophysis are expanded.

The apical divisions of the genera of the Pisaura genus-group show a
wide range of diversity and are genus-specific.

April 30, 1990 Petra Sierwald 33

Thalassius Simon, 1885 (Figs. 8-10, 40, 43)

The tibial apophysis is reduced (see Sierwald, 1987). The tegulum
(Figs. 40, 43) forms a tubular ring with a membraneous center. The
conductor is membraneous and very similar to the one found in Dolomedes.
The median apophysis is large; its tip is heavily sclerotized and species-
typical (Sierwald, 1987, figs. 54-55, 58-61, 126, 127).

In the rubromaculatus-group, the distal tegular projection is large; it
possesses a somewhat less sclerotized zone (Fig. 8, arrowhead; Fig. 40,
meandering line). The apical division consists of a single, solid sclerite,
whose shape is very similar to the distal sclerotized tube and fulcrum
combined as it occurs in Dolomedes tenebrosus (Fig. 3). The apical division
is connected to the distal tegular projection via a short basal membraneous
tube.

In the spinosissimus-group, the distal tegular projection appears to be
reduced. The apical division consists of a basal bulbous section, a
conspicuous, sclerotized finger-shaped apophysis (Fig. 43, fu), and a long,
whip-like embolus. The finger-shaped apophysis is attached to the
sclerotized bulbous section via a membrane and is considered here a
homologue of the fulcrum.

(5) Palp Morphology in the South American Trechalea_ genus-
group

For many of the genera in this group, the copulatory organs have
never been figured in sufficient detail. For the present study, the palps of
Trechalea (22 nominal species) and Paradossenus (three nominal species)
are analyzed. Additionally, I conducted preliminary studies on palps of
Dyrines striatipes, Enna sp., Hesydrus sp. and Syntrechalea porschi. The
results reveal features justifying the separation of these spiders from the
remaining nursery-web Pisauridae.

Trechalea Thorell, 1870 (Figs. 31-33, 46)

The tibial apophysis consists of two branches (Fig. 31; Carico &
Minch, 1981, figs. 1, 2). The body wall between both branches is
membraneous. The tegulum forms a ring. The ventral section of the tegular
ring is broad; the sperm duct performs seven loops within this section of
the tegulum. The conductor is a broad, low hump and only visible in the

expanded palp. The median apophysis is a massive structure with two stout
branches. Its shape and structure is considerably different from the median
apophyses occurring in nursery-web Pisauridae, thus representing another
| character state. The apical division consists of a short thin embolus and a
large terminal apophysis, bearing three small finger-shaped projections.

No. 35

NEMOURIA

34

April 30, 1990 Petra Sierwald 35

Paradossenus F. O. Pickard-Cambridge, 1903 (Figs. 34-36, 47)

The tibial apophysis consists of a spur pointing ventrally. The body
wall beneath it forms a deep membraneous pit. The tibial apophysis of P.
taczanowskii Caporiacco, 1948, agrees with the figure given by F. O.
Pickard-Cambridge (1903: figs. 7-9) in the original description of P.
nigricans. These species are here regarded to be synonymous. The tegulum
is ring-like, the ventral section is well developed as in Trechalea. The sperm
duct performs five loops in the ventral section of the tegulum. The
conductor is a small, inconspicuous projection of the tegular wall. As in
Trechalea, the median apophysis forms a massive cubic structure with two
stout branches. The distal tegular projection is well developed (Fig. 35).
The apical division consists of a short basal membraneous tube and a
slender sclerotized tube, which is fused to the embolus. There are no
terminal apophyses (Fig. 47).

The palps of Dyrines, Enna, Hesydrus and Syntrechalea possess several
identical characters with Trechalea and Paradossenus: a small, reduced
conductor, the massive median apophysis with two branches, a coiled sperm
duct with several switchbacks (except perhaps Dyrines), and a conspicuous
membraneous pit at the distal end of the tibia.

DISCUSSION

(1) Homology of Palpal Elements Within Pisauridae and Evaluation
of Character States

Tibial Apophysis. — The Trechalea genus-group possesses a synapo-
morphic character in the form of a membraneous pit that accompanies the
tibial apophysis. A membraneous ‘pit is also present in Rhoicinus Simon,
1898 (see Exline, 1960: fig. 2), whose relationship is uncertain, but has
been placed in the Pisauridae by various authors.

Distal Tegular Projection. — The distal tegular projection can be
identified by the loop of the sperm duct within it, its distal position and
the attachment site of the apical division being below the tip of the
projection. It occurs clearly in Dolomedes, Pisaura, Architis/Staberius,
Thaumasia, the Thalassius rubromaculatus-group, and Paradossenus. In
Pisaurina, the retrolaterally situated hump with the spur is considered here

Figures 31-33. Trechalea sp., left palp. 31. Unexpanded, ventral view. 32.
Expanded, retrolateral view. 33. Expanded, prolateral view. Figures 34-36.
Paradossenus nigricans, left palp. 34. Unexpanded, ventral view. 35. Expanded,
retrolateral view. 36. Expanded, prolateral view. Scale bars: 1 mm.

36

NEMOURIA

No. 35

April 30, 1990 Petra Sierwald 37

to be homologous with the distal tegular projection of the other genera
because it contains the loop of the sperm duct and because its position
relative to the attachment of the apical division. The genital bulbs of Tinus
and the Thalassius spinosissimus species-group probably lost this feature
secondarily and independently. The distal tegular projection is absent in
the Trechalea genus-group.

Median Apophysis. — All genera studied here possess a median
apophysis. The median apophysis occurs in a variety of different character
states, its general structure being always autapomorphic at the generic
level. Additionally, it can be species-typical (Thalassius), or species-group
specific (Dolomedes). The character state of the median apophysis in
Architis/Staberius is considered a reduction, therefore derived. In the
Trechalea genus-group, the morphology of the median apophysis differs
distinctly from the median apophysis of the nursery-web Pisauridae, thus
this character state is a synapomorphy for the South American Trechalea
genus-group.

Conductor. — The conductor occurs in several morphological condi-
tions (character states): it is membraneous in Dolomedes and Thalassius
(synapomorphy); it is long in Thaumasia, and large, stiff, and heavily
sclerotized in Tzmus. In the Pisaura genus-group it is large as well, but in
several genera the concave side of the conductor consists of an inflatable
membrane of variable size, which is a continuation of the upper-surface
of the tegulum. The conductor is small in Architis/Staberius and the
Trechalea genus-group, and absent in Pisaurina. These reductions are
independently derived.

Apical Division. — The apical division of the genital bulb shows a
wide range of diversity: its appearance is usually genus-specific. The basal
membraneous tube occurs in all pisaurid genera studied so far: the sclerites
of the apical division were always found to be connected to the distal
tegular process via an inflatable membrane. The basal membraneous tube

Figures 37-47. Schematic drawings of tegula, trajectory of sperm duct black,

arrows indicate insertion of conductor (c), sail-shaped apophysis (ssa) and median

-apophysis (unmarked arrow). 37. Dolomedes tenebrosus, conductor and median
_apophysis omitted. 38. Tinus peregrinus, median apophysis omitted, embolus cut
off. 39. Staberius spinipes. 40. Thalassitus rubromaculatus, conductor and median
_apophysis omitted. 41. Thaumasia cf. uncata, median apophysis omitted, embolus
cut short. 42. Architis tenuis. 43. Thalassius spinosissimus, conductor and median
_apophysis omitted. 44. Pisaurina mira, median apophysis and sail-shaped apophysis
omitted. 45. Pisaura mirabilis, tegulum including conductor, median apophysis and
distal apophysis. 46. Trechalea sp., median apophysis omitted. 47. Paradossenus

-migricans, median apophysis omitted.

38 NEMOURIA No. 35

Figure 48. Trochosa ruricola, left unexpanded palp, ventral view. Figures 49-51.
Charminus camerunensis. 49. Left unexpanded palp, ventral view. 50. Expanded left
palp, retrolateral view. 51. Unexpanded left palp, retrolateral view. Scale bars: 1
mm.

April 30, 1990 Petra Sierwald 39

varies in size. It is very small in Thalasstus. The membraneous sac in Tinus
is interpreted as a complexly developed basal membraneous tube of the
apical division, based on the fact that the sperm duct forms the ejaculatory
duct within it.

The distal sclerotized tube is clearly present in Dolomedes, Tinus, and
the Pisaura genus-group; the Thalassius rubromaculatus-group possesses a
homologue that is fused to the distal tegular projection and to the fulcrum.
The presence of the distal sclerotized tube is considered the plesiomorphic
condition. In Pisaurina, it may be represented by the second bulbous
section of its apical division (see below). In Thaumasia, several small
sclerites are present at the base of the embolus; they may represent
homologues of the sclerotized tube and/or terminal apophysis. In Architis
and Staberius, the basal membraneous tube and the distal sclerotized tube
appear to be fused to the broad hump that is attached to the distal tegular
projection. In Paradossenus, the distal sclerotized tube is reduced in size
and fused to the base of the embolus. The distal sclerotized tube is absent
in Trechalea.

Apophyses of the Apical Division. — The genera in question show a
variety of terminal apophyses. The fulcrum is defined as an apophysis
attached to the distal end of the distal sclerotized tube. The fulcrum in
both Thalassius species-groups is considered homologous to the Dolomedes
fulcrum because of its position and similarity in shape. The club-shaped
terminal apophysis in 7imus (Fig. 15) is considered here to represent a
homologue of the fulcrum as well. The subterminal lateral apophysis in
Dolomedes is an autapomorphic feature of this genus.

Distal Apophysis and Sail-Shaped Apophysis. — The distal apophysis
in the Pisaura genus-group may be homologous to the sail-shaped apophysis
in Pisaurina. Both are attached to the tegulum in the membraneous area
surrounding the distal tegular projection. Their position and connections
to the middle and apical divisions support this hypothesis. Since the
sail-shaped apophysis in Pisaurina is connected (via the sclerotized strap)
to the second bulbous section of the apical division, the second bulbous
section could be considered a homologue of the distal sclerotized tube.
This homology-hypothesis is speculative. It could be supported or refuted
by the presence or absence of further concordant homologies in the female
copulatory organs or somatic characters. Pisaurina displays a sexual

behavior (using silk to wrap the female’s legs [Bruce & Carico, 19887])
that is different from the copulatory behavior in Pisaura mirabilis.

Blandin (1976: 923, figs. 8, 9, 13, 14) labelled the distal apophysis in
Pisaura as the fulcrum. Since the term fulcrum is used for an apophysis

attached to the distal end of the sclerotized tube of the apical division, the
fulcrum sensu Comstock is not homologous to the distal apophysis in

40 NEMOURIA No. 35

Pisaura.

Trajectory of Sperm Duct. — In most genera studied, the sperm
duct forms a simple loop, which presumably represents the plesiomorphic
condition. The Trechalea genus-group is characterized by several switch-
backs of the duct in the well-developed ventral section of the tegulum.

The Spiral Embolus. — The spiral embolus, consisting of a truncus
and a pars pendula as in Thaumasia, Architis/Staberius, and the Pisaura
genus-group, occurs in many other spider families and is perhaps traceable
in Liphistius [Fig. 1; see below under (3)]. In several pisaurid genera, the
pars pendula is reduced and the truncus appears to be very thin, especially
distally, thus producing a whip-like embolus (as in Thalassius spinosissimus,
Pisaurina and the Trechalea genus-group).

(2) Systematic Implications

A definitive cladistic analysis of the approximately 55 pisaurid genera
based on genitalic characters would be premature. Too many pisaurid
genera, especially from Asia, are not known in sufficient detail and analyses
of the female copulatory organs in both the Pisaura and the Trechalea
genus-group are lacking. But the results obtained to date suggest the
following phylogenetic conclusions:

The Pisaura genus-group includes at least: Afropisaura Blandin, 1976;
Caripetella Strand, 1926; Charminus Thorell, 1899; Czspius Simon, 1898;
Cladycnis Simon, 1898; Euprosthenops Pocock, 1897; Euprosthenopsis Blan-
din, 1974; Maypacius Simon, 1898; Perenethis L. Koch, 1878; Pisaura;
Pisaurellus Roewer 1961; Tetragonophthalma Karsch, 1878; Thalassiopsis
Roewer, 1955; Vuattouxia Blandin, 1979 (1979a); Walrencea Blandin, 1979
(1979a); and presumably also: Chiasmopes Pavesi, 1883; Dendrolycosa
Doleschall, 1859; Paracladycnis Blandin, 1979 (1979a); Phalaeops Roewer,
1955; Ransonia Blandin, 1979 (1979a); Rothus Simon, 1898; Tallonia
Simon, 1889. The group is characterized by the distal apophysis, thus it
appears to form a well-supported monophyletic clade. Features in the
female copulatory organs (Sierwald, in prep.) support this hypothesis
through more synapomorphies. It would be important to study the
copulatory behavior of other genera of the Pisaura genus-group. The
copulatory behavior of Pisaura mirabilis is complex: the male presents a
prey item to the female (Gerhardt & Kastner, 1938: 543). If similar

behavioral features are found in other members of the Pzsaura-group, they |

would provide additional synapomorphies.

The Trechalea genus-group contains presumably the nominal genera —
Demolodes Mello-Leitao, 1943; Dossenus Simon, 1898; Dyrines Simon, 1903; —
Dyrinoides Badcock, 1932; Enna; Hesydrus; Paradossenus; Sisenna Simon, —
1898; Syntrechalea; Trechalea; and Xinguisella Mello-Leitao, 1940. The ©

April 30, 1990 Petra Sierwald 4]

group is well supported as a clade through the synapomorphies found in
the male palp, the structure of the egg sac, and through the parental
behavior. Preliminary studies on the female copulatory organs (Sierwald,
unpubl.) provide further evidence. In general, this group is poorly known.
Rhoicinus and its relative Barrisca Chamberlin and Ivie, 1936 (see Platnick,
1979), might belong to the Trechalea genus-group as well. The systematic
position of the Trechalea genus-group is unknown at this point. The
assignment of the genera of this group to the family Pisauridae was based
on plesiomorphic characters like the eye pattern, and similarities in the
general habitus and habitats (occurrence near fresh water). Thus, the
Trechalea genus-group is certainly not the sister group of the Pisaura
genus-group, but the latter is more closely related to the remaining
nursery-web pisaurid genera.

The remaining American nursery-web Pisauridae do not represent a
monophyletic group; the affinities and relationships of single members are
not known at this point. Architis and Staberius surely are monophyletic.
In case more Staberius-species are found (Staberius is currently monotypic)
synapomorphies will have to be discovered to warrant the existence of two
separate genera (see Platnick, 1976). These then would represent sister taxa.
Otherwise, the two nominal genera Architis and Staberius would be
synonyms and the nomenclatural priority will need to be established, since
both genera were described by Simon in 1898. Architis is the only member
of the American nursery-web Pisauridae that hunts on snares (Nentwig,
1985). Webs for prey capture occur in several pisaurid genera around the
world, e.g., in the African Euprosthenops (see Gerhardt & Kastner, 1938:
619), in the Asian Eurychoera and Polyboea (see Koh, 1989: 97, 100) and
others, and also in juveniles, e.g., in Pisaurina (see Carico, 1985) and Pisaura
(see Lenler-Eriksen, 1969). Web types and occurrences in pisaurid genera
are not been studied in sufficient detail to allow phylogenetic conclusions.
But Carico (1985) pointed out that the juvenile webs of Pisaurina and
Pisaura, and the web of Architis are strikingly similar, which could indicate
relationship.

Thalassius and Dolomedes share a greater number of characters with
each other than with any other pisaurid. Their genital bulbs, spine-pattern
on the legs (Sierwald, 1987), and the hunting behavior of most of their
species on the surface of the water provide additional evidence. Dolomedes
and Thalassius are considered close relatives. Tinus and Thaumasia share
the proximal tegular projection and a large conductor, but no other features

in the bulb are synapomorphic. The female copulatory organs of both
genera display similarities (Sierwald, 1989b: figs. 11-12, 13-16; see also

Carico, 1976: figs. 18-31): in both genera the copulatory duct is long and
at least in some species, complexly coiled. Additional similarities are:

42 NEMOURIA No. 35

habitus and color pattern (white longitudinal stripes along the lateral sides
of the body), as well as the hunting on the water surface in several species
of Tinus (Carico, 1976; Sierwald, pers. observ. in Belize, Central America)
and Thaumasia (Sierwald, pers. observ. on Barro Colorado Island, Panama).
These features could be evidence for a closer relationship of both genera.

The genus Pisaurina is isolated. If its sail-shaped apophysis is a
homologue of the distal apophysis in the Pisaura genus-group, it may be
a the sister taxon of the latter.

The present study shows that monophyletic units (Pisaura genus-
group, 7 rechalea genus-group) can be found using homologous morphologi-
cal elements of the male copulatory organs. These hypotheses can be tested
by finding corroborating features in female copulatory organs, somatic
characters and other evidence such as behavioral data (e.g., structure of
the egg sac in the Trechalea genus-group, mating behavior in the Pisaura
genus-group, hunting behavior in Thalassius and Dolomedes). A study of
the copulatory organs in potential sister taxa of Pisauridae either among
the Amaurobioidea sensu Forster & Wilton (1973) or Lycosoidea sensu
Homann (1971) will provide data to test homology-concepts of palpal
elements within a more inclusive taxon, i.e., at the superfamilial level.

(3) Homology of Pisaurid Palpal Elements with Those of Other
Families

Hypotheses on the homology of pisaurid palpal elements with those
in other families rely nearly exclusively on data furnished by other authors,
either in the form of descriptions or drawings. In the course of this study,
it became evident that unequivocal interpretations of drawings and
descriptions can not always be achieved: actual attachment points of
membranes (or hematodochae) and of apophyses, especially at the tegulum
in relation to the course of the sperm duct, are not always presented
clearly. Likewise, the actual limits of sclerites can not always be determined
with certainty from drawings. Therefore, the following homology concepts
have to be considered with caution. But certain striking morphological
similarities among unrelated groups should be mentioned.

Retrolateral Tibial Apophysis. — Apophyses at the distal end of the
tibia occur in many spider families®. Their shape and structure are often
species-typical and have been figured in many taxonomic studies. The
function of tibial apophyses is uncertain and not necessarily the same in
all groups. One observation in Dolomedes tenebrosus revealed that its
retrolateral apophysis arrests the expanded bulb during copula (Sierwald —
& Coddington, 1988). If the retrolateral tibial apophyses in the different —
groups are homologous, they represent a synapomorphy for a large taxon |
within the Araneoclada sensu Platnick (1977; including Araneomorphae |

April 30, 1990 Petra Sierwald 43

except Hypochiloidea and Austrochiloidea). Lehtinen (1967: 285, fig. 6)
suggested such a taxon including his Amaurobiides and Zodariides, but
his analysis regarding morphology and exact position of tibial apophyses
in the male palp (e.g., table 8, page 310; table 29, page 351) is too vague
to draw conclusions on homology/non-homology. Griswold [in press]
distinguished between apophyses at different positions on the male palpal
tibia, and also reached the conclusion that the retrolateral tibial apophyses
may be a synapomorphy for a large taxon within the Araneoclada. A
retrolateral tibial apophysis is present in Liphistius (Mesothelae) (see Platnick
& Sedgwick, 1984: 3, fig. 7), but its homology with the retrolateral
apophysis within Araneoclada is doubtful.

Basal and Middle Divisions of the Genital Bulb. — Subtegulum, basal
hematodocha (basal division), and tegulum (middle division) are considered
homologous among all spiders. The male genital bulb of Liphistius already
contains these basic elements (Kraus, 1978; Haupt, 1983). Furthermore,
subtegulum and tegulum occur in most entelegyne spider groups. Lack of
tegulum and subtegulum as distinct sclerites (in certain Mygalomorphae
and ‘“‘haplogyne” spiders) is currently interpreted as secondary loss (either
by fusion of formerly distinct sclerites or by paedomorphosis during
ontogeny, see Coddington [in press]). The ontogenetic evidence supports
this conclusion.

Subtegulum. — The pisaurid type of subtegulum with a sclerotized
distal ridge and anelli occurs in other families as well (Agelenidae
[Comstock, 1910: fig. 24; Gering, 1953: figs. 27, 31; see Bennett, 1987, for
a detailed description of the subtegulum in Agelenidae], Thomisidae
[Loerbrocks, 1984: fig. 9], and Phyxelidinae [Griswold, in press]). The
morphological structure of the subtegulum often does not receive detailed
attention in monographic studies. Details on its morphological diversity
are therefore lacking.

Distal Tegular Projection. — Enlargements and elaborate structure
of the distal tegular area occur in several spider groups independently. In
Linyphiidae, the suprategulum corresponds in its position to the distal
tegular projection as it is found in Pisauridae. The linyphiid suprategulum
| does not contain a loop of the sperm duct, thus the complex structure and
enlargement of the distal tegular area in both families is obviously
homoplasous. Griswold [in press] described an embolic sclerite (EBS) in
some genera of Phyxelidinae that also corresponds to the distal tegular area.
| Median Apophysis. — Haupt (1983: 276, figs. la-e) presented
hypotheses that (a) the median apophysis is a homologue in all spiders,
and (b) it can be traced back to the Kontrategulum in Liphistius and
Heptathela. The Kontrategulum sensu Haupt is a ring-segment broken out
of the ring-like tegulum in Liphistius. Haupt did not present any further

Aa, NEMOURIA INO: 35

details on the Kontrategulum = median apophyses hypothesis (e.g., by
comparing it in various families, giving morphological criteria or describing
different character states of the Kontrategulum in other spiders). He only
presented the araneid median apophysis as the representative for the
“median apophysis” in all entelegyne spiders. Raven (1985: 15) argued
convincingly that the segmentation of the tegular ring in liphistiids appears
to be an autapomorphy of subunits within the group or of certain Liphistius
species. Haupt’s claim that the median apophysis of all other spiders is a
derivative of the Kontrategulum of Liphistius is therefore unsubstantiated.

Nevertheless, Haupt’s concept of the median apophysis as an old
structure, and therefore homologous in a large clade within Araneoclada,
appears to be valid. The median apophysis as it occurs in Pisauridae - a
distinct sclerite, at a certain position on the tegulum, connected to the
tegulum via an inflatable or flexible membrane - occurs in many and
unrelated families (e.g., in Amaurobiidae [Griswold, in press]; in Araneidae
[Coddington, in press], in Anyphaenidae [Sierwald, 1988: fig. 1]; in
Lycosidae, Fig. 48). Lehtinen (1967: 285, fig. 6, and 289, table 1) placed
the “‘origin of the median apophysis with movable joint at the base”’ as a
synapomorphy for his Amaurobiides, thereby excluding the Araneoidea
from possessing a homologue. He did not comment on the striking
similarity between the araneid median apophysis with the median apophysis
in Amaurobiides.

Conductor. — As mentioned before, the term conductor has been
used for tegular as well as for apical apophyses. In the Linyphiidae, the
term conductor was used for a structure of the apical division. Because of
doubtful homology, Saaristo (1971: 465) recommended avoiding the term
conductor for Linyphiidae. Another example for inconsistent use of the
term conductor appears in the family Lycosidae. Whereas in most lycosid
genera a tegular process is labelled conductor (e.g., Trochosa ruricola, Fig.
48), the membraneous “conductor” in Pardosa (see Dondale & Redner,
1984, figs. 5-7) is an apophysis of the apical division and clearly not
homologous.

Haupt (1983) considered the conductor homologous in all entelegyne
spiders, but did not discuss morphological and positional differences of
conductors in the various groups, and thus gave no evidence for his claim.
He homologized the mygalomorph conductor with the Kontrategulum.
Raven (1985: 16, and fig. 11) presented convincing evidence that the
mygalomorph conductor (as in Atypus) may be a homologue of the stiff,
prominent ridges of the distal margin of the tegulum in the mesothelid
Liphistius (see Fig. 1), since ‘‘the mygalomorph conductor is a sclerotized
process arising opposite the embolus, separated from it by an extensive
haematodocha....””> Such tegular ridges occur in several Liphistius- and

April 30, 1990 Petra Sierwald 45

Heptathela-species (Kraus, 1978: figs. 2-4; Platnick & Sedgwick, 1984: figs.
39, 41, 66; Haupt, 1983: figs. 4c, 7).

Lehtinen (1967: 285, fig. 6) proposed “‘a membraneous, secondary
conductor”’ exclusively for Lycosoidea and Amaurobioidea sensu Lehtinen
(including Miturgidae, Amaurobiidae, Liocranidae, Agelenidae, Dictynidae
and Hahniidae). As discussed earlier, his argumentation lacks convincing
morphological details to distinguish between a primary and a secondary
conductor.

The pisaurid conductor as an immovable, tegular outgrowth is
considered here homologous in many spider groups (e.g., Lycosidae,
Anyphaenidae, Amaurobiidae, Psechridae, Araneidae). It is suggested that
the basic pattern of the genital bulb in Araneoclada contains two tegular
apophyses (median apophysis and conductor), as Coddington [in press]
presumed as well. Coddington [in press] considered the conductor to be a
homologue in Dictynoidea, Orbiculariae, and Amaurobioidea sensu Forster
& Wilton.

Conductor and Median Apophysis. — Independent modifications,
reductions or losses of one or both may have frequently occurred in various
groups. If only one tegular apophysis is present, it may be difficult to
identify which (the median apophysis or the conductor) has been lost.
Likewise, if both are present in two not closely-related spider groups, it
may be difficult or impossible to decide whether the conductor in one
group is homologous to the median apophysis in the other group or vice
versa. Slight changes in the growth pattern of the tegulum could result in
rather different insertion points of the tegular apophyses in relation to the
trajectory of the sperm duct in the mature genital bulb.

Apical Division. — In many Mygalomorphae and “‘haplogyne”’ spiders
the apical division consists of the embolus, forming a mere continuation
of the middle division rather than being separated from it (connate type
of embolus sensu Comstock, 1910: 182). The mygalomorph conductor, often
assigned to the apical division, appears to be a tegular apophysis, as Raven
(1985) showed (see above). In entelegyne spiders, the apical division is often
a complex structure.

Basal Membraneous Tube of Apical Division. — The large sclerites
of the genital bulb are usually connected via less sclerotized, membraneous
tubes, which are called hematodochae, when they are large and inflatable.
The column, as described by Saaristo (1971) for Linyphiidae (called duct
membrane by Merrett, 1963), connecting the tegulum and the apical
division corresponds in position to the basal membraneous tube as found
in Pisauridae. The character state as an inflatable, rather large membraneous
tube is supposedly independently derived.

Distal Sclerotized Tube of Apical Division. — In Araneidae, the

46 NEMOURIA No. 35

apical division consists of two tubes with the ejaculatory duct running
through them (Grasshoff, 1968: figs. 33a, b). The proximal tube, called
the radix, is fully sclerotized. The second, distal tube is partly sclerotized
and partly membraneous (see below, Stipes). A radix has been described
for other spider families (e.g., Uloboridae, Oecobiidae, Theridiidae) but
Coddington [in press] considered none of these homologues to the araneid
radix based on major morphological differences and arguments involving
parsimony. He argues that the true radix is most likely a synapomorphy
for Araneidae and Linyphiidae. The distal sclerotized tube of the apical
division in Pisauridae appears to be similar to the araneid radix,
morphologically and positionally. A similar sclerite was described by Gering
(1953: fig. 6) in Agelenopsis. The possible homology of these sclerotized
tubes of the apical division deserves further attention.

Embolus. — A detailed study on the morphology of emboli in spiders
is lacking. Whether emboli contain one or several sclerites, the shape of
such sclerites, and the extent of membraneous sections are often missing
in morphological descriptions. As mentioned earlier, only Comstock (1910)
described different morphological types of emboli. The opening of the
sperm duct always identifies the embolus itself unambiguously. But the
opening of the sperm duct only identifies the terminal end of the embolus,
not the proximal end of the structure.

A survey of 53 araneomorph spider families showed that spine-like
emboli are most common. Comstock called these spiral emboli, consisting
of a gutter-like sclerite, the truncus, and a membraneous pars pendula.
Together they form a tube. The typical spiral embolus has a broad base
and a tapering terminal section (e.g., in Architis, Fig. 42). The length of
the terminal section varies. Long terminal sections may perform a loop as
Thaumasia (Figs. 16, 17). Such loops of the terminal section occur also
independently in e.g., Linyphiidae (Lasiargus, Whiele, 1960: fig. 21a),
Theridiidae (Achaearanea, Levi, 1961: fig. 7), and Stiphidiidae (Stzphidion,
Davies, 1988: fig. 23). Very long emboli often form multispiral coils as in
Tinus (Fig. 11). Such coiled emboli are broadened and tape-like. This type
occurs independently in e.g., Deinopidae (Deiopis, Kraus, 1956: fig. 4),
Symphytognathidae (Patu, Forster & Platnick, 1977: fig. 53), Linyphiidae
(Labulla, Merrett, 1963: fig. 33), Mysmenidae (Mysmenopsis, Gertsch, 1960:
fig. 45, 46), Pacullidae (Sabahya, Deeleman-Reinhold, 1980: fig. 10), and
Theridiidae (Latrodectus, Wiehle, 1960: fig. 11).

In addition to a sclerite, emboli often contain membraneous parts,
especially in the basal region (e.g., Figs. 18, 23). Gering (1953: 26, figs.
29, 30; Agelenidae), Loerbrocks (1983: figs. 4, 6; Thomisidae), and
Griswold (in press; Amaurobiidae) labelled membraneous parts of emboli
pars pendula. Comstock (1910: fig. 6) used the term in a theraphosid

April 30, 1990 Petra Sierwald 47

embolus, obviously implying homology. Often, membraneous sections in
the base of the embolus are indicated in drawings, but not confirmed in
descriptions (e.g., Senoculidae [Senoculus, Petrunkevitch, 1925: fig. 105],
Linyphiidae [Cornicularia, Wiehle, 1960: fig. 10], Lycosidae [Schizocosa,
Dondale & Redner, 1978: fig. 10; Sosippus, Brady: 1962, fig. 36]).

It is my current hypothesis that all spine-like emboli, containing the
truncus and a pars pendula, are in fact homologous. The question is at
what level are they homologous?

Since the genital bulb of Mesothelae already contains the two large
sclerites, subtegulum and tegulum, the structure of its apical division is
of special importance. In Liphistius (Fig. 1), the apical division consists of
a large sclerite (sclerite III sensu Kraus, 1978) bearing one elongated
sclerotized projection and two lamellae (bl = broad lamella and hl = hyaloid
lamella; after Kraus, 1978). The sclerotized projection is U-shaped in cross
section and forms three sides of the embolus. The fourth, open side is
closed by the hyaloid lamella, thus forming a tube-like embolus’. Haupt
(1979: 360) described the embolus of Ryuthela nishihirai (Haupt, 1979)!°
as “‘ strongly sclerotized except for a longitudinal groove... [citation
translated],’’ presumably describing a homologue of the hyaloid lamella
sensu Kraus of Liphistius.

The resemblance of the liphistiid embolus to the “spiral type of
embolus” as described above is striking. This morphological similarity
could imply homology of at least the projection of the sclerite III (sensu
Kraus) with the truncus of the spiral embolus in entelegyne spiders.

Since all emboli in spiders seem to contain a sclerite or form a
sclerotized tube, I propose that it is most parsimonious to assume that the
embolic sclerite in all spiders is a homologue of either the complete sclerite
IiI or the projection of sclerite III in Liphistius. This hypothesis is in
concordance with the homology of subtegulum and tegulum in all spiders.
The pars pendula could be a homologue of the hyaloid lamella, but I am
less confident of such a hypothesis. Supporting evidence for the latter
hypothesis could come from ontogenetic studies of genital bulbs, or from
the occurrence of membraneous sections in the emboli of Paleo- and/or
Neocribellate spiders. But even then, homology may be difficult to
hypothesize since a membraneous section is not complex in structure.
Therefore, morphological similarity may not offer supportive evidence for
homology. If the pars pendula in the emboli of entelegyne spiders is not
homologous with the hyaloid lamella in Liphistius, the spiral embolus may
be a synapomorphy for a large group within the Araneoclada sensu Platnick.
A re-examination of the embolic area of spider groups with other types
of emboli (e.g., coniform and lamelliform sensu Comstock) may reveal
unexpected morphological similarities and may lead to new homology

48 NEMOURIA No. 35

concepts as the following indicates.

Stipes in Araneidae. — In addition to the proximal sclerotized tube
called the radix, the apical division of some araneids contains a second,
distal tube. The latter tube carries none to several terminal apophyses and
an embolus. Part of the wall of the second tube is sclerotized, forming a
sclerite called the stipes. The remaining part of the tube wall is
membraneous and inflatable (called distal hematodocha by Grasshoff, 1968:
fig. 33), carrying terminal apophyses and a short embolus. This embolus
was figured by Comstock (1910: fig. 25) and called the coniform embolus.
It is possible that the stipes-section of the araneid bulb is an enormously
developed spiral embolus, with the stipes representing the truncus and the
membraneous part representing the pars pendula, thus the distal hematodo-
cha would be the homologue of the pars pendula. If this is true, the
traditional araneid embolus corresponds only to a part of the embolus in
Pisauridae and other groups with spiral emboli.

Haupt (1983) and Heimer (1986) both discussed the striking similari-
ties in morphology of the genital bulbs in Mesothelae and entelegyne
spiders. Because of the “presumed great phylogenetic distance between
Araneoidea and Liphistiidae...,” Heimer (1986: 119) explicitly avoided the
term homologous but attributed the morphological similarity to ho-
moiology. This term, meaning analogous structures in homologous organs,
is not part of the current set of terms used in phylogenetic systematics.
Although divergent definitions exist!!, the term basically means parallelism.
Heimer’s statement makes it difficult to recognize on what level he
considers parts of the genital bulb homologous within Araneae.

As shown above in the discussion on the conductor, median apophysis,
and spiral embolus, I believe that careful morphological analysis and
comparison will enable us to establish homologies of palpal structures
within Araneae at all taxonomic levels. This knowledge of homologous
elements in the male palpal organ, as well as in the female copulatory
organ, will greatly advance the reconstruction of the phylogeny of Araneae.

SUMMARY

Morphology and Homology of Palpal Elements

It is now generally accepted that the tripartite genital bulb in male
spiders is plesiomorphic (genital bulb structure in Mesothelae, ontogenetic
evidence; see Platnick & Gertsch, 1976, Kraus, 1978, and Haupt, 1983).
The large sclerites (subtegulum and tegulum) are therefore homologous
in all spiders.

It is proposed here that morphological similarity and position of
tegular apophyses and processes often provide sufficient evidence for

April 30, 1990 Petra Sierwald 49

homology concepts (process connected via membrane to tegulum or mere
outgrowth of tegulum; process containing a loop of the sperm duct or
lacking such). The basic pattern in Araneoclada sensu Platnick (1977)
contains two tegular apophyses (median apophysis and conductor). A
conductor found in the apical division (as described in e.g., Linyphiidae)
is not a homologue of the true conductor.

It is suggested here that the spiral embolus sensu Comstock (1910) is
a plesiomorphic feature and homologous for a large clade. The sclerite of
the spiral embolus, the truncus, is considered to be a homologue of the
projection or the entire sclerite III in the apical division of Liphistzus.
Furthermore, it is suggested that other types of emboli, e.g., the coniform
embolus of Araneidae, may only be partly homologous to the spiral
embolus, but that the araneid stipes may represent a homologue of the
truncus. The araneid radix of the apical division is compared to the distal
sclerotized tube as it occurs in the apical division of Pisauridae and possible
homology is discussed.

The retrolateral tibial apophysis as it occurs in Pisauridae is homolo-
gous wherever it is found in other spider families and therefore represents
a synapomorphy for a large clade within Araneoclada.

Palp Morphology of American Nursery-Web Pisauridae

Architis/Staberius, Pisaurina, Thaumasia, Tinus, and Dolomedes consti-
tute the American nursery-web Pisauridae. Only Dolomedes is distributed
worldwide, the remaining genera are restricted to the Americas. All
American nursery-web Pisauridae possess a well-developed and species-
typical retrolateral tibial apophysis. The subtegulum bears a sclerotized
ridge and anelli. The tegulum is ring-like (bowl-like in Pisaurina). The
distal end of the tegulum (distal tegular projection) is enlarged and contains
a loop of the sperm duct.

Dolomedes possesses a membraneous conductor, the median apophysis
is flat, lamelliform, and species-group specific. Thaumasia and Tinus have
a large conductor, heavily sclerotized in Tinus; the genus-typical median
apophyses are short with a slightly sclerotized distal region. The conductor
is independently reduced in Pisaurina and Architis/Staberius. The median
apophysis in the latter is very similar to the one found in Tinus and
Thaumasia. Pisaurina possesses a well-developed, species-typical median
apophysis and an additional, tegular, sail-shaped apophysis that is presum-
ably a homologue of the distal apophysis as found in the Pisaura
genus-group. Zimus and Thaumasia both possess a sclerotized, proximal,
tegular process that is large in Tinus and short in Thaumasia.

The apical division of Pisauridae consists of a basal membraneous
tube connected to the distal end of the tegulum, a sclerotized distal tube

50 NEMOURIA No. 35

and a spiral embolus sensu Comstock (1910). Dolomedes carries a terminal
apophysis called the fulcrum. A homologue of the fulcrum is found in
Tinus in form of a small club-shaped apophysis. In Tznus, the basal
membraneous tube of the apical division is large, forming a species-typical,
coiled, inflatable sac.

The bulb structure of the African genus Thalassius is very similar to
Dolomedes, indicating closer relationship.

Palp Morphology of the Pisaura Genus-Group

The tegulum is bowl-like. The conductor is sclerotized and well
developed. The conspicuous, distal tegular apophysis in addition to the
median apophysis, and structure of the apical division are synapomorphies
of the group. Preliminary studies of the female copulatory organ corroborate
monophyly of this clade.

Palp Morphology of the Trechalea Genus-Group

The Trechalea genus-group possesses the following synapomorphies
in the genital bulb: a small, reduced conductor, a median apophysis with
two branches, a sperm duct with several switchbacks, a well-developed
ventral section of the tegulum and a conspicuous membraneous pit at the
distal end of the palpal tibia. Preliminary studies of the female copulatory
organs, the structure of the egg sac and maternal behavior (attaching the
egg sac to the spinnerets) corroborate the monophyly of this clade.

Systematics of the Family Pisauridae

So far, no morphological synapomorphies have been found to unite
the 55 genera assigned to the family Pisauridae. An assumed apomorphy,
the nursery-web, built by the female for the newly hatched spiderlings has
been observed to date in nine genera belonging to all three subfamilies of
Pisauridae. Thus, to define monophyletic units among pisaurid genera,
morphological synapomorphies have to be found. The sister group to
Pisauridae is unknown, but should be among the Lycosoidea sensu Homann
(1971), since this superfamily is united by the synapomorphy of a
grate-shaped tapetum in the secondary eyes. Lehtinen’s (1967) separation
of Pisauridae sensu lato in Dolomedidae and Pisauridae sensu stricto, placed
in two different superfamilies, is rejected at this point. The main distinction
between the two, the primary conductor in Pisauridae sensu stricto and the
secondary conductor in Dolomedidae appears to be unsubstantiated.

Based on the structure of the genital bulb, the Prsaura genus-group
contains at least: Afropisaura Blandin, 1976; Caripetella Strand, 1926;
Charminus Thorell, 1899; Czspzus Simon, 1898; Cladycnis Simon, 1898;
Euprosthenops Pocock, 1897; Euprosthenopsis Blandin, 1974; Maypacius
Simon, 1898; Perenethis L. Koch, 1878; Pisaura Simon, 1885; Pisaurellus

April 30, 1990 Petra Sierwald 51

Roewer, 1961; Tetragonophthalma Karsch, 1878; Thalasstopsis Roewer,
1955; Vuattouxia Blandin, 1979 (1979a); Walrencea Blandin, 1979 (1979a);
and perhaps also: Chiasmopes Pavesi, 1883; Dendrolycosa Doleschall, 1859;
Paracladycnis Blandin, 1979 (1979a); Phalaeops, Roewer, 1955; Ransonia
Blandin, 1979 (1979a); Rothus Simon, 1898; Tallonia Simon, 1889.

The remaining American nursery-web Pisauridae do not represent a
monophyletic clade. Tinus F. O. Pickard-Cambridge, 1901, and Thaumasia
Perty, 1833, are closely related to each other and that clade is related to
Dolomedes Latreille, 1804, and Thalasstus Simon, 1885. The affinities of
Architis Simon, 1898, and Staberius Simon, 1898, are unknown. The genus
Pisaurina Simon, 1898, is presumably most closely related to the Prsaura
genus-group.

The Trechalea genus-group contains presumably the nominal genera
Demolodes Mello-Leitao, 1943; Dossenus Simon, 1898; Dyrines Simon, 1903;
Dyrinoides Badcock, 1932; Enna O. Pickard-Cambridge, 1897; Hesydrus
Simon, 1898; Paradossenus F. O. Pickard-Cambridge, 1903; Sisenna Simon,
1898; Syntrechalea F. O. Pickard-Cambridge, 1902; Trechalea Thorell,
1870; and Xinguisella Mello-Leitao, 1940. Based on features in the
copulatory organs of both sexes, the structure of the egg sac, and parental
care, the Trechalea genus-group is not a close relative of the nursery-web
Pisauridae.

ACKNOWLEDGMENTS

The present study was supported in part by a postdoctoral fellowship of the
Smithsonian Institution, Washington, DC and in part by Research Grant Si 371/1-1
of the Deutsche Forschungsgemeinschaft (Federal Republic of Germany). This
publication was aided substantially by a grant from the Johanna und Fritz
Buch-Gedachtnisstiftung (Hamburg, Germany). Parts of this work were done while
in residence at the Harbor Branch Oceanographic Institution (HBOJ) in Ft. Pierce,
Florida. Use of office space and library is gratefully acknowledged. Dr. Kevin
Eckelbarger (HBOI) kindly supported the study through loan of equipment.
Observations on live Trechalea specimens were made during a visit at the Research
Station on Barro Colorado Island (Smithsonian Tropical Research Institute,
Panama) in August of 1983.

Dr. Jonathan Coddington acted as principal adviser during the Smithsonian
postdoctoral fellowship; his encouragement, help and many favors are gratefully
acknowledged. Dr. James Carico (Lynchburg College) provided advice, material
and helpful discussions. The following colleagues provided specimen loans: Dr. J.
A. Coddington (USNM), Dr. J. Gruber (NHMW), Dr. J. Hertault (MNHN),
Dr. H. W. Levi (MCZ), Ms. S. Mascherini (MZS), Dr. N. I. Platnick (AMNH),
and Ms. D. Silva (MHNSM). Thanks are due to Drs. C. Griswold and J.
Coddington for reading earlier drafts of the manuscript and helpful criticism. I
thank all reviewers for their suggestions and improvements on the manuscript.

52 NEMOURIA No. 35

LITERATURE CITED

Ax, P. 1984. Das phylogenetische System. Gustav Fischer Verlag, Stuttgart, New
York, 349 pp.

Barnes, R. D. 1980. Invertebrate Zoology. Saunders College/Holt, Rinehart and
Winston, Philadelphia, 1089 pp.

Barrows, W. M. 1925. Modification and development of the arachnid palpal claw
with special reference to spiders. Annals of the Entomological Society of
America, 18: 483-525.

Baum, S. 1972. Zum “‘Cribellaten-Problem”’: Die Genitalstrukturen der Oecobiinae
und Urocteinae (Arachn.: Aran.: Oecobiidae). Abhandlungen und Verhand-
lungen des naturwissenschaftlichen Vereins in Hamburg, (NF)16: 101-153.

Bennett, R. G. 1987. Systematics and natural history of Wadotes (Araneae,
Agelenidae). The Journal of Arachnology, 15: 91-128.

Berkum, F. H. van. 1982. Natural history of a tropical, shrimp-eating spider
(Pisauridae). The Journal of Arachnology, 10: 117-121.

Bertkau, P. 1875. Ueber den Generationsapparat der Araneiden. Archiv fiir
Naturgeschichte, 41(1): 235-262, pl. VII.

Bhatnagar, R. D. S. & J. G. Rempel. 1962. The structure, function, and
postembryonic development of the male and female copulatory organs of the
black widow spider Latrodectus curacaviensis (Miiller). Canadian Journal of
Zoology, 40: 465-510.

Blackwall, J. 1843. On the palpi of spiders. British Association for the Advancement
of Science. Report, 12, Notices: 66-68.

Blandin, P. 1976. Etudes sur les Pisauridae africaines. VI. Définition des genres
Pisaura Simon, 1885, Pisaurellus Roewer, 1961, Afropisaura n. gen. et mise
au point sur les espéces des genres Afropisaura et Pisaurellus (Araneae
Pisauridae Pisaurinae). Revue de Zoologie Africaines, 90(4): 917-939.

— 1979a. Etudes sur les Pisauridae africaines. XI. Genres peu connus ou
nouveaux des iles Canaries, du continent africain et de Madagscar (Araneae,
Pisauridae). Revue de Zoologie Africaines, 93(2): 347-375.

— 1979b. Cycle biologique et production de l’araignee Afropisaura valida (Simon,
1885) (Araneae-Pissauridae) dans une savane d’Afrique occidentale (Lamto,
Cote d’Ivoire). Tropical Ecology, 20(1): 78-93.

Blauvelt, H. H. 1936. The comparative morphology of the secondary sexual organs

of Linyphia and some related genera, including a revision of the group.
Festschrift Strand, 2: 81-171.

Brady, A. R. 1962. The spider genus Sosippus in North America, Mexico and —
Central America (Araneae, Lycosidae). Psyche, 69(3): 129-164.

— 1964. The lynx spiders of North America, north of Mexico (Araneae: —
Oxyopidae). Bulletin of the Museum of Comparative Zoology, 131(13):
429-518. |

April 30, 1990 Petra Sierwald 53

Brignoli, P. M. 1984. Zur Problematik der mediterranen Pisaura-Arten (Arachnida,
Araneae, Pisauridae). Zoologischer Anzeiger, 213(!/2): 33-43.

Bruce, J. A. & J. E. Carico. 1988. Silk use during mating in Pisaurina mira
(Walckenaer) (Araneae, Pisauridae). The Journal of Arachnology, 16(1): 1-4.

Caporiacco, L. di. 1948. Arachnida of British Guiana collected in 1931 and 1936
by Professors Beccari and Romiti. Proceedings of the Zoological Society of
London, 118: 607-747.

Carico, J. E. 1972. The nearctic spider genus Pisaurina (Pisauridae). Psyche, 79(4):
295-310.

— 1973. The nearctic species of the genus Dolomedes (Araneae: Pisauridae).
Bulletin of the Museum of Comparative Zoology, 144(7): 435-488.

—— 1976. The spider genus Timus (Pisauridae). Psyche, 83(1): 63-78.

— 1981. The neotropical spider genera Architis and Staberius (Pisauridae).
Bulletin of the American Museum of Natural History, 170(1): 140-153.

—— 1985. Description and significance of the juvenile web of Pisaurina mira
(Walck.) (Araneae: Pisauridae). Bulletin of the British Arachnological Society,
6(7): 295-296.

— 1986. Trechaleidae: A “‘new” American spider family [Abstract], p. 305. In:
Eberhard, W. G., Y. D. Lubin & B. C. Robinson (eds.), Proceedings of the
Ninth International Congress of Arachnology. Smithsonian Institution Press,
Washington, DC, London, 333 pp.

Carico, J. E., J. Adis & N. D. Penny. 1985. A new species of Trechalea (Pisauridae:
Araneae) from Central Amazonian inundation forests and notes on its natural
history and ecology. Bulletin of the British Arachnological Society, 6(7):
289-294.

Carico, J. E. & E. W. Minch. 1981. A new species of Trechalea (Pisauridae) from
North America. Bulletin of the American Museum of Natural History, 170(1):
154-156.

Chamberlin, R. V. 1904. Notes on generic characters in the Lycosidae. The
Canadian Entomologist, 36(6): 173-178.

Coddington, J. A. [in press]. A review of ontogeny and homology in the male
palpus of orb-weaving spiders and their potential outgroups. Smithsonian
Contributions to Zoology.

Comstock, J. H. 1910. The palpi of male spiders. Annals of the Entomological
Society of America, 3: 161-185.

Cooke, J. A. L. 1966. Synopsis of the structure and function of the genitalia in
Dysdera crocata (Araneae, Dysderidae). Senckenbergiana biologica, 47(1):
35-43,

Davies, V. T. 1988. Three new species of the spider genus Stiphidion (Araneae:
Amaurobioidea: Stiphidiidae) from Australia. Memoirs of the Queensland
Museum, 25(2): 265-271.

54 NEMOURIA IN@: 35

Davies, V. T. & R. J. Raven. 1980. Megadolomedes, nov. gen. (Araneae: Pisauridae)
with a description of the male of the type species, Dolomedes australianus
Koch 1865. Memoirs of the Queensland Museum, 20(1): 135-141.

Deeleman-Reinhold, C. L. 1980. Contributions to the knowledge of the southeast
Asian spiders of the families Pacullidae and Tetrablemmidae. Zoologische
Mededelingen, 56(5): 65-82.

Dondale, C. D. 1986. The subfamilies of wolf spiders (Araneae: Lycosidae), in:
Barrientos, J. A. (ed.), Actas X Congreso Internacional de Aracnologia:
327-332.

Dondale, C. D. & J. H. Redner. 1978. Revision of the nearctic wolf spider genus
Schizocosa (Araneida: Lycosidae). Canadian Entomology, 110: 143-181.

— 1984. Revision of the milvina group of the wolf spider genus Pardosa (Araneae:
Lycosidae). Psyche, 91(1-2): 67-117.

Dzwillo, M. 1978. Prinzipien der Evolution. B. G. Teubner, Stuttgart, 152 pp.

Eberhard, W. G. 1985. Sexual selection and animal genitalia. Harvard University
Press, Cambridge, 244 pp.

Exline, H. 1960. Rhoicinine spiders (Pisauridae) of western South America.
Proceedings of the California Academy of Sciences, 29(17): 577-620.

Foelix, R. F. 1987. Biology of spiders. Harvard University Press, Cambridge, 306
Pp.
Forster, R. R. & N. I. Platnick. 1977. A review of the spider family Sym-

phytognathidae (Arachnida, Araneae). American Museum Novitates, 2619:
1-29.

Forster, R. R. & C. L. Wilton. 1973. The spiders of New Zealand. Part IV.
Agelenidae, Stiphidiidae, Amphinectidae, Amaurobiidae, Neolanidae, Ctenidae,
Psechridae. Otago Museum Bulletin, 4: 1-309.

Gassmann, F. 1925. Die Entwicklung des mdnnlichen Spinnentasters, dargestellt
an Lepthyphantes nebulosus Sund. Zeitschrift fiir Morphologie und Okologie
der Tiere, 5: 98-118.

Gerhardt, U. & A. Kastner. 1938. Araneae. Jn: Kiikenthal, W. & T. Krumbach:
Handbuch der Zoologie, 3(2): 394-656.

Gering, R. L. 1953. Structure and function of the genitalia in some American
agelenid spiders. Smithsonian Miscellaneous Collections, 121(4): 1-84.

Gertsch, W. J. 1960. Descriptions of American spiders of the family Sym-
phytognathidae. American Museum Novitates, 1981: 1-40.

Grasshoff, M. 1968. Morphologische Kriterien als Ausdruck von Artgrenzen bei
Radnetzspinnen der Subfamilie Araneinae (Arachnida: Araneae: Araneidae).
Abhandlungen der senckenbergischen naturforschenden Gesellschaft, 516:
1-100.

April 30, 1990 Petra Sierwald 55

Griswold, C. E. [in press]. A revision and phylogenetic analysis of the spider
subfamily Phyxelidinae (Araneae, Amaurobiidae). Bulletin of the American
Museum of Natural History.

Harm, M. 1931. Beitrage zur Kenntnis des Baues, der Funktion und der
Entwicklung des akzessorischen Kopulationsorgans von Segestria bavarica
C. L. Koch. Zeitschrift fiir Morphologie und Okologie der Tiere, 22: 629-670.

— 1934. Bau, Funktion und Entwicklung des akzessorischen Kopulationsorgans
von Evarcha marcgravi Scop. Zeitschrift fiir wissenschaftliche Zoologie, 146:
123-134.

Hasselt, A. W. M. van. 1889. Le muscle spiral et la vésicule du palpe des araignées
males. Tijdschrift voor Entomologie, 32: 161-203, pls. 6, 7.

Haupt, J. 1979. Lebensweise und Sexualverhalten der mesothelen Spinne Heptathela
nishihirai n. sp. (Araneae, Liphistiidae). Zoologischer Anzeiger, 202(5-6):
348-374.

— 1983. Vergleichende Morphologie der Genitalorgane und Phylogenie der
liphistiomorphen Webspinnen. I. Revision der bekannten Arten. Zeitschrift
fiir zoologische Systematik und Evolutionsforschung, 21(4): 275-293.

Heimer, S. 1982. Interne Arretierungsmechanismen an den Kopulationsorganen
médnnlicher Spinnen (Arachnida, Araneae). Entomologische Abhandlungen,
45(3): 35-64.

—— 1986. From where are the Linyphiidae derived? Problems of Araneoidea
phylogeny (Arachnida: Araneae), pp. 117-120. In: Eberhard, W. G., Y. D.
Lubin & B. C. Robinson (eds.), Proceedings of the Ninth International
Congress of Arachnology. Smithsonian Institution Press, Washington, DC,
London, 333 pp.

Homann, H. 1971. Die Augen der Araneae. Zeitschrift fiir Morphologie der Tiere,
69: 201-272.

[ICZN] International Commission on Zoological Nomenclature. 1985. International
code of zoological nomenclature. Third edition. International Trust for
Zoological Nomenclature, etc., London, xx + 338 pp.

Koh, J. K. H. 1989. A guide to common Singapore spiders. Singapore Science
Centre, Singapore, 160 pp., numerous color photographs.

Kraus, O. 1956. Eine neue Deinopidae aus El Salvador (Arch., Araneae).
Senckenbergiana biologica, 37(!/2): 167-170.

— 1978. Liphistius and the evolution of spider genitalia. Symposium of the
Zoological Society of London, 42: 235-254.

—— 1984. Male spider genitalia: evolutionary changes in structure and function.
Verhandlungen des naturwissenschaftlichen Vereins in Hamburg, (NF)27:
373-382.

Lamoral, B. H. 1973. On the morphology, anatomy, histology and function of the
tarsal organ on the pedipalpi of Palystes castaneus (Sparassidae, Araneida).
Annals of the Natal Museum, 21(3): 609-648.

56 NEMOURIA No. 35

Lehtinen, P. T. 1967. Classification of the cribellate spiders and some allied families,
with notes on the evolution of the suborder Araneomorpha. Annales Zoologici
Fennici, 4(3): 199-468.

Lenler-Eriksen, P. 1969. The hunting-web of the young Pisaura mirabilis. Journal
of Zoology, London, 157: 391-398.

Levi, H. W. 1961. Evolutionary trends in the development of palpal sclerites in
the family Theridiidae. Journal of Morphology, 108: 1-9.

— 1982. Arthropoda. Chelicerata. Jn: Parker, S. P. (ed.), Synopsis and
classification of living organisms, 2: 71-95. McGraw Hill Book Company.

Loerbrocks, A. 1983. Revision der Krabbenspinnen-Gattung Heriaeus Simon
(Arachnida: Araneae: Thomisidae). Verhandlungen des naturwissenschaftli-
chen Vereins in Hamburg, (NF)26: 85-139.

— 1984. Mechanik der Kopulationsorgane von Misumena vatia (Clerck, 1757)
(Arachnida: Araneae: Thomisidae). Verhandlungen des naturwissenschaftli-
chen Vereins in Hamburg, (NF)27: 383-403.

Melchers, M. 1963. Zur Biologie und zum Verhalten von Cupiennius salei
(Keyserling), einer amerikanischen Ctenide. Zoologische Jahrbiicher, Sys-
tematik, 91: 1-90.

Menge, A. 1843. Ueber die Lebensweise der Arachniden. Neueste Schriften der
naturforschenden Gesellschaft in Danzig, 4(3): 1-64, 2 pls.

— 1866-1879. Preussische Spinnen. 11 Abtheilungen. Schriften der Naturforsch-
enden Gesellschaft in Danzig, 560 pp., pls. I-XCI.

Merrett, P. 1963. The palpus of male spiders of the family Linyphiidae. Proceedings
of the Zoological Society of London, 140(3): 347-467.

— 1988. Notes on the biology of the neotropical pisaurid, Ancylometes bogotensis
(Keyserling) (Araneae: Pisauridae). Bulletin of the British Arachnological
Society, 7(7): 197-201.

Millidge, A. F. 1977. The conformation of the male palpal organs of linyphiid
spiders and its application to the taxonomic and phylogenetic analysis of the
family (Araneae: Linyphiidae). Bulletin of the British Arachnological Society,
4(1): 1-60.

— 1980. The erigonine spiders of North America. Part 1. Introduction and
taxonomic background (Araneae: Linyphiidae). The Journal of Arachnology,
8(2): 97-107.

Nentwig, W. 1985. Architis nitidopilosa, a neotropical pisaurid with a permanent —
catching web (Araneae, Pisauridae). Bulletin of the British Arachnological —
Society, 6(7): 297-303. |

Osterloh, A. 1922. Beitrége zur Kenntnis des Kopulationsapparates einiger Spinnen. |
Zeitschrift fiir wissenschaftliche Zoologie, 119: 326-421.

April 30, 1990 Petra Sierwald By)

Patterson, C. 1982. Morphological characters and homology. Pp. 21-74. In: Joysey,
K. A. & A. E. Friday, (eds.), Problems of phylogenetic reconstruction.
Academic Press, New York, 442 pp.

Petrunkevitch, A. 1925. Arachnida from Panama. Transactions of the Connecticut
Academy of Arts and Sciences, 27: 51-248.

Pickard-Cambridge, F. O. 1903. On some new species of spiders belonging to the
families Pisauridae and Senoculidae; with characters of a new genus.
Proceedings of the Zoological Society of London, 1903(1): 151-168.

Platnick, N. I. 1975. A revision of palpimanid spiders of the new subfamily
Otiothopinae (Araneae, Palpimanidae). American Museum Novitates, 2562:
1-32.

— 1976. Are monotypic genera possible? Systematic Zoology, 25(2): 198-199.

— 1977. The hypochiloid spiders: a cladistic analysis, with notes on the Atypoidea
(Arachnida, Araneae). American Museum Novitates, 2627: 1-23.

— 1979. A revision of the spider genus Barrisca (Araneae, Rhoicininae). The
Journal of Arachnology, 6: 213-217.

Platnick, N. I. & W. J. Gertsch. 1976. The suborders of spiders: a cladistic analysis.
American Museum Novitates, 2607: 1-15.

Platnick, N. I. & W. C. Sedgwick. 1984. A revision of the spider genus Liphistius
(Araneae, Mesothelae). American Museum Novitates, 2781: 1-31.

Raven, R. J. 1985. The spider infraorder Mygalomorphae (Araneae): Cladistics and
systematics. Bulletin of the American Museum of Natural History, 182(1):
1-180.

Remane, A. 1952. Die Grundlagen des natiirlichen Systems der vergleichenden
Anatomie und Phylogenetik. Geest und Portig KG, Leipzig, 400 pp.

Riedl, R. 1975. Die Ordnung des Lebendigen. Paul Parey Verlag, Hamburg, Berlin,
372 pp.

Roewer, F. 1955. Lycosaeformia I. Exploration du Parc National de l’Upemba,
30: 1-420.

Saaristo, M. I. 1971. Revision of the genus Maro O. Pickard-Cambridge (Araneae,
Linyphiidae). Annales Zoologici Fennici, 8: 463-482.

Sadana, G. L. 1971. Studies on the postembryonic development of the palpal organ
of Lycosa chaperi Simon (Lycosidae: Araneida). Zoologischer Anzeiger, 186:
251-258.

Schult, J. 1983a. Taster haplogyner Spinnen unter phylogenetischem Aspekt
(Arachnida: Araneae). Verhandlungen des naturwissenschaftlichen Vereins
in Hamburg, (NF)26: 69-84.

—— 1983b. Simple bulbs in male spiders - primitive or derived? (Arachnida:
| Araneae). Verhandlungen des naturwissenschaftlichen Vereins in Hamburg,
(NF)26: 155-160.

58 NEMOURIA No. 35

Schult, J. & U. Sellenschlo. 1983. Morphologie und Funktion der Genitalstrukturen
bei Nephila (Arach., Aran., Araneidae). Mitteilungen des hamburgischen
zoologischen Museums und Instituts, 80: 221-230.

Sierwald, P. 1987. Revision der Gattung Thalassius (Arachnida, Araneae, Pisauri-
dae). Verhandlungen des naturwissenschaftlichen Vereins in Hamburg,
(NF)29: 51-142.

— 1988. Spiders of Bermuda. Nemouria, 31: 1-24.

— 1989a. Notes on the behavior of Thalassius spinosisstmus (Arachnida, Araneae,
Pisauridae). Psyche, 95(3-4): 243-252.

— 1989b. Morphology and ontogeny of female copulatory organs in American
Pisauridae with special reference to homologous features (Arachnida: Ara-
neae). Smithsonian Contributions to Zoology, 484: 1-24.

Sierwald, P. & J. A. Coddington. 1988. Functional aspects of the male palpal organ
in Dolomedes tenebrosus, with notes of the mating behavior (Araneae,
Pisauridae). The Journal of Arachnology, 16(2): 262-265.

Simon, E. 1890. Etudes arachnologiques. 22e Mémoire (1). XXXIV. Etude sur les
Arachnides de Yemen. -—- Remarques sur la classification des Araignées
[page 79-82]. Annales de la Société entomologique de France, (6)10: 77-124.

— 1898. Histoire naturelle des Araignées, 2(2): 193-380. Paris.

Song, D. & J. Haupt. 1984. Comparative morphology and phylogeny of liphisti-
omorph spiders (Araneae: Mesothelae). 2. Revision of new Chinese heptathe-
lid species. Verhandlungen des naturwissenschaftlichen Vereins in Hamburg,
(NF)27: 443-451.

Szombathy, C. 1915. Uber Bau und Funktion des Bulbus der méannlichen
Kopulationsorgane bei Agelena und Mygale. Annales Musei Nationales
Hungarici, 13: 252-276.

Wagner, W. A. 1886. Entwicklung und Bau der Kopulationsorgane bei den
Araneinen. Izwestia imperatorskago obtchiestwa lioubitelei iestiestwoznania,
antropology i etnografi, sostoiaschago pri imperatorskom Moskowskom
Ouniwersitete, 50: 206-236, 4 pls. [non vidi, cited from Osterloh, 1922].

— 1888. Copulationsorgane des Mannchens als Criterium fiir die Systematik der
Spinnen. Horae Societatis entomologicae Rossicae, 22: 3-132, pl. I-X.

Westring, N. I. 1861. Araneae svecicae. Géteborgs Konglige Vetenskaps- och |
Vitterhets-Samhdlles Handlingar, 7: 1-615. |

Wiehle, H. 1960. Der Embolus des miannlichen Spinnentasters. Verhandlungen |
der deutschen zoologischen Gesellschaft, 1960: 457-480 [= Supplement |
Zoologischer Anzeiger]. |

—— 1967. Meta - eine semientelegyne Gattung der Araneae (Arach.). Senckenber- |
giana biologica, 48(3): 183-196. |

April 30, 1990 Petra Sierwald 59

Wiley, E. O. 1981. Phylogenetics. John Wiley & Sons, New York, Chichester,
Brisbane, Toronto, Singapore, 439 pp.

ENDNOTES

1Members of Sparassidae (e.g., Heteropoda venatoria Linnaeus, 1767; pers. observ.)
carry the flat, discus-shaped egg sac in their chelicerae as well.

2The egg sac of Lycosidae is also attached to the spinnerets. Although the lycosid
egg sac possesses a seam, it consists of two equally shaped and equally constructed
halves, thus is significantly different from the egg sac of the Trechalea genus-group.
Attaching the egg sac to the spinnerets occurs also in the ctenid Cupiennius Simon,
1891 (see Melchers, 1963).

3Processes at the subtegulum are very rare. The only ones known to me occur in

Liphistius langkawi and L. murphyorum (see Platnick & Sedgwick, 1984: figs. 31-35,
38-42).

4Osterloh (1922: 333) distinguished between apophyses that arrest the bulb
(Retinulum) and those which engage in the female epigynum (Retinaculum).

-Exception: Thalassius Simon, 1885, is the only member of Pisauridae with a
reduced tibial apophysis (Sierwald, 1987).

©Roewer (1955) called the median apophysis in Thalassius conductor.

’This behavior occurs also in Ancylometes (see Merrett, 1988) and Thalassius (see
Sierwald, 1989a).

8A superficial count of spider families with such retrolateral tibial apophyses
exceeded over 20 families, including: Acanthoctenidae, Agelenidae, Amaurobiidae,
Ammoxenidae, Anyphaenidae, Clubionidae, Ctenidae, Dictynidae, Gallieniellidae,
Gnaphosidae, Hahniidae, Heteropodidae, Homalonychidae, Oxyopidae, Pisauridae,

Philodromidae, Psechridae, Salticidae, Selenopidae, Senoculidae, Tengellidae,

Thomisidae, Toxopidae, Zodariidae.

°Raven (1985: 15) misidentified the hyaloid lamella at the embolus of Liphistius as

being the mesothelid conductor.

10See Song & Haupt, 1984, for the taxonomic position of R. nishihirai.

Dzwillo, 1978: 96; Riedl, 1975: 60. Ax (1984: 63) considered the terms parallelism,
homoplasy, and homoiology superfluous. Patterson (1982: 45-48) gives an
enlightening analysis of the terms homology, homoiology, homonomy, parallelism,
and convergence.

Reviewers of this paper: Dr. Jonathan Coddington, National Museum of Natural
‘History, Washington, DC; Dr. Robert Raven, Queensland Museum, South
Brisbane, Australia; and Dr. William A. Shear, Hampden-Sydney College,
‘Hampden-Sydney, Virginia.

Founding Editor Managing Editor
JOHN E. DU PONT RUDIGER BIELER

Consulting Editors
WILLIAM K. EMERSON, American Museum of Natural History, New York,
NY.
M. G. HARASEWYCH, National Museum of Natural History, Smithsonian
Institution, Washington, DC.
KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh,
PA.
RICHARD W. THORINGTON, JR., National Museum of Natural History,
Smithsonian Institution, Washington, DC.

Published by the

DELAWARE MUSEUM OF NATURAL HISTORY
4840 Kennett Pike, Wilmington,
Delaware 19807, U.S.A.

a

A
3G

NEMOURIA

Occasional Papers of the Delaware Museum of Natural History

NUMBER 36 JuLy 30, 1991

ANNOTATED CATALOG OF TYPE SPECIMENS IN
THE MALACOLOGICAL COLLECTION OF
THE DELAWARE MUSEUM OF NATURAL HISTORY.

GASTROPODA
(PROSOBRANCHIA AND OPISTHOBRANCHIA)

Rudiger Bieler)? and Alison Bradford!

ABSTRACT. A brief outline of the history and sources of the malacological collection
at the Delaware Museum of Natural History is given. All type material (as defined
by ICZN Article 72) was separated from the main collection. This paper reviews
and lists the type holdings of Recent prosobranch and opisthobranch gastropods.

_ The type status of each specimen was studied by comparison with the original
publications. A total of 191 type lots of 160 species-group taxa from 65 authors
is recognized; 55 of these are holotypes or syntype lots. The majority of
name-bearing types consists of marine caenogastropod prosobranchs. An anno-
tated list of all type lots as well as an author index are given. The list of references
contains all papers in which the taxa are first described and also other major
publications based on the collection.

INTRODUCTION

The malacological collection of the Delaware Museum of Natural
History C(DMNH) with currently about 190,000 catalogued lots is one
of the largest of its kind in the country. In 1982, then containing 135,000
catalogued lots, it was ranked eighth in size among the U.S. collections
in a survey by Thompson (1982: 44).

The marine shell-bearing gastropods are the main strength of the
collection (about 52% of total lots), followed by land and freshwater
gastropods (27%), marine bivalves (16%), freshwater bivalves (4.5%),

'Delaware Museum of Natural History Serial Publication
2Current address: Field Museum of Natural History, ISSN 0085-3887
Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605

Z NEMOURIA No. 36

and miscellaneous other molluscan groups (0.5%). In addition to this
material in dry lots, there is a small alcoholic collection. The dry
collection is housed in steel cabinets and arranged in taxonomic order
by families, largely following the traditional arrangement based on the
Thiele (1929-1935), Wenz (1938-1944) and Zilch (1959-1960) mono-
graphs. It contains some fossil (mainly Tertiary) specimens, interfiled
with the Recent material. There is no geographical specialization. At
present, data for approximately 30% of the lots have been entered into
a micro-computer-based relational database system (dBase IJ]+, commer-
cially available from Ashton-Tate).

The DMNH malacological collection, amassed during only 30 years
(the museum was founded in 1959 and opened to the public in 1972),
came from various sources. The main origin has been donations and
purchases from private collectors and exchanges with other institutions.
The large collection storage areas (13,000 square feet), designed with a
holding capacity of 1.4 million lots, made it possible to absorb orphaned
collections from other institutions as well as extensive material originating
from expeditions and ecological studies. Additional deposited specimens
were collected as part of research projects of DMNH staff and associates,
or were placed here because of the publications associated with the
museum (Indo-Pacific Mollusca, The Nautilus, Delaware Museum of Natural
_ Aistory Monograph Series, Nemouria).

In addition to the large original collection by DMNH’s founder,
John E. du Pont, major private collections were obtained, by donation
or purchase, from the following individuals (3,000—100,000 specimens
each): W. Andrews, C. Aguayo, H. Boswell, M. P. Breithaupt, W. J.
Clench, T. Crow, W. E. Dodd, E. Doremus, L. Grantier, B. K. Harris,
R. Jackson, H. G. Jewell, C. L. Kauffman, G. & M. Kline, N. W.
Lermond, C. A. Long, P. M. Mikkelsen, J. E. Morrison, J. E. Norton,
R. E. M. Ostheimer, W. Over, J. D. Parker, E. Quammen, C. L.
Richardson, F. Schilling, H. L. Sears, M. & H. Schriner, D. Steger, M.
L. Walton, and R. E. Winter. Also, significant collections (1,500—3,000
specimens each) came from: J. C. Alexander, J. & C. Bennett, A.
Chadwick, H. Du Shane, W. M. Heilman, W. M. Ingram, J. W.
Lippincott, C. V. MacCoy, H. & M. Minzak, B. J. Piech, B. Roth, G.
M. Seymour, C. Weaver, and A. Westerfield.

Significant quantities of exchange material and donations were
received from the Academy of Natural Sciences of Philadelphia (ANSP,
Pennsylvania), the National Museum of Natural History (USNM,
Washington, DC), and the Ohio State University Museum (Columbus);
orphaned collections came from University of Kansas (Lawrence), Mystic
Seaport Museum (Mystic, Connecticut), Portland Society of Natural

July 30, 1991 Bieler & Bradford 3

History (Portland, Maine), Stamford Museum (Stamford, Connecticut),
and the Division of Historical and Cultural Affairs, State of Delaware.
The museum holds one permanent loan, the eastern U.S. Unionidae
collection of Stanford University (Stanford, California).

The type material (holo-, para-, syn-, lecto-, paralecto- and neotypes,
as defined by ICZN Article 72) has recently been separated from the
main collection. This allows us to comply with Recommendation 72G(4)
of the International Code of Nomenclature (ICZN, 1985: 72): “Every
institution in which name-bearing types are deposited should. . . publish
lists of name-bearing types in its possession or custody.’’ As can be
expected from the collections of a relatively young museum, the DMNH
type holdings are small when compared to other institutions. The type
material can be sorted into two groups. The first comprises specimens
that were in possession of the museum at the time of the original
description of the respective taxon. This usually is material based on
research by DMNH staff and associates, by outside scientists working
on DMNH material or using DMNH as a repository, or it is material
described in journals currently or in the past published by the museum.
The second group consists of material that reached DMNH subsequent
to the original publication date. For type specimens that have changed
ownership several times a type catalog such as this may be the only
means of locating the current repository.

It might come as a surprise that the DMNH holdings contain type
lots of taxa described by Australian and New Zealand authors, such as
W. O. Cernohorsky, B. C. Cotton, H. J. Finlay, C. Hedley, F. W.
Hutton, T. Iredale, and A. W. B. Powell.

This is not a critical catalog in the sense that the current taxonomic
status for the nominal taxa involved has not been researched in detail.
However, the type status was critically evaluated by comparison with the
original description and in strict application of the ICZN rules. Many
specimens previously recognized as para- and syntypes because they
were part of the original material of the describing author, were excluded
from the type collection when the original description explicitly restricted
the number or repository of types (ICZN Art. 72(b)(vi)). This type
material “grey zone”? most often originated from retained, but not cited,
specimens in the collections of the original collector or author. In
accordance with ICZN Article 73(b), DMNH specimens were recognized
as syntypes in cases when the original author failed to designate a holotype
and we were unable to locate a subsequent lectotype designation.

All recognized or suspected type specimens were measured. Speci-
mens with an overall size larger than 3 mm were measured using calipers,
smaller specimens with the aid of a dissecting microscope and calibrated

4 NEMOURIA No. 36

eye piece (accuracy + 0.1 mm in both cases). ““Height’’ (H) was defined
as the largest dimension parallel to the columellar axis, and ‘“‘width”’
(W) the largest dimension perpendicular to that axis. In cases when these
measurements were meaningless or the shell axis was obscured (e.g.,
Caecidae, Patellidae), other dimensions are given and _ individually
indicated. The new measurements not only provide additional informa-
tion when dimensions were not given in the original publication, but also
serve to evaluate identity of type material. Minor discrepancy between
the original measurements and ours may reflect different techniques
(e.g., measurements without reference to shell axis) or accuracy (e.g.,
original measurement by ruler instead of calipers). Large discrepancies,
however, indicate either error in the original publication or, worse, a
subsequent confusion of type material. In cases where the originally given
dimensions differ from ours, they are noted in parentheses and quotes.
In all other cases, the measurements agree, or there was no such
information in the original description.

Reflecting the strength of the general collection, the majority of the
DMNH type specimens, including most name-bearing types (holo-,
lecto-, neotypes and syntypic series), belong to the shell-bearing Proso-
branchia and Opisthobranchia (191 lots of 160 species-group taxa, all
of them of Recent forms). Of these, 38 are recognized as holotypes, 61
_ aS syntypes in 17 lots, and more than 350 as paratypes or paralectotypes
in 134 lots.

The listings in this catalog are arranged as follows:

(1) new species-group name, (2) original generic (subgeneric) and/or
species-level placement, (3) original status (e.g., “‘[as n. ssp.]’’), (4)
original author and short bibliographic reference, with figure reference
to DMNH specimen(s), (5) number and kind of types, (6) DMNH
collection catalog number, (7) locality data for specimen in type lot, taken
from label and/or original description [discrepancies noted, metric
conversion of depth data where originally missing]; in the case of
paratypes, a note whether this location is the type locality, (8) dimensions
as described above, (9) remarks, (10) family placement.

The ‘Remarks’ section provides a variety of information. In case the
original publication does not indicate DMNH as type repository, the
history of the lot as known to us is given (e.g., “ex T. L. McGinty, via
D. Steger’’). Other data include references to figures in publications
other than the original, and recognized errors in the original publication.

The family placement is given in its originally published form, and
often an updated family placement is added in square brackets (for
prosobranch family-group taxa according to Ponder & Warén, 1988).
The following listing of types is arranged in alphabetical order of species,

July 30, 1991 Bieler & Bradford 5

with Prosobranchia and Opisthobranchia listed separately. In addition,
an index by author as well as complete bibliographic references are
provided.

TYPE CATALOG

CLASS GASTROPODA
SUBCLASS PROSOBRANCHIA

abrolhosensis, Acmaea (Collisella) [as n. sp.]

Petuch, 1979b: 513.

4 paratypes: DMNH 121797; south side of Santa Barbara Island,
Abrolhos Archipelago, Bahia State, Brazil (19°57’S, 38°42’°W);
type locality. Dimensions (greatest lengths): 20.9 [1 shell];
21.4, 19.6, 18.8 [3 complete alcohol-preserved specimens]
(“lengths 21 mm, 21 mm, 20 mm, 19 mm7’’). Patellidae
[Acmaeidae].

adamsianum, Vexillum (Costellaria) [as n. sp.]
Cernohorsky, 1978: 64, fig. 9.
1 paratype: DMNH 118026; Keehi Lagoon, Oahu, Hawaiian
Islands, 50 fathoms [91.5 ml]; type locality. Dimensions: H=
9.0, W= 3.6 (“3.7”). Costellartidae.

ahiparana, Turritella (Zeacolpus) [as n. sp.]
Powell, 1927: 297.
6 paratypes: DMNH 027852; off Ahipara, Ninety Mile Beach, New
Zealand, 23 fathoms [42.1 m]; type locality. Dimensions: (1)
H= 17.4, W= 5.2; (2) H= 17.1, W= 4.7; 3) H= 17.0, W=
5.4; (4) H= 16.9, W= 4.8; (5) H= 16.2, W= 4.9; (6) H=
14.6, W= 4.1. Remarks: Ex J. H. Alexander. Turritellidae.

allyni, Terebra [as n. sp.]
Bratcher & R. D. Burch, 1970: 298.
1 paratype: DMNH 022421; California Academy of Sciences station
237792, east shore of Maria Madre Island, Tres Marias
Group, Baja California, Mexico (21°35’N, 106°26’W), 5-10
fathoms [9.1-18.3 mJ]; type locality. Dimensions: H= 28.4,
We= 7.2. Terebridae.

amosi, Metula [as n. sp.]
Vanatta, 1913: 22.
2 syntypes: DMNH 024523; Panama. Dimensions: (1) H= 35.9,
W= 11.8 [apex broken]; (2) H= 28.9, W= 10.9 [apex broken].

6 NEMOURIA No. 36

Remarks: Ex S. N. Rhoads, via ANSP 316700. Buccinidae.

aphrodite, Conus [as n. sp.|

Petuch, 1979a: 11, figs. 34-35 (holotype).

Holotype: DMNH 126398; off Panglao, Bohol Island, Philippines,
approximately 250 m. Dimensions: H= 20.8 (‘‘21”’), W= 11.3
(Ga Bina

1 paratype: DMNH 126399; type locality. Dimensions: H= 13.9
(“13”), W= 7.3.

Conidae.

arcas, Terebra (Strioterebrum) [as n. sp.]

Abbott, 1954: 40.

1 paratype: DMNH 104984; 35 miles E.N.E. of Arcas Cays, 17
fathoms [31.1 m] [or] 25 miles N.N.W. of Arcas Cays, sandy
mud, 26 fathoms [47.6 m]; southern Campeche Bank, Mexico.
Dimensions: H= 8.9, W= 2.5. Remarks: Ex D. Steger.

2 paratypes: DMNH 118028; off Arcas Cays, southern Campeche
Bank, Mexico. Dimensions: (1) H= 7.4, W= 2.5; (2) H=
5.4, W= 2.0. Remarks: Ex D. Steger.

Terebridae.

atkana, Littorina sitkana [as n. var.]
Dall, 1886: 211.
2 syntypes: DMNH 043607; Kyska Island, Alaska. Dimensions: (1)
H= 16.0, W= 13.7 [with operculum]; (2) H= 16.0, W= 13.5.
Remarks: Ex USNM. Littorinidae.

aureola, Pyrene [as n. sp.]
Howard, F. B., 1963: 2.
1 paratype: DMNH 066821; Norse Beach, Punta Penasco, Sonora,
Mexico (31°20’N, 113°38’W); type locality. Dimensions: H=
17.6, W= 11.0. Columbellidae.

aurora, Columbarium (Peristarium) [as n. sp.]

Bayer, 1971: 180.

1 paratype: DMNH 153519; R/V ‘Pillsbury’ station P-1309, north-
east of Fowey Light, Straits of Florida (25°40’N, 80°02’W),
247 m; type locality. Dimensions: H= 42.9 [apex missing],
W= 11.0. Remarks: Ex Rosenstiel School of Marine and
Atmospheric Sciences, University of Miami. Columbariidae
[Turbinellidae].

axelrodi, Conus [as n. sp.]
Walls, 1978: 1, fig. p. 5.

July 30, 1991 Bieler & Bradford 7

Holotype: DMNH 123127; Palawan, Philippines. Dimensions: H=
16.0, W= 8.7. Remarks: Figure of holotype (Walls, 1978: fig.
p. 5 upper right) erroneously labeled ‘“‘topotypic paratype.”’
Also described by Walls (1979b: 183 ff.) without reference
to earlier publication. Conidae.

bailey1, Oliva [as n. sp.]

Petuch, 1979a: 10, figs. 18-19 (holotype).

Holotype: DMNH 126395; off Russell Island, Solomon Islands,
180 m. Dimensions: H= 30.3 (‘30’), W= 14.6 (‘‘8”’ [sic]).

1 paratype: DMNH 126396; type locality. Dimensions: H= 29.0
(‘28’), W= 13.7. Remarks: Erroneously cited as_ plural
“paratypes”? in the original description. This is not the
specimen figured as “‘paratype’’ by Petuch (1979a: 13, figs.
16,17).

Olividae.

barbarae, Brachycythara [as n. sp.]

Lyons, 1972: 4.

1 paratype: DMNH 047902; R/V ‘Hourglass’ station D, off Egmont
Key, Florida (27°37’'N, 83°58’W), 55 m; type locality. Dimen-
sions: H= 3.2, W= 1.7.

1 paratype: DMNH 107553; type locality. Dimensions: H= 2.9,
W= 1.5. Remarks: Ex D. Steger.

Turridae.

barnardi, Fusivoluta [as n. sp.]

Rehder, 1969: 207, pl. 40 fig. 9, pl. 43 figs. 40-41.

Holotype: DMNH 010751; off Natal coast, South Africa, 120
fathoms [220 m]. Dimensions: H= 115.2, W= 34.6.

1 paratype: DMNH 010750; off mouth Tugela River, South
Zululand, South Africa, 160-180 fathoms [293-329 m]. Di-
mensions: H= 96.5, W= 35.0. Remarks: ‘‘Early whorls and
anterior end missing”? (Rehder, 1969: 208).

Volutidae.

bermudensis, Rissoina [as n. sp.]
Peile, 1926: 98.
2 paratypes: DMNH 135069; ‘“‘Near Baeotia, St. Georges’’ [label],
Bermuda. Dimensions: (1) H= 5.6, W= 2.4; (2) H= 5.2,
W= 2.1. Remarks: Ex J. D. Parker. Rissoidae.

berryi, Anachis [as n. sp.]
Shasky, 1970: 190.

8 NEMOURIA No. 36

2 paratypes: DMNH 040849; El Pulmo Reef, Baja California,
Mexico (23°26’N, 109°25’W), 1-3 m. Dimensions: (1) H=
9.0, W= 3.3; (2) H= 8.5, W= 3.2. Remarks: Ex D. Shasky.
Columbellidae.

betsyae, Hastula (Punctoterebra) [as n. sp.]

R. D. Burch, 1965: 243.

2 paratypes: DMNH 006001; off Honokohau, Maui, Hawaii
(21°04’‘N, 156°38’W), 20 feet [6.1 m]; type locality. Dimen-
sions: (1) H= 29.7, W= 6.6; (2) H= 22.7, W= 5.9.

4 paratypes: DMNH 010141; locality data as above. Dimensions:
(1) H= 35.7, W= 7.2; (2) H= 26.5, W= 5.6; (3) H= 23.1,
W= 5.4; (4) H= 22.6, W= 5.3.

Remarks: Ex R. D. Burch collection no. 172. Terebridae.

blanes1, Olivella [as n. sp.]
Ford, 1898: 66.
2 syntypes: DMNH_ 136363; Cardenas, Cuba. Dimensions: (1)
H= 9.2, W= 4.2; (2) H= 6.3, W= 3.1. Remarks: Ex J. Parker.
Olividae.

boholensis, Conus [as n. sp.]

Petuch, 1979a: 12, figs. 20-21 (holotype).

Holotype: DMNH 126400; off Panglao, Bohol Island, Philippines,
approximately 250 m. Dimensions: H= 37.7 (°538”’), W= 17.3
[lip damaged] (“‘18’’).

2 paratypes: DMNH 126401; type locality. Dimensions: (1) H=
25.6 (‘26’), W= 11.3; (2) H= 18.5 (“17”), W= 8.6.

Remarks: Holotype figured by Coomans et al. (1982: 62, fig. 250),
as junior synonym of Conus borneensis Adams & Reeve, 1848.
Conidae.

boschi, Cymatium [as n. sp.]
Abbott & Lewis, 1970: 87, figs. 1-4.
Holotype: DMNH 022862; Al Masirah Island, Oman Protectorate,
southeast Arabia. Dimensions: H= 75.0, W= 41.0. Cymatiidae
[Ranellidae].

boswellae, Pirenella {as n. sp.]
Barnard, 1963: 140.
Holotype: DMNH 011370; ex pisce, off Durban, South Africa.
Dimensions: H= 53.2 (‘52’), W= 19.8 (‘‘19’’). Remarks:

Ex H. Boswell. Holotype figured by Barnard (1969: 650, figs.

25 a, b). Potamididae.

July 30, 1991 Bieler & Bradford 9

boswellae, Volutocorbis [as n. sp.]
Rehder, 1969: 202.
1 paratype: DMNH 019308; off Saint Sebastian Bay, South Africa,
in 80-100 fathoms [146-183 m]. Dimensions: H= 24.7, W=
10.9 Guvenile). Volutidae.

bouvetia, Margarella [as n. sp.]
Powell, 1951: 97.
2 paratypes: DMNH 106921; R/V ‘Discovery IT station 456, 1
mile east of Bouvet Island, 40-45 m; type locality. Dimensions:
(1) H= 7.0, W= 7.8; (2) H= 5.4, W= 6.5; both with dried
animals. Remarks: Ex A. W. Powell. Trochidae.

brasiliana, Borsonia [as n. sp.]

Tippett, 1983: 136.

2 paratypes: DMNH 161061; R/V ‘Oregon’ station 4226, 200 miles
north of Sao Luis, Brazil (00°18’N, 44°17’W), on the edge
of the continental shelf in 150 fathoms [274.5 m]; type locality.
Dimensions: (1) H= 11.1, W= 4.2; (2) H= 10.6, W= 4.0.
Remarks: In the original description, longitude erroneously
given as ‘‘004x17’°W.”’ Turridae.

burgessi, Distorsio [as n. sp.]
Lewis, 1972: 46.
1 paratype: DMNH 051956; Barber’s Point, S.W. Oahu Island,
Hawaii; type locality. Dimensions: H= 42.8, W= 28.8.
1 paratype: DMNH 093303; Haleiwa, Oahu, Hawaii. Dimensions:
H= 60.5, W= 36.0. Remarks: Ex C. Weaver.
Cymatiidae [Ranellidae].

clarkei, Fusivoluta [as n. sp.]

Rehder, 1969: 206, pl. 40 fig. 8, pl. 43 fig. 37 (holotype), pl. 43,
figs. 38, 39 (paratype DMNH 019307).

Holotype: DMNH 012833; off Joao Belo, District Gaza, Mozam-
bique, 240 fathoms [439 m]. Dimensions: H= 96.7, W= 31.3.

2 paratypes: DMNH 019305, 80 km east of Inhaca Island, District
Laurenco Marques, Mozambique. Dimensions: (1) H= 75.6,
W= 26.9; (2) H= 74.3 (“74.5”), W= 25.7.

1 paratype: DMNH 019307; 92 km east of Inhaca Island,
Mozambique. Dimensions: H= 79.8, W= 27.1.

Volutidae.

cloveri, Conus [as n. sp.]
Walls, 1978: 2, fig. p. 5 bottom left.

10 NEMOURIA No. 36

Holotype: DMNH 123128; harbor near Anse Bernard, Dakar,
Senegal. Dimensions: H= 25.7, W= 13.3. Remarks: Also
described by Walls (1979b: 338 ff.) without reference to earlier
publication. Conidae.

cloveriana, Lyria [as n. sp.]
Weaver, 1963b: 31.
1 paratype: DMNH 010139; 20 miles from ‘Tangalla, southern
Ceylon, 50 feet [15.2 m]. Dimensions: H= 43.6, W= 18.0
(‘17,3’) [juvenile]. Remarks: Ex R. Jonklaas. Subsequently
figured by Weaver & du Pont (1970: pl. 4 figs. M-N).
Volutidae.

cona, Acmaea [as n. sp.]|

Test, 1945: 92.

1 paratype: DMNH 023433; Point Fermin, San Pedro, California;
same locality data as “type series.’? Dimensions: greatest
length= 15.1, width= 11.7, height= 4.1. Remarks: Ex A. R.
G. Test, via ANSP 181024. Author mentions a “‘type series”’
of 20 specimens in addition to numerous “‘paratypes.’? Only
the California Academy of Sciences in San Francisco and the
United States National Museum are originally listed as
repositories. Acmaeidae.

corticata, Acmaea [as n. sp.]

Hutton, 1880: 89.

1 paralectotype: DMNH 023489; New Zealand. Dimensions: great-
est length= 12.0, width= 9.5, height= 5.1. Remarks: Ex F.
W. Hutton, via ANSP 50047. Powell (1979: 47), in the
redescription of Patelloida corticata (Hutton, 1880), refers to
a “‘type of corticata’’ with a length of 14 mm and the type
locality ‘“Dunedin.”’ This (or an earlier reference of this kind)
qualifies as designation of lectotype (ICZN Art. 74(b)).
Acmaeidae.

cosmani, Chicoreus [as n. sp.]
Abbott & C. J. Finlay, 1979: 159, fig. 8.
1 paratype: DMNH 028503; Discovery Bay, Jamaica, 35-100 feet
[10.7-30.4 m]. Dimensions: H= 39.0, W= 18.5. Muricidae.

craket, Duplicaria (Duplicaria) [as n. sp.]
R. D. Burch, 1965: 245.
2 paratypes: DMNH 006002; in fine sand at low tide on Cable
Beach, Broome, Western Australia (18°00’S, 122°15’E). Di- |

July 30, 1991 Bieler & Bradford 11

mensions: (1) H= 19.6, W= 4.5; (2) H= 16.8, W= 4.1.
Remarks: Ex R. D. Burch collection no. 336. Terebridae.

crosnieri, Cancilla scrobiculata [as n. ssp.]

Cernohorsky, 1970: 95, fig. 1 (holotype).

Holotype: DMNH 022392; Point Noire, Congo, West Africa
(4°57’S, 11°22’E), 135 m. Dimensions: H= 73.1, W= 18.5.

2 paratypes: DMNH 028221; Coast of Dahomey (6°05’N, 2°37’E),
West Africa; 150-200 m. Dimensions: (1) H= 71.0 (‘°71.4’’),

= 18.8; (2) H= 56.4, W= 14.6.
Mitridae.

cuvierensis, Microvoluta [as n. sp.]
H. J. Finlay, 1930: 242.
1 paratype: DMNH 012209; off Cuvier Island, Bay of Plenty, New
Zealand, 38 fathoms [69.5 ml]; type locality. Dimensions:
H= 5.7, W= 2.4. Volutomitridae.

daisyae, Volutoconus hargreavesi [as n. ssp.]

Weaver, 1967: 302, pl. 41 figs. 5-6 (holotype), fig. 7 (paratype
DMNH 010557), figs. 8-9 (paratype DMNH 010142).
Holotype: DMNH 010022; southwest of North West Cape, central
Western Australia (22°00’S, 113°45’E), 80 fathoms [146 m].

Dimensions: H= 85.3, W= 34.3.

1 paratype: DMNH 010557; Houtman Abrolhos Islands, southern
Western Australia; 85 fathoms [155 m]. Dimensions: H=
106.7 (‘96.0” [sic]), W= 42.6 (39.0’’). Remarks: Ex H.
Boswell.

1 paratype 010142; off Point Cloates, central Western Australia,
118 fathoms [216 m]. Dimensions: H= 85.2 (‘685.5’), W=
34.2 (35.0’). Remarks: Ex C. Weaver.

Volutidae.

dalli, Latiaxis (Babelomurex) [as n. sp.]

Emerson & D’Attilio, 1963: 4.

2 paratypes: DMNH 105930; locality description: ‘“‘southwest of
Egmont Key, Florida, 220-230 fathoms [403-421 m]‘‘; Steger
collection label: “‘from 120 fathoms [220 m], about 200 miles
SW of St. Petersburg, Fla.”’ Dimensions: (1) H= 35.7, W=
37.2 (incl. spines); (2) H= 34.5, W= 32.2 (incl. spines) [with
opercula]. Remarks: Ex D. Steger. Coralliophilidae [Muri-
cidae].

dalliana, Acmaea [as n. sp.]

12 NEMOURIA No. 36

Pilsbry, 1891: 13.

1 paratype: DMNH 023404; La Libertad, Sonora, Mexico. Dimen-
sions: greatest length= 39.3, width= 27.0, height= 8.0.
Remarks: Ex H. N. Lowe, via ANSP 167967. This locality
not listed in original description. Acmaeidae.

dampieria, Amoria (Amoria) [as n. sp.]

Weaver, 1960: 1, 3.

1 paratype: DMNH 010140; 25 miles off Bezout Island, Dampier
Archipelago, Western Australia, 25 fathoms [45.8 m]. Dimen-
sions: H= 33.2 (33.3), W= 15.2 (‘515.0’). Remarks: Ex
C. Weaver. Originally introduced in Hawazan Shell News by
Weaver (1960: 1, 3) when referring to “‘Amoria (Amoria)
dampteria Iredale, 1914, nomen nodum [sic],”’ with figure
and short description. Redescribed by Weaver (1963a: 28).
DMNH paratype figured and listed by Weaver (1963a: pl. 5,
two center figures; table 1, “‘paratype #2’’). Also figured by
Weaver & duPont (1970: pl. 63 figs. C-D). Volutidae.

darwint, Cancellaria [as n. sp.]

Petit, 1970: 85.

1 paratype: DMNH 027196; south of Academy Bay, Isla Santa
Cruz, Galapagos Islands, 170-200 m; type locality. Dimen-
sions: H= 17.6, W= 10.1. Remarks: Ex R. E. Petit. Cancel-
lariidae.

delli, Penion benthicola [as n. ssp.]
Powell, 1971: 222.
1 paratype: DMNH 026798; off Aldermen Islands, northern New
Zealand, 350 fathoms [640.5 m]. Dimensions: H= 68.4, W=
29.3. Remarks: Ex A. W. Powell. Buccinidae.

dispar, Neritina [as n. sp.]
Pease, 1867: 285.
6 syntypes: DMNH 153352; Rarotonga, Cook Islands. Dimensions
(heights): 7.2, 6.8, 6.8, 6.6, 6.3, 4.5 (with opercula). Remarks:
Ex N. W. Lermond. Neritidae.

disparilis, Volutocorbis [as n. sp.]
Rehder, 1969: 203, pl. 41 figs. 20-21.
3 paratypes: DMNH 010760; off Agulhas Bank, South Africa, 300
fathoms [549 mJ]. Dimensions: (1) H= 31.5, W= 17.1
(17.4); (2) H= 30.9, W= 13.5; (3) H= 30.2, W= 13.9
(14.0’’). Remarks: Specimens in DMNH 10760 match the

July 30, 1991 Bieler & Bradford 13

original description and measurements but do not fully agree
with the original figures (Rehder, 1969: pl. 41 figs. 20-21).
An additional specimen (DMNH 10761), figured by Weaver
& duPont (1970: pl. 3 figs. N-O) and cited as “‘paratype,”’ is
from the type locality but does not qualify as type material
because it was not included in the original description.
Volutidae.

donmooret, Caecum [as n. sp.|

Mitchell-Tapping, 1979: 105, figs. 29, 30.

Holotype: DMNH 119521; Sprat Bay, Water Island, United States
Virgin Islands, 5 m. Dimensions (greatest length): 2.2 (“1.4”
[sic]). Remarks: According to the original description (Mitchell-
Tapping, 1979: 105), the holotype was “deposited in 1977
in the Delaware Museum of Natural History #119521.’ The
DMNH specimen matches the published figure, but measures.
2.2 mm overall length instead of the dimension given in the
text (‘‘1.4 mm/’’) and those deduced from the scale accompa-
nying the published figure (also 1.4 mm). Caecidae.

dorothyae, Terebra [as n. sp.]
Bratcher & R. D. Burch, 1970: 297.
1 paratype: DMNH 022422; Allan Hancock Foundation station
770-38, 7-11 fathoms [12.8-20.1 m] on black sand off San
Jose Point, Guatemala (13°53’N, 91°09’W). Dimensions: H=
20.9, W= 5.6. Remarks: DMNH number erroneously given
as “22421” in original publication. Terebridae.

duponti, Festilyria [as n. sp.]
Weaver, 1968: 444, pl. 60 figs. 1-2.
Holotype: DMNH 013706; off Boa Paz, Mozambique, 40 fathoms
[73 m]. Dimensions: H= 124.4, W= 56.5. Volutidae.

eccentricus, Zelippistes [as n. sp.]
Petuch, 1979a: 5, figs. 7-9.
Holotype: DMNH 126391; China Strait, off Samarai Island, Eastern
Papua, Papua New Guinea, 250 m. Dimensions: H= 11.0
(“width 11 mm’), W= 17.9 (“length 18 mm/’’). Remarks:
Erroneously cited as “DMNH No. 123691” in original
description. Trichotropidae [Capulidae].

echinophilus, Africonus [as n. sp.]
Petuch, 1975: 180.
1 paratype: DMNH 121795; N’Gor, Cape Verde, Senegal, West

14 NEMOURIA No. 36

Africa (14°43’N, 17°33’W); type locality. Dimensions: H=
10.9, W= 6.1. Conidae.

ecphoroides, Fulgurofusus (Fulgurofusus) [as n. sp.]
Harasewych, 1983a: 20, fig. 28.
1 paratype: DMNH 153515; R/V ‘Oregon’ station 4226, 180 miles
N.E. of Tijoca, Para, Brazil (0°18’N, 44°17'W), 150 fathoms
[274 mJ]; type locality. Dimensions: H= 15.9 (“‘16”), W=
6.1. Turbinellidae.

electra, Columbarium (Peristarium) [as n. sp.]

Bayer, 1971: 176.

1 paratype: DMNH 153517; R/V ‘Gerda’ station G-289, S.S.E. of
Key West, Straits of Florida, (24°11’N, 81°36’W), 594-604
m; type locality. Dimensions: H= 16.2, W= 4.0. Remarks:
Ex Rosenstiel School of Marine and Atmospheric Sciences,
University of Miami. Protoconch of paratype illustrated by
Harasewych (1983a: 28, fig. 41). Columbariidae [Turbinelli-
dae].

elegans, Vitrinella (Striovitrinella) [as n. sp.]
Olsson & McGinty, 1958: 31.
1 paratype: DMNH 104277; Bocas Island, Panama [Caribbean].
Dimensions: H= 3.5, W= 1.6. Remarks: Ex D. Steger.
Vitrinellidae.

exquisita, Pseudocypraea [as n. sp.]
Petuch, 1979a: 6, figs. 5, 6.
Holotype: DMNH 126392; 250 m depth off Panglao, Bohol Island,
Philippines. Dimensions: H= 8.8 (‘9’), W= 5.4 (°5’’).
Ovulidae.

fayae, Anachis (?Costanachis) [as n. sp.]

Keen, 1971: 579.

2 paratypes: DMNH 047037; Playa Caracol, Nuevo Guayamas,
Sonora, Mexico; type locality. Dimensions: (1) H= 7.6, W=
2.9; (2) H= 6.6, W= 2.5. Remarks: Ex F. B. Howard, via
Santa Barbara Museum. The original description mentions a
holotype and ‘“‘[a]bout 200 specimens.’”’ Three illustrations
(Keen, 1971: 953, fig. 1178) are labeled “‘holotype,”’ “‘para-
type,”’ and “‘hypotype,”’ the two former from the type locality,
the last from San Marcos, California. Columbellidae.

finlayi, Murex (Murex) [as n. sp.]
Clench, 1955: 1.

July 30, 1991 Bieler & Bradford 15

1 paratype: DMNH 105880; Matanzas Bay, Cuba, 200 fathoms
[366 mJ]; type locality, but holotype is from “100 fathoms
[183 m].”? Dimensions: H= 74.7, W (apertural aspect, incl.
spines)= 44.7; with operculum. Remarks: Ex C. J. Finlay, via
D. Steger. Muricidae.

finlayi, Neritopsis [as n. sp.]
Hoerle, 1974: 104.
1 paratype: DMNH 099360; Banes Sands, Pifiar del Rio Province,
Cuba. Dimensions: H= 18.4, W= 17.3. Neritopsidae.

floridana, Stenacme [as n. sp.]

Pilsbry, 1945: 114.

1 paratype: DMNH 103823; Lake Worth, Boynton Beach, Florida.
Dimensions: H= 8.0, W= 4.9. Remarks: Ex T. L. McGinty,
via D. Steger. Described as a pulmonate in the nominal family
Stenacmidae; the species was subsequently recognized as a
member of the family Epitoniidae (Robertson & Oyama, 1958:
68-69).

floridanus, Microdochus [as n. sp.]
Rehder, 1943: 193.
70+ paratypes: DMNH 023747; Missouri Key, Florida. Dimensions:
heights from 1.6 to 2.5 mm. Remarks: Ex B. R. Bales, via
ANSP 220264. Rissoidae [Elachisinidae].

fragilissimus, Conus [as n. sp.]

Petuch, 1979a: 14, figs. 22-23 (holotype), figs. 24-25 (paratype
DMNH 126403).

Holotype: DMNH 126402; off south coast of Harmil Island, Dahlak
Archipelago, Eritrea Province, Ethiopia, Red Sea, 3 m.
Dimensions: H= 34.0 (‘30’), W= 15.6 (‘412’).

3 paratypes: DMNH 126403; type locality. Dimensions: (1) H=
30.2, W= 13.5 (figured); (2) H= 28.4, W= 12.8; (3) H= 26.8,
W= 13.5 (‘Lengths 34 mm, 31 mm, 26 mm’’).

Conidae.

gaylordi, Olivella gracilis [as n. var.]
Ford, 1895: 104.
2 syntypes: DMNH 024743; Gulf of California. Dimensions: (1):
H= 9.9, W= 3.9; (2) H= 8.5, W= 3.3. Remarks: Ex J. Ford,
via ANSP 316716. Olividae.

germonae, Tractolira [as n. sp.]
Harasewych, 1987: 3.

16 NEMOURIA No. 36

2 paratypes: DMNH 169441; R/V ‘Islas Orcada’ cruise 575, station
75, E.N.E. of Zavadovski Island, South Sandwich Islands
(56°03.5’S, 26°58.3°W), 2128-2161 m. Dimensions: (1) H=
48.7, W= 19.6; (2) H= 41.6, W= 16.6. Remarks: Both
specimens alcohol-preserved. Volutidae.

gtirardi, Cerodrillia [as n. sp.]
Lyons, 1972: 3.
1 paratype: DMNH 047901; R/V ‘Hourglass’ station D, off Egmont
Key, Florida (27°37’N, 83°58’W), 55 m; type locality. Dimen-
sions: H= 8.0, W= 3.5. Turridae.

gracilior, Benthovoluta [as n. sp.]

Rehder, 1967: 185.

1 paratype: DMNH 015456; U.S.B.F. ‘Albatross I’ station 5423,
off Cagayan Islands, northern Sulu Sea, Philippines (9°38’30’N,
121°11’E), 508 fathoms [930 m]. Dimensions: H= 49.9, W=
12.8. Remarks: Ex USNM 238408. Volutidae [Turbinellidae].

greegori, Cypraea cruenta [as n. var. |
Ford, 1893: 112.
1 syntype: DMNH 043613; no locality given. Dimensions: greatest
length= 24.2, width= 18.3. Remarks: Ex USNM (from I.
Greegor, via J. Ford). Cypraeidae.

guesti, Natica (Natica) [as n. sp.]

Harasewych & Jensen, 1984: 99, figs. 8, 10.

7 paratypes: DMNH 164500; 2.5 miles off Castle Roads, South
Shore, Bermuda, Lightbourn-Guest ‘Northstar’ expeditions;
in traps set in 347 m. Remarks: Lot contains 7 paratypes,
largest (H= 26.5, W= 27.0) illustrated in original fig. 8, plus
SEM stub with mounted shell apex shown in fig. 10.

1 [0] paratype: DMNH 096986; 2.5 miles off Castle Roads, South
Shore, Bermuda, Lightbourn-Guest ‘Northstar’ expeditions;
in traps set in 402 m. Remarks: Cited in original description
(1984: 100); not located.

Naticidae.

guesti, Pterynotus (Pterynotus) [as n. sp.]
Harasewych & Jensen, 1979: 3, figs. 6, 14.
Holotype: DMNH 122258; E.S.E. of Key West, Florida, 275 m.
Dimensions: H= 28.6. Muricidae.

habe1, Assiminea habei [as n. sp.]
Abbott, 1958b: 251.

July 30, 1991 Bieler & Bradford 17

2 paratypes: DMNH 025311; estuary, 2 miles south of Tanauan,
Leyte Island, Philippines; type locality. Dimensions: (1) H=
3.8, W= 2.5; (2) H= 3.6, W= 2.4. Remarks: Ex R. T. Abbott,
via ANSP 193366. Assimineidae.

habet, Distorsio constricta [as n. ssp.]
Lewis, 1972: 38.
1 paratype: DMNH 050943; Tosa Bay, Shikoku, Japan; type locality.
Dimensions: H= 50.6, W= 29.0. Cymatiidae [Ranellidae].

hancocki, Terebra [as n. sp.]
Bratcher & R. D. Burch, 1970: 299.
1 paratype: DMNH 051367; off La Libertad, Ecuador (2°08’S,
81°00’W); type locality. Dimensions: H= 27.4, W= 7.8.
‘Terebridae.

harrisae, Columbarium [as n. sp.]
Harasewych, 1983b: 28.
1 paratype: DMNH 153524; east of Lady Musgrave Island,
Queensland, Australia, 140 fathoms [256 mJ]. Dimensions:
H= 99.2 (99), W= 29.5. Remarks: With operculum.
Turbinellidae.

haysae, Thais floridana [as n. ssp.]

Clench, 1927: 6.

2 paratypes: DMNH 060612; Grand Bayou, Mississippi Delta,
Lousiana; type locality. Dimensions: (1) H= 84.3, W= 51.8;
(2) H= 78.1, W= 46.3.

2 paratypes: DMNH 152981; type locality. Dimensions: (1) H=
75.7, W= 44.3; (2) H= 66.5, W= 44.4.

Remarks: According to the original description, an unspecified
number of paratypes was originally deposited in the collections
of Museum of Comparative Zoology (Cambridge, MA),
ANSP, and Museum of Zoology, University of Michigan.
DMNH collection contains specimens labelled ‘“‘paratypes’’
ex E. Doremus (DMNH 060612) and ex N. W. Lermond
(DMNH 152981).

Muricidae.

_ hesperina, Blasicrura coxeni [as n. ssp.]
Schilder & Summers, 1963: 68.
1 paratype: DMNH 121541; Talasea, New Britain, Australia; type
locality. Dimensions: greatest length= 14.4, width= 7.9.
Remarks: Ex R. Summers. Cypraeidae.

18 NEMOURIA No. 36

hoffmeyeri, Terebra (Strioterebrum) [as n. sp.]|
Abbott, 1952: 78.
1 paratype: DMNH 024990; Manila Bay, Luzon Island, Philippines;
type locality. Dimensions: H= 19.8, W= 4.8. Remarks: Ex
E. M. Wistar, via ANSP 234286. Terebridae.

huttoni, Trochus (Canthiridus [sic, error for Cantharidus]) [as n. sp.]
E. A. Smith, 1876: 558.
2 syntypes: DMNH 136368; New Zealand. Dimensions: (1) H=
11.4, W= 10.5; (2) H= 9.1, W= 9.8. Remarks: Both with
dried animals. Trochidae.

inglei, Oceanida [as n. sp.]
Lyons, 1978: 543.
1 paratype: DMNH 121798; off St. Augustine, St. Johns County,
Florida, 38 m. Dimensions: H= 2.2, W= 0.9. Eulimidae.

johnstoneti, Melongena corona [as n. ssp.]

Clench & Turner, 1956: 178.

1 paratype: DMNH 126416; Little Lagoon, Gulf Shores, Alabama;
type locality. Dimensions: H= 82.5, W= 48.3.

1 paratype: DMNH 186770; type locality. Dimensions: H= 82.0,
W= 49.6.

5 paratypes: DMNH 072197; type locality.

2 paratypes: DMNH 092770; type locality.

Remarks: According to the original description, an unspecified |
number of paratypes is said to be in several institutional and |
private collections. DMNH collection contains specimens |

labelled ‘‘paratypes’? ex H.I. Johnstone (DMNH 126416, |
186770), ex G. & M. Kline (DMNH 072197) and ex H. & ©

M. Minzak (ODMNH 092770). Melongenidae [Buccinidae].

Raicherae, Vexillum (Costellaria) [as n. sp.]
Petuch, 1979b: 523.

2 paratypes: DMNH 121796; south end of Guaratibas Reefs, —
Abrolhos reef complex, Bahia State, Brazil (17°25’S, 39°08°W), |
1 m; type locality. Dimensions: (1) H= 9.1 (‘9’), W= 3.1; |

(2) H= 8.5 (‘9’), W= 3.1. Costellariidae.

Rellneri, Aulica [as n. sp.]
Iredale, 1957: 91.
1 paratype: DMNH 010032; Arnhem Land, Northern Territory,

Remarks: Ex C. Weaver. Volutidae.

Australia; type locality. Dimensions: H= 65.5, W= 34.5.

July 30, 1991 Bieler & Bradford 19

kermadecensts, Haurakia [as n. sp.]
Oliver, 1915: 518.
3 paratypes: DMNH 025359; off Sunday Island, Kermadec Islands;
type locality. Dimensions: (1) H= 1.8, W= 0.9; (2) H= 1.6,
W= 0.8; (3) H= 1.5, W= 0.8. Remarks: Ex R. S. Bell, via
ANSP 257017. Rissoidae.

kerstitchi, Conus [as n. sp.]
Walls, 1978: 2, fig. p. 6.
Holotype: DMNH 123130; Nayarit, off Isla Tres Marias (between
Isla San Juanito and Isla Maria Madre), Mexico. Dimensions:
H= 31.5 (‘31.8’), W= 15.9. Remarks: Also described by
Walls (1979a: 615 ff.) without reference to earlier publication,
with a poor figure of the holotype on p. 618. Conidae.

Ringae, Xenoturris [as n. sp.]
Powell, 1964: 325 [= p. 22-965].
2 paratypes: DMNH 010138; Waikiki, Oahu, Hawaiian Islands, 100
feet [30.4 m]. Dimensions: (1) H= 18.4, W= 7.0; (2) H=
15.6, W= 5.7. Remarks: Ex C. N. Cate. Original description
lists Waikiki material in D. Thaanum collection. Turridae.

Roto, Lamellaria [as n. sp.]

Schwengel, 1944: 17.

1 syntype: DMNH 024084; Bra[i]denton Beach, Florida. Dimen-
sions: H= 16.8, W= 14.0. Remarks: Ex J. S. Schwengel, via
ANSP 254013.

1 syntype: DMNH 139447; Bra[i]denton Beach, Florida. Dimen-
sions: H= 17.6, W= 14.0. Remarks: Ex J. Finlay.

Lamellariidae.

Rurzi, Guildfordia [as n. sp.]

Petuch, 1979a: 2, figs. 1-2 (holotype).

Holotype: DMNH 126389; off Balicasag, Bohol Island, Philippines,
300 m. Dimensions: H= 22.0, W (incl. spines)= 48.0 (“‘length
49 mm, width 47 mm’’). Remarks: With operculum.

4 paratypes: DMNH 126390; type locality. Dimensions: W (meas-
ured incl. spines)= 49.5, 48.4, 46.5, 42.2. Remarks: With
opercula.

Turbinidae.

Rurzi, Morum (Oniscidia) [as n. sp.]
Petuch, 1979: 7, figs. 10-13.
Holotype: DMNH 126393; off Panglao, Bohol Island, Philippines,

20 NEMOURIA No. 36

250 m. Dimensions: H= 22.4 (“423”), W= 15.2 (°15’’).
Cassidae [Harpidae].

lahainaenstis, Volva (Phenacovolva) [as n. sp.]

Cate, C., 1969: 365.

2 paratypes: DMNH 041616; off the southwest shore of the
Hawaiian Island of Maui, ‘‘Lahaina Roads’? between Lanai
and Maui (20°52’N, 156°41’E), on black coral, 185 feet [56.4
m]; type locality. Dimensions (greatest length and width): (1)
27.6, 8.5; (2) 14.5, 3.8 Guvenile, “‘expelled from the large
shell during cleaning’? according to label). Remarks: Ex C.
N. Cate. Ovulidae.

langi, Thais (Thaisella) [as n. sp.]
Clench & Turner, 1948: 1.
6 paratypes: DMNH 068665; Lohito Bay, Angola, Africa; type
locality. Dimensions: (1) H= 47.8, W= 34.1; (2) H= 47.5,
W= 35.4; (3) H= 47.0, W= 35.0; (4) H= 44.1, W= 33.0; (5)
H= 42.3, W= 30.6; (6) H= 39.9, W= 32.0. Remarks: Ex G.
& M. Kline. With opercula. Muricidae.

lenoreae, Phenacovolva (Subsimnia) [as n. sp.]
Cardin & Walls, 1980: 1, fig. p. 2.
Holotype: DMNH 157331; Isla Gobernador, Perlas Islands, Panama
(Pacific coast). Dimensions: H= 16.6 (“16.5”), W= 7.3.
Ovulidae.

lightbourni, Conus (Floraconus) [as n. sp.]

Petuch, 1986: 16, figs. 1-2 (holotype).

Holotype: DMNH 134938; 1.5 miles (2.5 km) due south of Castle
Island, Bermuda, from 180 fathoms [translates to 329.4 m,
not “497 m” as given in original description]. Dimensions:
H = “35”, W= ‘16’.

3 paratypes: DMNH 134939; type locality. Dimensions: (1) H=
47.8, W= 21.9; (2) H= 46.9, W= 22.0; (3) H= 39.7, W=
17.8; (‘lengths 26.0 mm [sic]—47.7 mm7’’).

Remarks: Holotype was listed in original description as deposited
in DMNH 134938, but has not been received. The original
publication mentions five additional paratypes (“lengths 22.4—
44 mm’) in the collection of Mr. J. R. H. Lightbourn,
Bermuda. In an attempt to trace the missing holotype and to
explain the mismatch of the DMNH paratypes with the
original description, Mr. Lightbourn’s material was re-
examined by us. It consists of four specimens. Three of these

|
|

July 30, 1991 Bieler & Bradford 21

are apparently part of the paratype series: (1) H= 36.7, W=
17.5 [lip damaged]; (2) H= 36.5, W= 17.2 [lip damaged];
(3) H= 34.0, W= 15.9. A fourth, much larger, specimen (H=
50.5, W= 22.0) was presumably added later. None of the
specimens studied matches the original illustration of the
holotype or the figure given by Réckel (1987: no. 78 Suppl.
[581]) as of a DMNH paratype (“45 x 19.5 mm/’’). Conidae.

lightbourni, Fusinus [as n. sp.]

Snyder, 1984: 28, figs. 1-2 (holotype), fig. 3 (paratype DMNH
154462).

Holotype: DMNH 154461; crabbed specimen from fish pot in
100-120 fathoms [183-220 m] taken 1.5 miles south of
Gurnet Rock, South Shore of Bermuda. Dimensions: H= 60.9
(617’), W= 20.5.

3 paratypes: DMNH 154462; type locality, but “‘depths down to
200 fms.” [366 m]. Dimensions: (1 [figured]) H= 66.4
(‘66’); (2) H= 60.6 (“61”); (3) H= 45.8 (‘*46’’).

Fasciolariidae [Buccinidae].

lightbourni, Pterynotus (Pterynotus) [as n. sp.]
Harasewych & Jensen, 1979: 8, fig. 7.
Holotype: DMNH 122259; off St. David’s, Bermuda, 275-600 m.
Dimensions: H= 33.8. Muricidae.

lyalliana, Rissoia (Alvania) plicata [as n. var.]
Suter, 1898: 6.
1 paratype: DMNH 135066; Lyall Bay, near Wellington, New
Zealand; type locality. Dimensions: H= 2.5, W= 1.05. Re-
marks: Ex J. D. Parker. Rissoidae.

macleant, Calliostoma [as n. sp.]
Shasky & Campbell, 1964: 117.
1 paratype: DMNH 040850; N.W. side of Ensenada Lalo, Guaymas,
Sonora, Mexico, 30-45 feet [9.1-13.7 m]. Dimensions: H=
10.1, W= 9.0. Remarks: Ex D. Shasky. Trochidae.

macleani, Coralliophila [as n. sp.]

Shasky, 1970: 189.

2 paratypes: DMNH 040846; Saladita Bay, Guaymas, Sonora,
Mexico (27°53715’N, 110°59’W), 3-4 m, on the bases of white
gorgonid sea whips; type locality. Dimensions: H= 10.9, W=
8.1. Remarks: Ex D. Shasky. Coralliophilidae [Muricidae].

macquariensis, Trophon [as n. sp.]

22 NEMOURIA No. 36

Powell, 1957: 143.

3 paratypes: DMNH 106911; B.A.N.Z.A.R.E. station 83, off
Lusitania Bay, Macquarie Island, 69 m; type locality. Dimen-
sions: (1) H= 22.7, W= 10.6; (2) H= 19.8, W= 11.0; (@)
H= 16.4, W= 9.0. Remarks: Ex A. W. Powell. Muricidae.

major, Buccinum pemphigus [as n. var.]

Dall, 1919: 325.

1 syntype: DMNH 043618; U.S. Fish Commission (‘Albatross’)
station 3643, in the western part of Bering Sea, S.E. of
Kamchatka, in 100 fathoms [183 m]. Dimensions: H= 65.5,
W= 34.7. Remarks: Ex USNM. Buccinidae.

marmosa, Olivella minuta [as n. ssp.]
Olsson & McGinty, 1958: 38.
1 paratype: DMNH 104276; Bocas Island, Bocal del Toro, Panama,
Caribbean; type locality. Dimensions: H= 6.0, W= 2.7.
Remarks: Ex T. L. McGinty, via D. Steger. Olividae.

martini, Latirus [as n. sp.]
Snyder, 1988: 54, fig. 2.
1 paratype: DMNH 169442; north coast, Roatan Island, Honduras,
15-20 feet [4.6-6.1 m]. Dimensions: H= 23.8, W= 11.9.
Fasciolariidae [Buccinidae].

maryleeae, Hastula (Hastula) [as n. sp.]
R. D. Burch, 1965: 242.
2 paratypes: DMNH 006003; in sand at low tide, Surfside Beach,
Freeport, Texas (28°57'N, 95°38’W). Dimensions: (1) H=
21.6, W= 4.7; (2) H= 15.8, W= 4.1. Remarks: Ex R. D.
Burch collection no. 023. Terebridae.

mastirana, Persicula [as n. sp.]
Roth & Petit, 1972: 80, figs. 1, 2.
Holotype: DMNH 051327; Ras Jidufah (local name: Monument
Beach) at northeast tip of Masirah Island (21°N, 59°E),
Arabian Sea, off the coast of the Sultanate Muscat and Oman.
Dimensions: H= 9.8 (‘10.0’), W= 6.9 (*°7.2”’). Marginellidae.

mauiensis, Cypraea [as n. sp.]

Burgess, 1967: 6.

2 paratypes: DMNH 048221; reef at Olowalu, Maui Island, Hawaii
(20°48.5'N, 156°37.3’W); type locality. Dimensions [greatest
lengths]: (1) 13.9; (2) 13.4. Remarks: Ex C. S. Weaver.
Cypraeidae.

July 30, 1991 Bieler & Bradford 23

mazatlanica, Ruthia [as n. sp.]
Shasky, 1970: 192.
1 paratype: DMNH 040848; east side of Chivos Island, Mazatlan,
Sinaloa, Mexico. Dimensions: H= 7.8, W= 2.6. Remarks: Ex
D. Shasky. Columbellidae.

megintyt, Eubela [as n. sp.]
Schwengel, 1943: 76.
1 paratype: DMNH 108007; off Lake Worth, Palm Beach County,
Florida, 400-500 feet [122-152 m, label] or 80 fathoms [146.4
m, description]; type locality. Dimensions: H= 6.4, W= 2.4.
Remarks: Ex T. L. McGinty, via D. Steger. Turridae.

mendozana, Strombina (Cotonopsis) [as n. sp.]
Shasky, 1970: 194.
1 paratype: DMNH 040847; Gulf of Fonseca, El Salvador (15°57’N,
95°32’°W), 33-73 m; type locality. Dimensions: H= 23.1, W=
9.3. Remarks: Ex D. Shasky. Columbellidae.

merope, Columbarium (Peristarium) [as n. sp.]
Bayer, 1971: 178.
1 paratype: DMNH 153518; R/V ‘Gerda’ station G-970, south of
Marquesas Keys, Straits of Florida (24°24’N, 82°08’W), 512
m. Dimensions: H= 23.3, W= 6.1. Remarks: Ex Rosenstiel
School of Marine and Atmospheric Sciences, University of
Miami. Paratype illustrated by Harasewych (1983a: 34, fig.
54). Columbariidae [Turbinellidae].

meyeri, Conus biliosus [as n. ssp.]

Walls, 1979b: 3.

Holotype: DMNH 122117; Genezzano, Natal, South Africa. Dimen-
sions: H= 44.0, W= 25.3 (‘‘24.6’’). Remarks: The original
description refers to a figure by Walls (1978b”’ [error for
1979a]: 160, bottom). This is in error, the holotype was
figured by Walls (1979a: 161, bottom left). Conidae.

michaeli, Ocinebra [as n. sp.]
Ford, 1888: 188.
2 syntypes: DMNH 152945; Cayucos, San Luis Obispo County,
California. Dimensions: (1) H= 14.3, W= 7.3; (2) H= 14.3,
W= 7.1. Remarks: Ex W. E. Webb, via N. W. Lermond.
Muricidae.

nanus, Strombus raninus [as n. ssp.]
Bales, 1942: 19.

24 NEMOURIA No. 36

5 paratypes: DMNH 001490; southern end of Lake Worth, Palm
Beach County, Florida; type locality. Dimensions: (1) H=
57.4, W= 38.0; (2) H= 52.7, W= 37.1; (3) H= 51.5, W=
38.7; (4) H= 51.0, W= 33.6; (5) H= 48.8, W= 33.9. Remarks:
Original description lists types in ANSP, and further paratypes
in “‘the Bales, McGinty and Koto collections.’”» DMNH
records do not indicate origin of material. Strombidae.

nereis, Conus [as n. sp.]

Petuch, 1979a: 18, figs. 32-33 (holotype).

Holotype: DMNH 126404; off Panglao, Bohol Island, Philippines,
approximately 250 m. Dimensions: H= 22.8 (‘‘23’’), W= 12.3
Gage?

1 paratype: DMNH 126405; type locality. Dimensions: H= 14.7
(147), W= 7.1.

Conidae.

nivosa, Ovirissoa [as n. sp.]
Powell, 1957: 141.
3 paratypes: DMNH 106944; B.A.N.Z.A.R.E. station 83, off
Lusitania Bay, Macquarie Island, 69 m; type locality. Dimen-
sions: (1) H= 2.5, W= 1.4; (2) H= 2.3, W= 1.4; (3) H= 2.3,
W= 1.3. Remarks: Ex A. W. Powell. Rissoidae.

nodosivaricosa, Agatrix (Olssonella) [as n. sp.]
Petuch, 1979a: 11, figs. 26, 27.
Holotype: DMNH 126397; off Balicasag, Bohol Island, Philippines,
300 m. Dimensions: H= 12.9 (“117), W= 8.4 (‘9’).
Cancellariidae.

orcutti, Caecum [as n. sp.]
Dall zn Orcutt & Dall, 1885: 541.
2 syntypes: DMNH 161855; San Diego, California. Dimensions
(greatest lengths): 2.1; 1.7. Remarks: Ex N. W. Lermond.
Caecidae.

parkinsoni, Terebra [as n. sp.]
Cernohorsky & Bratcher, 1976: 137.
1 paratype: DMNH 112104; Nordup, East New Britain, Papua
New Guinea; type locality. Dimensions: H= 28.1, W= 5.3.
Terebridae.

parvatus, Conus musicus [as n. ssp. |]
Walls, 1979b: 4.
Holotype: DMNH 122118; Natal, South Africa. Dimensions: H=

July 30, 1991 Bieler & Bradford 25

21.5 (‘21.4’), W= 13.0. Remarks: The original description
refers to figures by Walls (“‘1978b’’ [error for 1979a]: 485
bottom left, 752) which are not of the holotype specimen.
Ex DMNH 017449.

6 paratypes: DMNH 133931; type locality. Dimensions: H= 21.9;
21.7; 19.6; 18.9, 18.2; 17.4. Remarks: The original description
refers to figures by Walls (“‘1978b” [error for 1979a]: 485,
bottom left; 752) which are not of these specimens. Ex
DMNH 017449.

Conidae.

patae, Conus [as n. sp.]

Abbott, 1971: 49, fig. 1 (holotype), figs. 4-5 (paratypes DMNH
040595), fig. 6 (paratype DMNH 044095).

Holotype: DMNH 044097; off Pompano Beach, Broward County,
Florida, 10 fathoms [18.3 m]. Dimensions: H= 24.4, W=
14.1 (‘°13.8’?). Remarks: Holotype also figured by Walls
(1979a: 524, upper left).

1 paratype: DMNH 044096; type locality. Dimensions: H= 23.9,
W= 14.4.

3 [2] paratypes: DMNH 040595; 0.5 miles off Lauderdale-by-the-
Sea, Florida, 50-100 feet [15.2-30.4 m]. Dimensions: (1)
H= 25.2 (25.07), W= 14.5 (“14.2”); (2) H= 24.7 (“‘24.5’’),
W= 13.6. Remarks: Original description cites three paratypes
under this lot number. Only two were originally catalogued.
The smaller one was figured by Walls (1979a: 524, upper
right).

2 [1] paratypes: DMNH 044095; Ocho Rios, north central coast
of Jamaica. Dimensions: H= 24.0, W= 13.3 (‘513.0’). Re-
marks: The original description cites two paratypes under this
lot number. Both were entered into the museum’s catalog,
but only one specimen remains in the lot. It is not the shell
figured by Walls (1979a: 524, bottom).

Conidae.

perplicata, Voluta [as n. sp.]

Hedley, 1902: 309.

1 syntype: DMNH 062301; “taken. . . either on the neighbouring
reefs of the Great Barrier, or on one of the atolls of the Coral
Sea.’ Dimensions: H= 59.5, W= 28.7. Remarks: Ex Australian
Museum, Sydney, no. C.9679. Hedley mentioned three
specimens and did not select one as holotype. Weaver &
duPont’s (1970: 96) erroneous reference to a “‘holotype’”’ does

26 NEMOURIA No. 36

not qualify as lectotype designation (ICZN Art. 74[b]).

philippinensis, Axymene [as n. sp.]
Petuch, 1979a: 8, figs. 14, 15.
Holotype: DMNH 126394; off Panglao, Bohol Island, Philippines,
250 m. Dimensions: H= 15.0 (‘‘16’’), 7.7 (‘8’). Muricidae.

pseudanaloga, Puncturella [as n. sp.]

Powell, 1957: 140.

2 paratypes: DMNH 022463; off Macquarie Island, Sub-Antarctica,
69 m; apparently from type locality. Dimensions [height from
aperture, greatest shell length]: (1) 6.3, 10.5; (2) 5.8, 9.3;
both with dried animals. Remarks: Ex J. E. du Pont. |

3 paratypes: DMNH 106910; B.A.N.Z.A.R.E. station 83, off —
Lusitania Bay, Macquarie Island, Sub-Antarctica; type locality. —
Dimensions [height from aperture, greatest shell diameter]:
(1) 7.5, 11.5; (2) 5.3, 9.4; (3) 5.1, 9.15 all with dried animals.
Remarks: Ex A. W. Powell.

Fissurellidae.

queenslandica, Leucosyrinx [as n. sp.]
Powell, 1969: 338.
2 paratypes: DMNH 026584; off Cape Moreton, Queensland,
Australia, in 100 fathoms [183 m]; type locality. Dimensions:
(1) H= 48.5, W= 15.0; (2) H= 47.6, W= 13.3. Turridae.

radwini, Pterynotus (Pterynotus) [as n. sp.]
Harasewych & Jensen, 1979: 11, figs. 10, 18 (radula).
Holotype: DMNH 122424; R/V “Pillsbury’’ station P-610, east of
Turneffe Islands, Belize, 296-329 m [17°2.0’N, 87°38.4'W].
Dimensions: H= 30.9. Muricidae.

randalli, Cymbiola [as n. sp.]
Stokes, 1961: [3].
1 paratype; DMNH 010136; 3 miles from Undine Reef, North
Queensland, Australia; type locality. Dimensions: H= 39.1,
W= 20.3. Remarks: Ex A. J. Stokes, via C. Weaver. DMNH
paratype figured by Weaver & duPont (1970: pl. 41 fig. C).
Volutidae.

reductaspiralis, Conus nielsenae [as n. ssp.]
Walls, 1979b: 5, fig. p. 5.
Holotype: DMNH 122119; Geraldton, Western Australia. Dimen-
sions: H= 33.3 (‘33.4’), W= 18.3. Remarks: The original
description refers to figures in Walls (‘‘1978b” [error for

July 30, 1991 Bieler & Bradford 27

1979a]: 496, bottom right; 762) which are not of the holotype
specimen.
Conidae.

rejecta, Oliva [as n. sp.]
J. Q. Burch & R. L. Burch, 1962: 166.
1 paratype: DMNH 005398; tide flats at La Paz, Baja California,
Mexico; type locality. Dimensions: H= 37.6, W= 16.3.
Olividae.

riost, Adelomelon (Weaveria) [as n. sp.]

Clench & Turner, 1964: 162.

1 paratype: DMNH 010767; about 150 miles E.S.E. of Cabo San
Antonio, Argentina (36°40’S, 53°08'W [54°08'W?]), 95 fath-
oms [174 m]. Dimensions: H= 231.0, W= 103.0. Remarks:
This is not the specimen illustrated under this DMNH
number by Weaver & duPont (1970: pl. 44, figs. C-D).
Volutidae.

rosselli, Zoila [as n. sp.]
Cotton, 1948: 30.
1 paratype: DMNH 005403; beach near North Wharf, Fremantle,
Western Australia. Dimensions: greatest length= 54.5, width=
34.4. Remarks: Ex Rossell via C. Weaver. Cypraeidae.

rubra, Alcyna [as n. sp.]
Pease, 1860: 436.
2 syntypes: DMNH 022968; Sandwich Islands [Hawaiian Islands].
Dimensions: (1) H= 2.8, W= 1.8; (2) H= 2.6, W= 1.7.
Remarks: Ex ANSP 37834. Trochidae.

salisburyt, Mitrolumna [as n. sp.]
Cernohorsky, 1978: 66.
1 paratype: DMNH 112101; Maile Point, Oahu, Hawaiian Islands,
32 fathoms [58.6 mJ]; type locality. Dimensions: H= 4.0
(‘°3.9’’), W= 1.9. Turridae.

salisburyi, Vexillum (Pusia) [as n. sp.]
Cernohorsky, 1976: 114.
3 paratypes: DMNH 112102; Pupukea beach, Oahu, Hawaiian
Islands; type locality. Dimensions: (1) H= 5.0, W= 2.4; (2)
H= 5.0, W= 2.3; (3) H= 4.8, W= 2.3. Mitridae [Costel-
lariidae].

_ sandwichensis, Conus suturalis [as n. ssp.]

28 NEMOURIA No. 36

Walls, 1978: 3, figs. p. 7.

Holotype: DMNH 123126; Pokai Bay, Oahu, Hawaii. Dimensions:
H= 14.4, W= 7.2 (juvenile).

Conidae.

scalaroides, Rissoa [as n. sp.]
C. B. Adams, 1850: 113.
7 paralectotypes: DMNH 135070; Jamaica; type locality. Dimen-
sions: (1) H= 4.6, W= 2.0; (2) H= 4.6, W= 2.0; (3) H= 4.6,
W= 1.9; (4) H= 4.5, W= 1.8; (5) H= 4.2, W= 2.0; (6) H=
3.6, W= 1.6; (7) fragment. Remarks: Ex J. D. Parker.
Lectotype in Museum of Comparative Zoology (Cambridge,
Massachussetts), number 177103, see Clench & Turner (1950:
340, pl. 33 fig. 6). Rissoidae.

scotiana, Sinuber sculpta [as n. ssp.]
Powell, 1951: 120.
2 paratypes: DMNH 106920; R/V ‘Discovery II’ station 167, off
Signy Island, South Orkneys (60°50’30”S, 46°15’00"W), 244-
344 m; type locality. Dimensions: (1) H= 9.1, W= 7.2; (2)
H= 7.6, W= 6.5; both with dried animals. Remarks: Ex A.
W. Powell. Naticidae.

signeyana, Chlanidota [as n. sp.]
Powell, 1951: 141.
1 paratype: DMNH 106917; R/V ‘Discovery II’ station 167, off
Signy Island, South Orkneys (60°50’30”S, 46°15’00"W), 244-
344 m; type locality. Dimensions: H= 28.6, W= 17.8.
Remarks: Ex A. W. Powell. Buccinulidae [Buccinidae].

smithi, Prodallia [as n. sp.]
Bartsch, 1942: 11.
1 paratype: DMNH 015454; U.S. Bureau of Fisheries (‘Albatross’)
station 5124, Maestre de Campo Island, 281 fathoms [514
m, off east coast of Mindoro, Philippines]. Dimensions: H=
51.0, W= 14.5. Remarks: Ex USNM 235394. Volutidae.

stegeri, Daphnella [as n. sp.]
McGinty, 1955: 76.
1 paratype: DMNH 119526; W.S.W. of John’s Pass, Gulf Beaches,
W. Florida, 35 fathoms [64 m]. Dimensions: H= 17.2, W=
7.1. Remarks: Ex D. Steger. Turridae.

stegeri, Olivella (Olivella) [as n. sp.]
Olsson, 1956: 176.

July 30, 1991 Bieler & Bradford 29

4 paratypes: DMNH 105286; S.W. of Egmont Key, Florida, 78-90
fathoms [143-165 m]. Dimensions: (1) H= 11.1, W= 5.0; (2)
H= 10.3, W= 4.5, (3) H= 9.7, W= 4.9, (4) H= 9.5, W=
4.5. Remarks: Ex D. Steger. Olividae.

stegeri, Terebra (Myurella) floridana [as n. ssp.]
Abbott, 1954: 39.
1 paratype: DMNH 105856; W.N.W. of Progresso, Mexico, 18
fathoms [32.9 m]. Dimensions: H= 78.4, W= 15.5. Remarks:
Ex D. Steger. Terebridae.

stilesi, Neptunea [as n. sp.]
A. G. Smith, 1968: 117.
1 paratype: DMNH 106558; between La Perouse Bank and Cape
Scott, Vancouver Island, British Columbia, ca. 100 fathoms
[183 ml]; type locality. Dimensions: H= 80.0 (apex broken),
W= 60.0. Remarks: Ex E. C. Stiles, via California Academy
of Sciences and A. Smiley. Buccinidae.

summerst, Bistolida (Blasicrura) pallidula [as n. ssp.]
Schilder, 1958: 85.
1 paratype: DMNH 121540; Nukalofa, Tonga-Tabu, Tonga; type
locality. Dimensions: greatest length= 16.7, width= 9.4.
Remarks: Ex J. M. Ostergaard, via R. Summers. Cypraeidae.

teaguet, Olivancillaria [as n. sp.]
Klappenbach, 1964: 132.
1 paratype: DMNH 024729; La Coronilla, Department of Rocha,
Republic of Uruguay. Dimensions: H= 18.9, W= 10.2.
Remarks: Ex ANSP 290783. Olividae.

textimattum, Epitonium (Asperiscala) [as n. sp.]
DuShane, 1977: 89.
1 paratype: DMNH 121769; Playa Los Angeles, Tenacatita Bay,
Jalisco, Mexico (19°18’N, 104°50’W), 7-18 m; type locality.
Dimensions: H= 10.9, W= 3.5. Epitoniidae.

thompson, Eudolium [as n. sp.]
McGinty, 1955: 80.
1 paratype: DMNH 105924; off Tampa, Florida, Gulf of Mexico
[publication], approx. 125 miles SW Egmont Key, Florida
[label]. Dimensions: H= 38.8 [apex missing], W= 27.2.
Remarks: Ex D. Steger. Tonnidae.

tumor, Fulgurofusus (Peristarium) [as n. sp.]

30 NEMOURIA No. 36

Harasewych, 1983a: 34, fig. 49.
1 paratype: DMNH 153516; off North Carolina, deep water.
Dimensions: H= 53.6, W= 12.4. Turbinellidae.

tokyoensis, Rissoa [as n. sp.]
Pilsbry, 1904: 26.
1 syntype: DMNH 136367; Tokyo Harbor, Japan. Dimensions:
H= 2.2, W= 1.0. Remarks: “‘From type lot’ ex Y. Hirase, via
J. Parker [ANSP 70910]. Rissoidae.

tribblei, Conus [as n. sp.]
Walls, 1977: 2, figs. p. 3.
Holotype: DMNH 123125; China Sea, off Taiwan. Dimensions:
H= 62.4 (‘°62.3’), W= 30.4.
Conidae.

trujilloi, Oliva [as n. sp.]
Clench, 1938: 111.
7 paratypes: DMNH 063513; Puerto Plata, Santo Domingo,
dredged at about 5 fathoms [9.1 m]; type locality. Dimensions
(heights): 41.6, 39.2, 38.7, 37.6, 36.5, 35.4, 31.8. Olividae.

venustula, Pareuthria [as n. sp.]

Powell, 1951: 134.

1 paratype: DMNH 106912; R/V ‘Discovery II’ station 388, between
Cape Horn and Staten Island (56°19.5’S, 67°09"W), 121 m;
type locality. Dimensions: H= 14.8, W= 6.5. Remarks: Ex
A. W. Powell. Buccinulidae [Buccinidae].

veracruzensis, Cerithidea pliculosa [as n. ssp.]

Bequaert, 1942: 5.

1 paratype: DMNH 023859; Estero de Espantaperros, Isthmus of
Tehuantepec, Mexico. Dimensions: H= 17.2, W= 7.5 [apex
broken]. Remarks: Ex ANSP 178508 [= old number ANSP
124569, as cited in original publication]. Potamididae
[Cerithideidae].

vernicosum, Mangilia (Calliotectum) [as n. sp.]
Dall, 1889: 304.
1 syntype: DMNH 010135; U.S. Fish Commission (‘Albatross’)
station 2807, near the Galapagos Islands, Pacific, 812 fathoms
[1486 m]. Dimensions: H= 47.0, W= 18.7. Remarks: Ex C.
Weaver. Pleurotomidae [Turridae].

vina, Assiminea succinea [as Nn. ssp.]

July 30, 1991 Bieler & Bradford 31

Marcus, Eveline & Ernst Marcus, 1965: 51.

31 syntypes: DMNH 033740; Cananéia, Brazil (25°01’S, 47°50°W).
Dimensions: heights from 1.2 to 1.7 mm. Remarks: Original
description is based on “a great number’ of specimens.
Assimineidae.

weavert, Turridrupa [as n. sp.]
Powell, 1967: 423.
1 paratype: DMNH 010143; Mokumanu Island, north shore of
Oahu, Hawaiian Islands, 65-70 feet [19.8-21.3 m]. Dimen-
sions: H= 17.6, W= 7.1. Turridae.

wittigi, Conus [as n. sp.]
Walls, 1977: 1, figs. p. 3.
Holotype: DMNH 123129; Lesser Sunda Islands north of Timor
[‘‘Straits of Flores’? according to specimen label]. Dimensions:
H= 31.7 (31.8), W= 15.7.
Conidae.

wolfet, Vexillum (Costellaria) [as n. sp.]

Cernohorsky, 1978: 63.

il paratype: DMNH 112103; Pokai Bay, Oahu Island, Hawaiian
Islands, 60-70 fathoms [110-128 ml]; type locality. Dimen-
sions: H= 5.9 (‘6.0’), W= 2.3 (‘2.2’). Mitridae [Costel-
lariidae].

wormaldi, Columbarium (Coluzea) [as n. sp.]

Powell, 1971: 221.

1 paratype: DMNH 026797; Aldermen Island, northern New
Zealand, 350 fathoms [640.5 m]. Dimensions: H= 82.3, W=
28.5. Remarks: Localities given in description: East of Poor
Knights Islands, 549 m (holotype), and 23 miles NE of Cuvier
Island, 260-270 fathoms [476-494 m]. Columbariidae [Turbi-
nellidae].

xenismatis, Fulgurofusus (Histricosceptrum) [as n. sp.]

Harasewych, 1983a: 26, fig. 36.

1 paratype: DMNH 153523; R/V ‘Pillsbury’ station P-1354, off
northeastern Nicaragua (14°21’N, 81°55’W), 192-263 m; type
locality. Dimensions: H= 32.4 (‘°33’’), W= 12.4. Turbinelli-
dae.

zebrina, Zediloma (Fractarmilla) corrosa [as n. ssp.]
Powell, 1946: 137.
3 paratypes: DMNH 022483; Beach Harbour, Breaksea Sound,

52 NEMOURIA No. 36

Fiordland, W. Ortago, New Zealand; type locality. Dimen-
sions: (1) H= 11.0, W= 12.7; (2) H= 11.0, W= 12.5, (3)
H= 10.3, W= 12.2. Remarks: Ex J. E. du Pont.

3 paratypes: DMNH 027487; type locality. Dimensions: (1) H=
12.2, W= 12.5, (2) H= 11.3, W= 12.5, (3) H= 11.0, W=
12.5. Remarks: Ex J. H. Alexander.

Trochidae.

zulu, Conus [as n. sp.]

Petuch, 1979a: 19, figs. 28-29 (holotype), figs. 30-31 (paratypes
DMNH 126407).

Holotype: DMNH 126406; off the mouth of the Umfolozi River,
Zululand coast, Natal, South Africa, approximately 40 m.
Dimensions: H= 62.3 (“62”), W= 37.0 (“35’’).

4 [3+2] paratypes: DMNH 126407; type locality. Dimensions: (1)
H= 57.3, W= 33.9 [figured]; (2) H= 61.6, W= 35.6; (3)
H= 57.2, W= 30.8. Remarks: The original description cites
four paratypes in DMNH 126407 (“‘Lengths 58 mm, 62 mm,
65 mm, 68 mm7’’). The lot contains five specimens: three of
them bearing the lot number (listed above), and two smaller,
unnumbered ones (H= 56.4, W= 33.1; H= 56.2, W= 30.9)
of dubious status.

1 paratype: DMNH 126408; off Inhaca Island, Mozambique, 35
m. Dimensions: H= 66.3 (‘‘74”’ [sic]), W= 39.5.

Conidae.

SUBCLASS OPISTHOBRANCHIA

eloiseae, Acteon [as n. sp.]

Abbott, 1973: 91, figs. 1-3 (holotype).

Holotype: DMNH 027213; Al Masirah Island, Oman Protectorate,
southeast Arabia. Dimensions: H= 30.9 (‘31.0’), W= 20.0
(“19.5”’). Remarks: With operculum.

2 paratypes: DMNH 067746; type locality. Dimensions: (1) H=
34.5, W= 22.2; (2) H= 31.0, W= 19.5.

Acteonidae.

mauganuica, Austrodiaphana [as n. sp.]

Powell, 1952: 185.

10 paratypes: DMNH 022727; ex pisce, 2.5 miles north of Mount
Maunganui, Bay of Plenty, New Zealand; type locality.
Dimensions: lengths ranging 1.17—1.92. Remarks: Ex J. E.
duPont.

July 30, 1991 Bieler & Bradford 33

9 paratypes: DMNH 027082; type locality. Dimensions: lengths
ranging 1.33—1.67. Remarks: Ex G. Williams.

19 paratypes: DMNH 027428; type locality. Dimensions: lengths
ranging 1.08—1.75. Remarks: Ex J. H. Alexander. One Azys-like
shell removed from this lot.

Austrodiaphanidae [Diaphanidae].

tenerum, Parvaplustrum [as n. sp.]
Powell, 1951: 180.
6 paratypes: DMNH 106916; R/V ‘Discovery II’ station WS-219,
north-west of Falkland Islands (47°06’S, 62°12’°W), 114-116
m; type locality. Dimensions: lengths= 2.92, 2.83, 2.75, 2.67,
2.50, 2.17. Remarks: Ex A. W. Powell. All specimens with
dried animals. Aplustridae [Hydatinidae].

34 NEMOURIA No. 36
ALPHABETIC INDEX BY AUTHOR

Abbott, R. T.
arcas, Terebra (Strioterebrum), Terebridae
eloiseae, Acteon, Acteonidae
habe1, Assiminea habei, Assimineidae
hoffmeyert, Terebra (Strioterebrum), Terebridae
patae, Conus, Conidae
stegert, Terebra (Myurella) floridana, Terebridae

Abbott, R. T. & C. J. Finlay
cosmani, Chicoreus, Muricidae

Abbott, R. T. & H. Lewis
boschi, Cymatium, Cymatiidae [Ranellidae]

Adams, C. B.
scalaroides, Rissoa, Rissoidae

Bales, B. R.
nanus, Strombus raninus, Strombidae

Barnard, K. H.
boswellae, Pirenella, Potamididae

Bartsch, P.
smithi, Prodallia, Volutidae

Bayer, F. M.
aurora, Columbarium (Peristarium), Columbariidae [Turbinellidae]
electra, Columbarium (Peristarium), Columbariidae [Turbinellidae]
merope, Columbarium (Peristarium), Columbariidae [Turbinellidae]

Bequaert, J. C.
veracruzensis, Cerithidea pliculosa, Potamididae [Cerithideidae]

Bratcher, T. (see also under Cernohorsky, W. O.)

Bratcher, T. & R. D. Burch
allyni, Terebra, Terebridae
dorothyae, Terebra, ‘Terebridae
hancocki, Terebra, ‘Terebridae

Burch, R. D. (see also under Bratcher, T.)
betsyae, Hastula (Punctoterebra), Terebridae
craket, Duplicaria (Duplicaria), Terebridae
maryleeae, Hastula (Hastula), Terebridae

July 30, 1991 Bieler & Bradford

Burch, J. Q. & R. L. Burch
rejecta, Oliva, Olividae

Burch, R. L. (see under Burch, J. Q.)

Burgess, C. M.
mauiensis, Cypraea, Cypraeidae

Campbell, G. B. (see under Shasky, D. R.)

Cardin, C. & J. G. Walls
lenoreae, Phenacovolva (Subsimnia), Ovulidae

Cate, C. N.
lahainaensis, Volva (Phenacovolva), Ovulidae

Cernohorsky, W. O.
adamsianum, Vexillum (Costellaria), Costellariidae
crosniert, Cancilla scrobiculata, Mitridae
salisburyi, Mitrolumna, Turridae
salisburyi, Vexillum (Pusia), Mitridae [Costellariidae]
wolfe, Vexillum (Costellaria), Mitridae [Costellariidae]

Cernohorsky, W. O. & T. Bratcher
parkinsoni, Terebra, Terebridae

Clench, W. J.
finlayi, Murex (Murex), Muricidae
haysae, Thais floridana, Muricidae
trujilloi, Oliva, Olividae
Clench, W. J. & R. D. Turner
johnstone1, Melongena corona, Melongenidae [Buccinidae]
langi, Thais (Thaisella), Muricidae
riosi, Adelomelon (Weaveria), Volutidae

Cotton, B. C.
rosselli, Zoila, Cypraeidae

Dall, W. H.
atkana, Littorina sitkana, Littorinidae
major, Buccinum pemphigus, Buccinidae
vernicosum, Mangilia (Calliotectum), Pleurotomidae [Turridae]

Dall, W. H. in C. R. Orcutt & W. H. Dall
orcutti, Caecum, Caecidae

D’ Attilio, A. (see under Emerson, W. K.)

35

36 NEMOURIA No. 36

DuShane, H.
textimattum, Epitonium (Asperiscala), Epitoniidae

Emerson, W. K. & A. D’Attilio
dalh, Latiaxis (Babelomurex), Coralliophilidae [Muricidae]

Finlay, C. J. (see under Abbott, R. T.)

Finlay, H. J.
cuvierensis, Microvoluta, Volutomitridae

Ford, J.
blanesi, Olivella, Olividae
gaylordi, Olivella gracilis, Olividae
greegori, Cypraea cruenta, Cypraeidae
michaeh, Ocinebra, Muricidae

Harasewych, M. G.
ecphoroides, Fulgurofusus (Fulgurofusus), Turbinellidae
germonae, Tractolira, Volutidae
harrisae, Columbarium, Turbinellidae
tumor, Fulgurofusus (Peristarium), ‘Turbinellidae
xenismatis, Fulgurofusus (Histricosceptrum), Turbinellidae

Harasewych, M. G. & R. H. Jensen
guesti, Natica (Natica), Naticidae
guesti, Pterynotus (Pterynotus), Muricidae
lightbourni, Pterynotus (Pterynotus), Muricidae
radwini, Pterynotus (Pterynotus), Muricidae

Hedley, C.
perplicata, Voluta, Volutidae

Hoerle, R. C.
finlayi, Neritopsis, Neritopsidae

Howard, F. B.
aureola, Pyrene, Columbellidae

Hutton, F. W.
corticata, Acmaea, Acmaeidae

Iredale, T.
kellneri, Aulica, Volutidae

Jensen, R. H. (see under Harasewych, M. G.)

July 30, 1991 Bieler & Bradford

Keen, A. M.
fayae, Anachis (?Costanachis), Columbellidae

Klappenbach, 1964: 132.
teaguet, Olivancillaria, Olividae.

Lewis, H. (see also under Abbott, R. T-)
burgessi, Distorsio, Cymatiidae [Ranellidae]
habei, Distorsio constricta, Cymatiidae [Ranellidae]

Lyons, W. G.
barbarae, Brachycythara, Turridae
girardi, Cerodrillia, ‘Turridae
inglet, Oceanida, Eulimidae

Marcus, Ernst (see under Marcus, Eveline)

Marcus, Eveline & Ernst Marcus
vina, Assiminea succinea, Assimineidae

McGinty, T. L. (see also under Olsson, A. A.)
stegert, Daphnella, Turridae
thompson, Eudolium, Tonnidae

Mitchell-Tapping, H. J.
donmoorei, Caecum, Caecidae

Oliver, W. R. B.
kermadecensis, Haurakia, Rissoidae

Olsson, A. A.
stegert, Olivella (Olivella), Olividae

Olsson, A. A. & T. L. McGinty
elegans, Vitrinella (Striovitrinella), Vitrinellidae
marmosa, Olivella minuta, Olividae

Pease, W. H.
dispar, Neritina, Neritidae
rubra, Alcyna, Trochidae
Peile, A. J.
bermudensis, Rissoina, Rissoidae

Petit, R. E. (see also under Roth, B.)
darwini, Cancellaria, Cancellariidae

af

38 NEMOURIA No. 36

Petuch, E. J.
abrolhosensis, Acmaea (Collisella), Patellidae [Acmaeidae]
aphrodite, Conus, Conidae
baileyi, Oliva, Olividae
boholensis, Conus, Conidae
eccentricus, Zelippistes, Trichotropidae [Capulidae]
echinophilus, Africonus, Conidae
exquisita, Pseudocypraea, Ovulidae
fragilissimus, Conus, Conidae
katcherae, Vexillum (Costellaria), Costellariidae
kurzi, Guildfordia, Turbinidae
kurzt, Morum (Ontscidia), Cassidae [Harpidae]
lightbourni, Conus (Floraconus), Conidae
nereis, Conus, Conidae
nodosivaricosa, Agatrix (Olssonella), Cancellariidae
philippinensis, Axymene, Muricidae
zulu, Conus, Conidae

Pilsbry, H. A.
dalhana, Acmaea, Acmaeidae
floridana, Stenacme, Stenacmidae [Epitoniidae]
tokyoensis, Rissoa, Rissoidae

Powell, A. W. B.
ahiparana, Turritella (Zeacolpus), Turritellidae
bouvetia, Margarella, Trochidae
del, Penion benthicola, Buccinidae
kingae, Xenoturris, Turridae
macquariensis, Trophon, Muricidae
mauganuica, Austrodiaphana, Austrodiaphanidae [Diaphanidae]
nivosa, Ovtirissoa, Rissoidae
pseudanaloga, Puncturella, Fissurellidae
queenslandica, Leucosyrinx, Turridae
scotiana, Sinuber sculpta, Naticidae
signeyana, Chlanidota, Buccinulidae [Buccinidae]
tenerum, Parvaplustrum, Aplustridae [Hydatinidae]
venustula, Pareuthria, Buccinulidae [Buccinidae]
weavert, Turridrupa, Turridae
wormaldi, Columbarium (Coluzea), Columbariidae [Turbinellidae]
zebrina, Zediloma (Fractarmilla) corrosa, Trochidae

July 30, 1991 Bieler & Bradford

Rehder, H. A.
barnardi, Fusivoluta, Volutidae
boswellae, Volutocorbis, Volutidae
clarket, Fusivoluta, Volutidae
disparilis, Volutocorbts, Volutidae
floridanus, Microdochus, Rissoidae [Elachisinidae]
gracihor, Benthovoluta, Volutidae [Turbinellidae]

Roth, B. & R. E. Petit
masirana, Persicula, Marginellidae

Schilder, F. A.
summers, Bistolida (Blasicrura) pallidula, Cypraeidae

Schilder, F. A. & R. Summers
hesperina, Blasicrura coxeni, Cypraeidae

Schwengel, J. S.
koto, Lamellaria, Lamellariidae
mceginty1, Eubela, Turridae

Shasky, D. R.
berryi, Anachis, Columbellidae
maclean, Coralliophila, Coralliophilidae [Muricidae]
mazatlanica, Ruthia, Columbellidae
mendozana, Strombina (Cotonopsis), Columbellidae

Shasky, D. R. & G. B. Campbell
macleani, Calliostoma, Trochidae

Smith, A. G.
stilesi, Neptunea, Buccinidae

Smith, E. A.
huttonu, Trochus (Cantharidus), Trochidae

Snyder, M. A.
lightbourn, Fusinus, Fasciolariidae [Buccinidae]
martini, Latirus, Fasciolariidae [Buccinidae]

Stokes, A. J.
randall, Cymbiola, Volutidae

Summers, R. (see under Schilder, F. A.)

Suter, H.
lyalhana, Rissota (Alvania) plicata, Rissoidae

9

40 NEMOURIA

Test, A. R.
cona, Acmaea, Acmaeidae

Tippett, D. L.
brasiliana, Borsonia, Turridae

Turner, R. D. (see under Clench, W. J.)

Vanatta, E. G.
amosi, Metula, Buccinidae

Walls, J. G. (see also under Cardin, J.)
axelrodi, Conus, Conidae
clovert, Conus, Conidae
kerstitcht, Conus, Conidae
meyerl, Conus biliosus, Conidae
parvatus, Conus musicus, Conidae
reductasptralis, Conus nielsenae, Conidae
sandwichensis, Conus suturalis, Conidae
tribble1, Conus, Conidae
wittigl, Conus, Conidae

Weaver, C. S.
cloveriana, Lyria, Volutidae
daisyae, Volutoconus hargreavesi, Volutidae
dampteria, Amoria (Amoria), Volutidae
duponti, Festilyria, Volutidae

No. 36

July 30, 1991 Bieler & Bradford 41

LITERATURE CITED

Abbott, R. T. 1952. A New Terebra (hoffmeyert) from the Philippines. The
Nautilus, 65(3): 77-80, pl. 5.

— 1954. New Gulf of Mexico gastropods (Jerebra and Ocenebra). The Nautilus,
68(2): 37-44, pl. 2.

— 1958a. The Marine mollusks of Grand Cayman Island, British West Indies.
Monographs of the Academy of Natural Sciences of Philadelphia, 11:
1-138, pls. 1-5.

—— 1958b. The gastropod genus Assiminea in the Philippines. Proceedings of
the Academy of Natural Sciences of Philadelphia, 110: 213-278, pls. 15-25.

— 1971. Conus patae, a new Caribbean gastropod. The Nautilus, 85(2): 49-51.

— 1973. Acteon eloiseae, a new opisthobranch from Arabia. The Nautilus,
87(4): 91-92.

Abbott, R. T. & C. J. Finlay. 1979. Chicoreus cosmani, a new muricid gastropod
from the West Indies. The Nautilus, 93(4): 159-162 (issue erroneously °
labeled ‘“94(4)”’ in part).

Abbott, R. T. & H. Lewis. 1970. Cymatium boschi, new species from the Arabian
Sea. The Nautilus, 83(3): 86-88.

Adams, C. B. 1850. Descriptions of supposed new species of marine shells, which
inhabit Jamaica. Contributions to Conchology, No. 7: 109-123.

Bales, B. R. 1942. A new subspecies of Strombus raninus Gmelin. The Nautilus,
56(1): 18-19, pl. 4.

Barnard, K. H. 1963. Contributions to the knowledge of South African Marine
Mollusca. Part III. Gastropoda: Prosobranchiata: Taenioglossa. Annals of
the South African Museum, 47(1): 1-199.

— 1969. Contributions to the knowledge of South African Marine Mollusca.
Part VI. Supplement. Annals of the South African Museum, 47(4): 595-661,
pls. 1-2.

Bartsch, P. 1942. Some deep-sea Philippine volutids. The Nautilus, 56(1): 9-13,
pl. 2.

Bayer, F. M. 1971. New and unusual mollusks collected by R/V John Elliott
Pillsbury and R/V Gerda in the tropical western Atlantic. Biological results
of the University of Miami deep-sea expeditions. 79. Bulletin of Marine
Science, 21(1): 111-236 [also issued as pp. 111-236 of Bayer, F.M. &
G.L. Voss (eds.), Studies in tropical American mollusks, University of
Miami Press, Coral Gables, Florida, 1971].

Bequaert, J. C. 1942. Cerithidea and Batillaria in the western Atlantic. Johnsonia,
1(5): 1-11, incl. pls. 1-5.

Bratcher, T. & R. D. Burch. 1970. Five new species of Terebra from the Eastern
Pacific. The Veliger, 12(3): 295-300, pl. 44.

42 NEMOURIA No. 36

Burch, J. Q. & R. L. Burch. 1962. New species of Oliva from West Mexico.
The Nautilus, 75(4): 165-166, pl. 17.

Burch, R. D. 1965. New terebrid species from the Indo-Pacific Ocean and from
the Gulf of Mexico, with new locality records and provisional lists of species
collected in Western Australia and at Sabah, Malaysia (Mollusca: Gastro-
poda). The Veliger, 7(4): 241-253, pl. 31.

Burgess, C. M. 1967. A new Hawaiian Cypraea. The Nautilus, 81(1): 6-11, pl. 2.

Cardin, C. & J. G. Walls. 1980. A new false cowry from Pacific Panama
(Ovulidae: Simniini). The Pariah, 8: 1-2.

Cate, C. N. 1969. Two new species of the genus Volva Réding, 1798 (Ovulidae
Fleming, 1828). The Veliger, 11(4): 364-366, pl. 56.

Cernohorsky, W. O. 1970. New Mitridae and Volutomitridae. The Nautilus,
83(3): 95-104.

— 1976. The taxonomy of some Indo-Pacific Mollusca. Part 4. With
descriptions of new taxa and remarks on Nassarius coppingeri (Smith).
Records of the Auckland Institute and Museum, 13: 111-129.

— 1978. New species of Mitridae, Costellariidae and Turridae from the
Hawaiian Islands with notes on Mitra sphoni in the Galapagos Islands. The
Nautilus, 92(2): 61-67.

Cernohorsky, W. O. & T. Bratcher. 1976. Notes on the taxonomy of Indo-Pacific
Terebridae (Mollusca: Gastropoda), with description of a new species.
Records of the Auckland Institute and Museum, 13: 131-140.

Clench, W. J. 1927. A new subspecies of Thais from Louisiana. The Nautilus,
41(1): 6-8.

— 1938. A new species of Oliva from Santo Domingo, with notes on other
marine forms. The Nautilus, 51(4): 109-114, pl. 9.

— 1955. A new Murex from Matanzas, Cuba. Breviora, 44: 1-2, 1 pl.

Clench, W. J. & R. D. Turner. 1948. A new Thais from Angola and notes on
Thats haemastoma Linné. American Museum Novitates, no. 1374: 1-4.

Clench, W. J. & R. D. Turner. 1950. The western Atlantic marine mollusks
described by C. B. Adams. Occasional Papers on Mollusks, 1(15): 233-403,
incl. pls. 29-49.

Clench, W. J. & R. D. Turner. 1956. The family Melongenidae in the western
Atlantic. Johnsonia, 3(35): 161-188.

Clench, W. J. & R. D. Turner. 1964. The subfamilies Volutinae, Zidoninae,
Odontocymbiolinae and Calliotectinae in the western Atlantic. Johnsonia,
4(43): 129-180.

Coomans, H. E., R. G. Moolenbeek & E. Wils. 1982. Alphabetical revision of
the (sub)species in recent Conidae. 5. baccatus to byssinus, including Conus
brettinghami nomen novum. Basteria, 46: 3-67.

Coomans, H. E., R. G. Moolenbeek & E. Wils. 1986. Alphabetical revision of

July 30, 1991 Bieler & Bradford 43

the (sub)species in recent Conidae. 9. ebraeus to extraordinarius with the
description of Conus elegans ramalhoi, nov. subspecies. Basteria, 50: 93-150.

Cotton, B. C. 1948. Southern Australian Gastropoda Part II. Transactions of
the Royal Society of South Australia, 72(1): 30-32, pl. 1.

Dall, W. H. 1886. Contributions to the Natural History of the Commander
Islands. No. 6.—Report on Bering Island Mollusca collected by Mr. Nicholas
Grebnitzki. Proceedings of the Unites States National Museum, 9: 209-219.

— 1889. Scientific results of explorations by the U.S. Fish Commission
Steamer Albatross. No. VII.— Preliminary report on the collection of
Mollusca and Brachiopoda obtained in 1887-’88. Proceedings of the United
States National Museum, 12: 219-362, pls. 5-14.

— 1919. Descriptions of new species of Mollusca from the North Pacific in
the collection of the United States National Museum. Proceedings of the
United States National Museum, 56(2295): 293-371.

DuShane, H. 1977. Epitonium textimattum, a new gastropod from the west coast
of Mexico. The Nautilus, 91(3): 89-91.

Emerson, W. K. & A. D’Attilio. 1963. A new species of Latiaxis from the western
Atlantic (Mollusca, Gastropoda). American Museum Novitates, 2419: 1-9.

Finlay, H. J. 1926. A further commentary on New Zealand molluscan systematics.
Transactions and Proceedings of the New Zealand Institute, 57: 320-485,
pls. 18-23.

— 1930. Additions to the Recent molluscan fauna of New Zealand. No. 3.
Transactions and Proceedings of the New Zealand Institute, 61(2):
222-258, pls. 42-45.

Ford, J. 1888. Description of a new species of Ocinebra. Proceedings of the
Academy of Natural Sciences of Philadelphia, 1888: 188.

—— 1893. Description of a new form of Cypraea. The Nautilus, 6(10): 112-113.
— 1895. A new variety of Olivella. The Nautilus, 8(9): 103-104, pl. 2.
—— 1898. Description of a new species of Olivella. The Nautilus, 12(6): 66-67.

Harasewych, M. G. 1983a. A review of the Columbariinae (Gastropoda:
Turbinellidae) of the western Atlantic with notes on the anatomy and
systematic relationships of the subfamily. Nemouria, 27: 1-42.

—— 1983b. A new species of Columbarium (Gastropoda: Muricacea) from off
eastern Australia. The Nautilus, 97(1): 28-29.

— 1987. Tractolira germonae, a new abyssal Antarctic volute. The Nautilus,
101(1): 3-8.

Harasewych, M. G. & R. H. Jensen. 1979. Review of the subgenus Prerynotus
(Gastropoda: Muricidae) in the western Atlantic. Nemouria, 22: 1-16.

Harasewych, M. G. & R. H. Jensen. 1984. Natica guesti, a new naticid gastropod
from the western Atlantic. The Nautilus, 98(3): 99-101.

44 NEMOURIA No. 36

Hedley, C. 1902. A new Australian volute. Records of the Australian Museum,
4(7): 309.

Hoerle, R. C. 1974. A Recent Caribbean species of the genus Neritopsis (Mollusca:
Gastropoda). Tulane Studies in Geology and Paleontology, 11: 103-104.

Howard, F. B. 1963. Description of a new Pyrene from Mexico (Mollusca—
Gastropoda). Santa Barbara Museum of Natural History Occasional Papers,
no. 7: 1-12, incl. 2 pls.

Hutton, F. W. 1880. Manual of the Mollusca. A systematic and descriptive
catalogue of the marine and land shells, and of the soft mollusks and
Polyzoa of New Zealand and the adjacent islands. Colonial Museum and
Geological Survey Department, Wellington, 224 pp.

[ICZN] International Commission on Zoological Nomenclature. 1985. Interna-
tional Code of Zoological Nomenclature (3rd ed.). xx + 338 pp.;
International Trust for Zoological Nomenclature, etc.

Iredale, T. 1957. Another Australian volute. Proceedings of the Royal Zoological
Society of New South Wales, 1955-1956: 91-92.

Kay, E. A. & W. J. Clench. 1975. A biobibliography of William Harper Pease,
malacologist of Polynesia. Nemouria, 16: 1-50.

Keen, A. M. (with the assistance of J. H. McLean). 1971. Sea Shells of Tropical
West America. Marine Mollusks from Baja California to Peru. Second
edition. Stanford University Press, Stanford, California, 1064 pp.

Klappenbach, M. A. 1964. A new species of Olivancillaria from Uruguay and
Brazil. The Nautilus, 77(4): 132-134, pl. 8.

Lewis, H. 1972. Notes on the genus Distorsio (Cymatiidae) with descriptions of
new species. The Nautilus, 86(2-4): 27-50.

Lyons, W. G. 1972. New Turridae (Gastropoda: Toxoglossa) from south Florida
and the eastern Gulf of Mexico. The Nautilus, 86(1): 3-7.

— 1978. Status of the genus Oceanida DeFolin (Gastropoda: Eulimidae), with
a description of a new species. Proceedings of the Biological Society of
Washington, 91(2): 539-545.

Marcus, Eveline & Ernst Marcus. 1965. On Brazilian supratidal and estuarine
snails. Boletim da Faculdade de Filosofia, Ciéncias e Letras, Universidade
de Sao Paulo, 287, Zoologia no. 25: 19-82, pls. 1-10.

McGinty, T. L. 1955. New marine mollusks from Florida. Proceedings of the
Academy of Natural Sciences of Philadelphia, 107: 75-86, pls. 1, 2.
Mitchell-Tapping, H. J. 1979. The Caecidae (Gastropoda: Rissoacea) of Water
Island, U.S. Virgin Islands, with a new species. The Nautilus, 93(2-3):

103-105.

Olsson, A. A. 1956. Studies on the genus Olivella. Proceedings of the Academy

of Natural Sciences of Philadelphia, 108: 155-225, pls. 8-16.

Olsson, A. A. & T. L. McGinty. 1958. Recent marine mollusks from the

July 30, 1991 Bieler & Bradford 45

Caribbean coast of Panama with the description of some new genera and
species. Bulletins of American Paleontology, 39(177): 1-59, pls. 1-5.

Oliver, W. R. B. 1915. The Mollusca of the Kermadec Islands. Transactions
and Proceedings of the New Zealand Institute, 47: 509-568, pls. 9-12.

Orcutt, C. R., with comments by W. H. Dall. 1885. Notes on the mollusks of
the vicinity of San Diego, Cal., and Todos Santos Bay, Lower California.
Proceedings of the Unites States National Museum, 8: 534-552.

Pease, W. H. 1860. Descriptions of forty-seven new species of shells from the
Sandwich Islands, in the collection of Hugh Cuming. Proceedings of the
Zoological Society of London, 28: 431-438 [‘‘? 1861” teste Kay & Clench,
1975: 47].

— 1867. Descriptions of sixty-five new species of marine Gasteropodae,
inhabiting Polynesia. American Journal of Conchology, 3: 271-279, pls.
23, 24.

Peile, A. J. 1926. The Mollusca of Bermuda. The Proceedings of the
Malacological Society of London, 17(2-3): 71-98.

Petit, R. E. 1970. Notes on Cancellariidae (Mollusca: Gastropoda)-II. Tulane
Studies in Geology and Paleontology, 8(2): 83-88.

Petuch, E. J. 1975. Two new cone species from Senegal, West Africa. The
Veliger, 18(2): 180-183, 1 pl.

— 1979a. Twelve new Indo-Pacific gastropods. Nemouria, 23: 1-21.

— 1979b. New gastropods from the Abrolhos Archipelago and reef complex,
Brazil. Proceedings of the Biological Society of Washington, 92(3): 510-526.

— 1986. The Austral-African conid subgenus Floraconus Iredale, 1930, taken
off Bermuda (Gastropoda: Conidae). Proceedings of the Biological Society
of Washington, 99(1): 15-16.

Pilsbry, H. A. 1891. Monographs of the Acmaeidae, Lepetidae, Patellidae and
Titiscaniidae. Manual of Conchology, 13: 1-195, pls. 1-74.

—— 1904. New Japanese marine Mollusca: Gastropoda. Proceedings of the
Academy of Natural Sciences of Philadelphia, 56: 3-37, pls. 1-6.

— 1945. Stenacme floridana, an American member of the Amphibolacea. The
Nautilus, 58(4): 112-116, pls. 5, 6.

Ponder, W. F. & A. Warén. 1988. Classification of the Caenogastropoda and
Heterostropha-—a list of the family-group names and higher taxa. Malacologi-
cal Review, Suppl. 4: 288-326.

Powell, A. W. B. 1927. Mollusca from twenty-three fathoms off Ahipara, N. Z.
Transactions and Proceedings of the New Zealand Institute, 58(3):
295-300, pl. 34.

—— 1946. New species of New Zealand Mollusca from the South Island, Stewart

Island, and Chatham Islands. Records of the Auckland Institute and
Museum, 3(2): 137-144, pls. 11, 12.

46 NEMOURIA No. 36

— 1951. Antarctic and Subantarctic Mollusca: Pelecypoda and Gastropoda
collected by the ships of the Discovery Committee during the years
1926-1937. Discovery Reports, 26: 47-196, pls. 5-10.

— 1952. New Zealand molluscan systematics, with descriptions of new species,
part 1. Records of the Auckland Institute and Museum, 4(3): 169-185,
pls. 35, 36.

— 1957. Mollusca of the Kerguelen and Macquarie Islands. B.A.N.Z. [British,
Australian and New Zealand] Antarctic Research Expedition, 1929-1931,
Reports, Series B, 6(7): 107-150, incl. pls. 1-3.

— 1964. The family Turridae in the Indo-Pacific Part 1. The subfamily
Turrinae. Indo-Pacific Mollusca, 1(5): 227-346.

— 1967. The family Turridae in the Indo-Pacific Part la. The subfamily
Turrinae concluded. Indo-Pacific Mollusca, 1(7): 409-432.

— 1969. The family Turridae in the Indo-Pacific. Part 2. The subfamily
Turriculinae. Indo-Pacific Mollusca, 2(10): 215-416.

— 1971. New Zealand molluscan systematics with descriptions of new species:
Part 7. Records of the Auckland Institute and Museum, 8: 209-228.

— 1979. New Zealand Mollusca. Marine, land and freshwater shells. xiv +
500 pp.; Collins; Auckland, Sydney, London.

Rehder, H. A. 1943. New marine mollusks from the Antillean region. Proceedings
of the United States National Museum, 93(3161): 187-203, pls. 19-20.

— 1967. A new genus and two new species in the families Volutidae and
Turbinellidae (Mollusca: Gastropoda) from the western Pacific. Pacific
Science, 21(2): 182-187.

— 1969. Volutocorbis and Fusivoluta, two genera of deepwater Volutidae from
South Africa. The Veliger, 11(3): 200-209, pls. 40-43.

Robertson, R. & K. Oyama. 1958. The family Stenacmidae. The Nautilus, 72(2):
68-69.

Roéckel, D. 1979-1988. Die Familie Conidae. 34 series with 606 plates. Club
Conchylia, Darmstadt.

Roth, B. & R. E. Petit. 1972. A new species of Persicula from the Arabian Sea
(Marginellidae). The Nautilus, 86(2-4): 80.

Schilder, F. A. 1958. Uber drei seltene Cypraeacea. Archiv fiir Molluskenkunde,
87(1/3):. 81-87.

Schilder, F. A. & R. Summers. 1963. A further note on Blasicrura coxeni (Cox).
The Cowry, 1(5): 68-69, pls. VIII, XI.

Schwengel, J. S. 1943. New marine shells from Florida. The Nautilus, 56(3):
75-78, pl. 7.

— 1944. A new Florida Lamellaria. The Nautilus, 58(1): 17-18, pl. 1.
Shasky, D. R. 1970. New gastropod taxa from tropical western America. The

July 30, 1991 Bieler & Bradford 47

Veliger, 13(2): 188-195, pl. 1.

Shasky, D. R. & Campbell, G. B. 1964. New and otherwise interesting species
of mollusks from Guaymas, Sonora, Mexico. The Veliger, 7(2): 114-120,
pls: 21, 22;

Smith, A. G. 1968. A new Neptunea from the Pacific Northwest. The Veliger,
11(2): 117-120, pl. 14.

Smith, E. A. 1876. A list of marine shells, chiefly from the Solomon Islands,
with descriptions of several new species. The Journal of the Linnean
Society, Zoology, 12: 535-567, pl. 30.

Snyder, M. A. 1984. Fusinus lightbourni (Gastropoda: Fasciolariidae), a new
species from Bermuda. The Nautilus, 98(1): 28-30.

—— 1988. Latirus martini (Gastropoda: Fasciolariidae), a new species from
Honduras. The Nautilus, 102(2): 54-55.

Stokes, A. J. 1961. A new species of volute from Undine Reef, Queensland [:]
Cymbiola randalli sp. nov. Royal Society of South Australia, Malacological
Section, Publication No. 16: [3].

Suter, H. 1898. Revision of the New Zealand Rissoidae. Proceedings of the
Malacological Society of London, 3: 2-8.

Test, A. R. (G.). 1945. Description of new species of Acmaea. The Nautilus,
58(3): 92-96.

Thiele, J. 1929-1935. Handbuch der systematischen Weichtierkunde. Gustav
Fischer, Jena [Vol. 1(1): 1-376, 1929; 1(2): 377-778, 1931; 2(3): 779-1022,
1934; 2(4): i-iv, 1023-1154 and i-vi for vol. 1; 1935].

Thompson, F. G. 1982. The Recent mollusk collections of North America-II.
ASC [Association of Systematics Collections] Newsletter, 10(4): 43-45.

Tippett, D. L. 1983. A new sinistral turrid from Brazil (Gastropoda: Turridae).
The Nautilus, 97(4): 135-138.

Vanatta, E. G. 1913. Descriptions of new species of marine shells. Proceedings
of the Academy of Natural Sciences of Philadelphia, 65: 22-27, pl. 2.

Walls, J. G. 1977. Two new cones from the western Pacific. The Pariah, 1: 1-3.
—— 1978. Diagnoses of four new cones (Mollusca: Conidae). The Pariah, 2: 1-7.

—— [1979a]. Cone shells. A Synopsis of the living Conidae. 1011 (+ 9) pp.,
T.F.H. Publications Inc. Ltd., Hong Kong [March 1979].

—— 1979b. Three new Indian Ocean cones (Mollusca: Conidae). The Pariah,
5: 1-6 [30 April 1979].

Weaver, C. S. 1960. Hawaiian scientific expedition fnds [sic] rare Western
Australian volutes. Hawaiian Shell News, 8(12): 1, 3, pl. on p. 1.

— 1963a. Redescription of Amoria dampieria Weaver, 1960 (Gastropoda:
Volutidae). Journal of the Malacological Society of Australia, 7: 28-31, pls.
5-6.

48 NEMOURIA No. 36

— 1963b. A new species of Lyria from Ceylon (Gastropoda: Volutidae). Journal
of the Malacological Society of Australia, 7: 31-32, pl. 6.

— 1967. A new subspecies of Volutoconus hargreavest (Angas, 1872) from
central Western Australia (Gastropoda: Volutidae). The Veliger, 9(3):
301-304, pl. 41.

— 1968. A new marine mollusk from Mozambique in the genus Festilyria
Pilsbry and Olsson, 1954 (Gastropoda: Volutidae). The Veliger, 10(4):
442-445, pl. 60.

Weaver, C. S. & J. E. duPont. 1970. The living volutes. A monograph of the
Recent Volutidae of the world. Delaware Museum of Natural History
Monograph Series No. 1, Greenville, xv + 375 pp., incl. 79 pls.

Wenz, W. 1938-1944. Gastropoda. Teil 1: Allgemeiner Teil und Prosobranchia.
Handbuch der Paldozoologie, 6(1): i-xii, 1-1639. Borntraeger, Berlin [pp.
1-240, March 1938; 241-480, October 1938; 481-720, July 1939; 721-960,
August 1940; 961-1200, October 1941; 1201-1506, October 1943; 1507-
1639, i-xii, November 1944].

Zilch, A. 1959-1960. Gastropoda von Wilhelm Wenz. Teil 2. Euthyneura.
Handbuch der Paldozoologie, 6(2): i-xii, 1-835, 2515 figs. Borntraeger,
Berlin [pp. 1-200, 17 July 1959; 201-400, 25 November 1959; 401-600,
30 March 1960; 601-835, i-xii, 15 August 1960].

ACKNOWLEDGMENTS

We thank DMNH curatorial staff and volunteers, especially Dr.
Albert F. Chadwick, Mr. Russell H. Jensen and Ms. Wendeline K.
Knoedler, for technical assistance and for providing background informa-
tion on earlier collection accessions. Dr. R. Tucker Abbott (Melbourne,
Florida), Dr. Arthur E. Bogan (ANSP), Dr. George M. Davis (ANSP),
Dr. M. G. Harasewych (USNM), Mr. John R. H. Lightbourn (Bermuda),
Mr. William G. Lyons (Florida Marine Research Institute, St. Peters-
burg), Ms. Paula M. Mikkelsen (Harbor Branch Oceanographic Museum,
Ft. Pierce, Florida), Dr. Robert Robertson (ANSP), and Dr. Dieter
Réckel (Eberbach, Germany) kindly provided information on type
material and/or literature.

The manuscript was reviewed by Drs. William K. Emerson
(American Museum of Natural History, NY) and M. G. Harasewych
(National Museum of Natural History, DC).

a
t eS
*.
=_
=
~ =

—

ao

Founding Editor Managing Editor
JOHN E. DU PONT RUDIGER BIELER

Consulting Editors

WILLIAM K. EMERSON, American Museum of Natural History, New
York, NY.

M. G. HARASEWYCH, National Museum of Natural History, Smithsonian
Institution, Washington, DC.

KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh,
PA.

RICHARD W. THORINGTON, JR.,; National Museum of Natural History,
Smithsonian Institution, Washington, DC.

Published by the

DELAWARE MUSEUM OF NATURAL HISTORY
4840 Kennett Pike, Wilmington,
Delaware 19807, U.S.A.

NEMOURIA

Occasional Papers of the Delaware Museum of Natural History

NUMBER 37 SEPTEMBER 30, 1991

REDISCOVERY OF MARGINELLONA GIGAS
(MARTENS, 1904), WITH NOTES ON
THE ANATOMY AND SYSTEMATIC POSITION OF
THE SUBFAMILY MARGINELLONINAE
(GASTROPODA: MARGINELLIDAE)

M. G. Harasewych! and Yuri I. Kantor?

ABSTRACT. Based on anatomical data, the genus Sigaluta Rehder, 1967, and its
type species S. pratasensis Rehder, 1967, originally described in the Volutidae,
are synonymized with the genus Marginellona Martens, 1904, and its type species
M. gigas (Martens, 1904), of the family Marginellidae. The originally questionable
inclusion of this species in Marginellidae is supported by additional anatomical
characters. The subfamily Marginelloninae, type genus Marginellona, differs from
other marginellids not only in its much larger size and distinctive radular
morphology, but also in having a relatively unmodified neogastropod alimentary
system with a large valve of Leiblein, and a large gland of Leiblein that lacks a
separate duct to the buccal cavity.

Key words: Marginellidae; Marginellona; Sigaluta; South China Sea; anatomy.

INTRODUCTION

The genus Sigaluta and its type species S. pratasensis were described
from two shells and provisionally assigned to the volutid subfamily
Cymbiinae on the basis of general conchological similarities (Rehder,
1967). An additional, larger specimen, also without preserved animal,
was reported by Rehder (1970). Weaver and duPont (1970: 99) followed
Rehder’s subfamilial assignment, but remarked that Sigaluta pratasensis
““, . . does not resemble any other species in the family Volutidae.”

1Department of Invertebrate Zoology, National Museum Serial Publication
of Natural History, Smithsonian Institution, Washington, ISSN 0085-3887
DC 20560, U.S.A.

2A. N. Severtzov Institute of Animal Evolutionary

Morphology and Ecology, Academy of Sciences of the U.S.S.R.,

Lenin Avenue, 33, Moscow 117071, U.S.S.R.

2 NEMOURIA No. 37

Rokop (1972) assigned a second species, from abyssal depths off Baja
California, to this genus, also on the basis of shell characters.

During our recent visit to the Zoological Institute in Leningrad,
Dr. Boris Sirenko brought to our attention several specimens of Sigaluta
pratasensis, three with preserved animal. Preliminary dissections revealed
this species to have a marginellid radula. A survey of the literature on
Marginellidae disclosed that Sigaluta pratasensis is a synonym of Marginel-
lona gigas (Martens, 1904).

As Marginellona gigas is the type species of the type genus of the
small and poorly known subfamily Marginelloninae Coan, 1965, we take
this opportunity to describe the anatomy of this unusual marginellid, to
compare it with that of other marginellids, and to discuss its relationship
within the family Marginellidae. The systematics of Sigaluta cukri Rokop,
1972, is discussed in Harasewych & Kantor (1991).

MATERIALS AND METHODS

In addition to the holotype and paratype of Sigaluta pratasensis in
the collections of the National Museum of Natural History, Smithsonian
Institution (USNM), we examined six specimens in the collections of
the Zoological Institute, Leningrad (ZIN). Anatomical data is based on
gross dissections of three of these specimens, a large, mature male (fig.
3), a small female (fig. 4), and an immature male (fig. 5). The
terminology for shell ultrastructure follows Hedegaard (1990).

SYSTEMATICS

Family MARGINELLIDAE Fleming, 1828

Subfamily MARGINELLONINAE Coan, 1965
The subfamily was proposed by Coan (1965: 186) to include the

monotypic genera Marginellona Martens, 1904, and Afrivoluta Tomlin,

1947, and was characterized by having unusually large (>4 cm) shells

and short radulae consisting of extremely broad rachidian teeth with

numerous (60-90) cusps. The unique radular morphology has been
termed “Type 7 radula’” by Coovert (1989: 33). The two species
comprising this subfamily have a Type 1 animal in the terminology of

Coovert (1987: 19).

Genus Marginellona Martens, 1904.

Marginella (Marginellona) Martens, 1904: 108-109, pl. 5, fig. 16; Thiele,
1904: 170, pl. 9, fig. 64; Tomlin, 1917: 268 [type species: Marginella
(Marginellona) gigas Martens, 1904, by monotypy].

Marginellona Martens, 1903 [sic], Thiele, 1929: 355-356, fig. 431; Wenz,
1943: 1380, fig. 3903.

September 30, 1991 Harasewych & Kantor 3

Marginelloma Cossmann, 1906: 225 [error in spelling].
Sigaluta Rehder, 1967: 182-183, text-figs. 1-4 (type species: Sigaluta
pratasensis Rehder, 1967, by original designation).

Martens (1904) observed that his taxon had affinities to the
Volutidae, but included it in the Marginellidae on the basis of radular
and shell morphology. Although Marginellona was originally described
as a subgenus of Marginella, Thiele (1929: 355-356) considered it to be
one of but three genera within Marginellidae.

Marginellona gigas (Martens, 1904)
Figures 1-19, Table 1

Marginella (Marginellona) gigas Martens, 1904: 108-109, pl. 5, fig. 16;
Thiele, 1904: 170, pl. 9, fig. 64; Tomlin, 1917: 268.
Marginellona gigas Martens, Thiele, 1929: 356; Wenz, 1943: 1380, fig.
3903.
Sigaluta pratasensis Rehder, 1967: 182-183, text-figs. 1-4.
Description: Shell (Figs. 2-5, Table 1) to 157 mm, thin, translucent,
porcellaneous, narrowly ovate. Protoconch of 2-2 rapidly expanding
(diameter 0.5 mm to 11.5 mm in 2 whorls), smooth, conical, glassy
whorls, deflected from coiling axis of teleoconch by up to 15°. Transition
to teleoconch abrupt, marked by growth line (Fig. 6, t), and accompanied
by formation of thin parietal callus (Fig. 6, c). Teleoconch with up to
3 smooth, inflated, convex, rapidly descending whorls. Suture with
abutting whorls. Shell surface smooth, glazed, lacking spiral and axial
sculpture. Aperture ovate, narrow posteriorly, broad anteriorly. Outer lip
smooth. Inner lip smooth, with thin, whitish inductural overglaze in
some specimens. Columella with single sharp, axially oriented columellar
fold (Fig. 3, cf) and sharp siphonal fold (Fig. 3, sf) of nearly equal
magnitude. Outer shell surface uniformly tan to greenish-tan; aperture
darker brown.
Ultrastructure: Shell composed of five crystalline layers (Fig. 7). Innermost
layer (Figs. 7, 9, i), thin (to 65 wm), of simple prismatic crystals.
Subsequent layer (Figs. 7, 9, t) thickest (to 300 um), composed of linear
simple crossed lamellar crystals oriented perpendicular to growing edge.
Next outer layer (Figs. 7, 8, c) nearly as thick (to 250 um), orthogonally
oriented, with faces of linear simple crossed lamellar crystals parallel to
growing edge. Next outer layer (Figs. 7, 8, p) thin (to 20 um), with
intersected crossed platy structure. Outermost layer (Fig. 8, 0) thin (to
70 um), again of simple prismatic crystals.
External anatomy: Soft parts of largest and only complete specimen (Figs.
3, 14) comprising 2 2 whorls, excluding foot. Mantle cavity spanning

4 NEMOURIA No. 37

Figure 1. Marginellona gigas (Martens). Original illustration of holotype, repro-
duced at 0.9 X.

Figure 2. Holotype of Sigaluta pratasensis Rehder, 1967, apertural, lateral and
dorsal views. W. of Pratas Reef, South China Sea, 20°37'N, 115°43’E, in 380
m, gray mud and sand bottom, 0.9 X.

September 30, 1991 Harasewych & Kantor 5

Figures 3-6. Marginellona gigas (Martens). 3. ZIN 44141, E. of Phan Thiet,
Vietnam, South China Sea, 10°41'08”N, 109°53'08”E, in 495-500 m, 6, 0.85
X. 4. ZIN 56353, E. of Phan Ly, Vietnam, South China Sea, 11°10'00"N,
110°10’00”E, in 1280-1000 m, mud, @, 0.85 XK. 5. ZIN 45513, E. of Phan Ly,
Vietnam, South China Sea, 11°09’06”N, 110°02'00”E, in 700 m, immature ¢,
0.85 X. 6. Protoconch of specimen in Fig. 5, 2.6 X.

c, parietal callus; cf, columellar fold; sf, siphonal fold; t, transition from
protoconch to teleoconch.

6 NEMOURIA No. 37

Figures 7-9. Shell ultrastructure of Marginellona gigas. 7. Fracture surface parallel
to growing edge, '% whorl behind aperture, circled region in Fig. 3. Scale bar =
50 um. 8. Enlargement, showing outer surface of shell. Scale bar = 20 um. 9.
Enlargement, showing inner surface of shell. Scale bar = 20 um.

c, colabrally oriented crossed lamellar crystals; i, innermost layer of simple
prismatic crystals; 0, outermost layer of simple prismatic crystals; p, layer with
crossed platy structure; t, transversely oriented crossed lamellar crystals.

September 30, 1991 Harasewych & Kantor 7

Figures 10-13. Radular morphology. 10. Radular tooth of holotype of
Marginellona gigas, redrawn from Thiele (1904: pl. 9, fig. 64). Scale bar = 500
Um. 11-13. Radula of specimen in Fig. 3. 11. Distal end of radula. Scale bar
= 500 um. 12. Region along central portion of tooth. Scale bar = 50 um. 13.
Region along right margin of tooth. Scale bar = 100 um.

8 NEMOURIA No. 37

—

Figures 14-16. External anatomical features of Marginellona gigas. 14. Shell-less |
animal from specimen in Fig. 3, dorsal view, 0.65 X. 15. Dorsal view of head, |
1.0 X. 16. Ventral view of head, 1.0 X. |

ag, anal gland; c, channel along mid-line of head; ct, ctenidium; dg, digestive
gland; em, efferent margin of mantle cavity; f, foot; hg, hypobranchial gland; k,
kidney; Ip, lateral lappets; mg, anterior mucus gland; os, osphradium; p, penis;

r, rectum; rh, rhynchostome; s, siphon; t, tentacle; te, testis; tr, transverse ridge. |

September 30, 1991 Harasewych & Kantor 9

19

Figure 17-19. Anatomical features of Marginellona gigas. 17. Alimentary system,
0.66 X. 18. Female pallial oviduct, 1.1 X. 19. Male pallial gonoduct, 0.9 X.

a, anus; ae, anterior esophagus; ag, anal gland; al, albumen gland; bc, bursa
copulatrix; bp, buccal pouch; bu, buccal cavity; cp, capsule gland; dgL, duct of
gland of Leiblein; dsg, duct of salivary gland; fo, female opening; gL, gland of
Leiblein; int, intestine; od, oviduct; p, penis; pap, penial papilla; pe, posterior
esophagus; pr, prostate; r, rectum; rs, receptaculum seminis; sg, salivary glands;
Sl, sperm ingesting gland; sto, stomach; vd, vas deferens; vL, valve of Leiblein.

10 NEMOURIA No. 37

Table 1. Measurements of shells and radulae of specimens of Marginel-
— lona gigas in Figs. 3-5. All measurements in mm.

Specimen in Figure no. 3 4 5
Shell length 157.0 68.3 B12
Aperture length 109.9 51.6 30.7
No. protoconch whorls 23 24 Dae
No. teleoconch whorls 27/4 1% 1/16
Radula length 10.1 5.3 3.4
Radula width 25 1.6 1.0
No. rows teeth 80 56 44
No. cusps/tooth* 83-85 58-59 69-71

* Number varies among teeth within the same radula.

approximately 2 whorl; kidney (Fig. 14, k) % whorl; digestive gland
(Fig. 14, dg) and testis (Fig. 14, te) 1 *%4 whorl. Foot (Fig. 14, f)
extremely large (~ 1.2 Xaperture length), broad (L/W = 1.0), with deep
medial indentation along anterior edge, tapering posteriorly. Anterior
mucus gland (Fig. 14, mg) deep. Dorsal surface of foot uniformly colored
in preserved specimens, rugose in large specimen, smoother in smaller
specimens. Operculum absent. Sole of foot thin, glandular. Tentacle
bases fused ventrally, producing deep, tubular channel along dorsal
mid-line of head (Fig. 15, c). Ventral surface of head with transverse
ridge (Fig. 16, tr) anterior to rhynchostome (Fig. 16, rh). Proximal region
of head with large, triangular lateral lappets (Fig. 15, Ip). Eyes absent.
Tentacles (Fig. 15, t) long, laterally directed. Siphon (Figs. 14,15, s)
fused to head just left of posterior margin of mid-line channel. Siphon
bifurcated in largest specimen, possibly due to injury; simple, tubular in
smaller two specimens.

Mantle cavity: Mantle cavity spanning approximately '/2 whorl, with right
side substantially foreshortened due to posterior displacement of its
efferent margin (Fig. 14, em). Arrangement of mantle cavity organs as
in generalized muricacean neogastropod (Ponder, 1973: fig. 5). Mantle
edge thickened, smooth, particularly broad and thick along columellar
margin. Osphradium (Fig. 14, os) dark greenish-brown, half as wide and
77, as long as ctenidium (Fig. 14, ct). Hypobranchial gland (Fig. 14, hg)
of parallel, deep, pendent folds. Pallial gonoduct and rectum short. Anal
gland (Figs. 14, 17, a) large.

September 30, 1991 Harasewych & Kantor 11

Alimentary system: Proboscis short, broad, pleurembolic, extending through
rhynchostome (Fig. 16, rh) at base of head. Proboscis retractor muscles
attached to dorsal and left walls of asymmetrical cephalic hemocoel.
Mouth large, leading to buccal cavity (Fig. 17, bu). Buccal cavity joined
dorsally by anterior esophagus (Fig. 17, ae), ventrally by buccal pouch
(Fig. 17, bp). Buccal pouch, containing buccal mass, extending nearly
to rear of retracted proboscis. Radula (Figs. 11-13, Table 1) short (3-10
mm), broad (1.0-2.5 mm), uniserial, with 44-80 asymmetrical teeth.
Teeth comb-like, with slightly curved basal plates that give rise to
numerous (58-85) short cusps. Cusps shorter at margins (Fig. 13),
irregular in thickness and occasionally fused along central portions of
teeth (Fig. 12). Anterior esophagus thin, muscular, extending from
buccal pouch to large, muscular valve of Leiblein (Fig. 17, vL). Posterior
to valve of Leiblein, esophagus doubling in diameter and becoming
sacculate, further expanding posterior to duct from gland of Leiblein
before constricting to original diameter at rear of cephalic hemocoel.
Gland of Leiblein (Fig. 17, gL) dark brown, extremely large, sacculate,
broad anteriorly, tapering posteriorly, with large central lumen, joined
to mid-esophagus posterior to nerve ring by thick, glandular stalk with
narrow duct (Fig. 17, dgL). Salivary glands (Fig. 17, sg) paired,
composed of extremely long, highly convoluted and intertwined tubules,
situated dorsal to the gland of Leiblein in expanded right anterior portion
of cephalic hemocoel. Ducts of salivary glands (Fig. 17, dsg) running
anteriorly along floor of proboscis, emptying into buccal cavity laterally.
Posterior esophagus reflected dorsally before joining stomach (Fig. 17,
sto). Stomach simple, sack-like, with ducts from digestive gland joining
to form single broad opening. Intestine (Fig. 17, int) short, narrow,
running anteriorly from stomach to form rectum (Fig. 17, r). Rectum
more than doubled in diameter alongside kidney, expanding further upon
entering mantle cavity. Anal gland (Fig. 17, ag) extremely large,
cavernous in large male, disproportionally smaller in smaller specimens,
spanning the pallial portion of the rectum, entering rectum by single
duct near anus.

Female reproductive system: [Only a single, poorly preserved pallial oviduct
was available for examination. The functions of the two glandular
structures between the capsule and albumen glands are surmised on the
basis of positional homologies with the pallial oviduct of Volvarina
taeniolata’ Mérch, 1860 (Fretter, 1976: fig. 2A) and will require
verification when additional specimens become available.] Oviduct (Fig.
18, od) joining albumen gland (Fig. 18, al) at rear of mantle cavity.
Pallial oviduct joined by sperm ingesting gland (?) (Fig. 18, si), seminal
receptacle (?) (Fig. 18, rs) before expanding into broad capsule gland

12 NEMOURIA No. 37

(Fig. 18, cp). Bursa copulatrix (Fig. 18, bc) large, situated ventral to
anus. Female opening (Fig. 18, fo) along lateral wall of bursa copulatrix.

Male reproductive system: Testis large, tan colored, lining anteriormost !/3
of digestive gland, broadest adjacent to rear wall of kidney. Testicular
duct convoluted along kidney, passing adjacent to pericardium before
entering mantle cavity. Prostate (Fig. 19, pr) large, glandular, not joined
to rectum by connective tissue, with slit-like opening extending along
posterior 3/5 of length. Vas deferens (Fig. 19, vd) descending abruptly,
running anteriorly to base of penis, surrounded by layer of muscle beneath
mantle floor. Penis (Fig. 19, p) long, broad, dorsoventrally compressed,
with terminal papilla (Fig. 19, pap) along right distal edge.

Type Locality: W. of Sombrero Channel, Nicobar Islands, Indian
Ocean, 07°48’N, 92°07’E, in 805 m, coarse sand.

Material examined: USNM 237018 [holotype Sigaluta pratasensis],
54.1 mm, USNM 637251 [paratype S. pratasensis] 61.1 mm, both from
W. of Pratas Reef, South China Sea, 20°37'N, 115°43’E, in 380 m, gray
mud and sand bottom, U.S.B.F. Albatross I, sta. 5301; ZIN 44141, E.
of Phan Thiet, Vietnam, South China Sea, 10°41'08”’N, 109°53'08”E,
in 495-500 m, R/V Odessey, trawl no. 55, 1 male specimen, 157.0 mm;
ZIN 45513, E. of Phan Ly, Vietnam, South China Sea, 11°09'06’N,
110°02’00”E, in 700 m, R/V Odessey, trawl no. 52, 1 immature male
specimen, 37.2 mm; ZIN 56351, E. of Phan Thiet, Vietnam, South
China Sea, 11°01'00”N, 109°55’00”E, in 460 m, mud, R/V Odessey, 2
shells, 98.5 mm, 106.9 mm; ZIN 56352, E. of Ba Ria, Vietnam, South
China Sea, 10°40’08”N, 110°03’00’E, in 760-800 m, mud, R/V Odessey,
1 shell, 108.8 mm; ZIN 56353, E. of Phan Ly, Vietnam, South China
Sea, 11°10’00”N, 110°10'00”E, in 1280-1000 m, mud, R/V Odessey,
trawl no. 51, 1 female specimen, 68.3 mm.

Distribution: This species has been collected in the eastern Indian
Ocean (Nicobar Islands) and on the upper continental slope along the
western margin of the South China Sea. The bathymetric range is
380-1000 m, with a mean station depth (n=7) of 661 m.

Ecology: Little is known of the ecology of this species other than it
inhabits substrates ranging from coarse sand to fine mud. Gut contents
of three dissected specimens were not identifiable.

Remarks: Marginellona gigas is the largest known marginellid. The largest
specimen of Marginellona gigas recorded in this study measures 157 mm
in length, significantly larger than the published record size of 130.9
mm for Afnvoluta pringlei (Wagner & Abbott, 1990: 80-039).

September 30, 1991 Harasewych & Kantor 13

DISCUSSION

The genus Marginellona and its type species M. gigas were described
largely on the basis of anatomical and radular characters, supplemented
with a shell description reconstructed from fragments of the unique
holotype. Sigaluta and its type species S. pratasensis, on the other hand,
were erected exclusively on features of shell morphology. The availability
of preserved specimens in which the shell morphology matches that of
S. pratasensis (compare Figs. 2 and 4), while the unusual morphologies
of the radula (compare Figs. 10 and 11), head, and external anatomy
agree with published descriptions of these features in M. gigas (Martens,
1904: 109; Thiele, 1904: 170; 1929: 356) leaves little doubt of the
synonymy of these two species and the nominal genera based upon them.

Martens (1904: 109) and Thiele (1904: 170; 1929: 356) noted the
similarities of M. gigas to certain volutids, especially in features of the
head and columellar folds, but both authors included this taxon in
Marginellidae on the basis of radular morphology, absence of operculum,
polished shell surface and glazed suture. Tomlin (1917: 268) disputed
the inclusion of this taxon in Marginellidae, but did not offer an
alternative familial placement. It is interesting to note that Afrivoluta
pringlei (Tomlin, 1947), the only other species in the subfamily
Marginelloninae, was also originally described as a volutid based on shell
morphology. This genus and species were subsequently transferred to
the Marginellidae when its radula was examined (Barnard, 1963).

In a survey of the literature dealing with anatomy of marginellids,
we have identified a number of anatomical features of possible utility for
phylogenetic inference (Table 2), and noted their distribution among the
few taxa studied to date (Table 3). Marginellona gigas has several features
that distinguish it from other marginellids, while possessing others that
appear to be widespread in the family and may prove to be diagnostic
of the Marginellidae.

As noted by Ponder (1970: 61), the fusion of the siphon to the left
side of the head appears to be an adaptation for burrowing. This feature
occurs in each of the three Recent marginellid subfamilies, and is likely
symplesiomorphic within Marginellidae. A deep channel on the dorsal
surface of the head, apparently formed by fusion of the ventral surfaces
of the tentacle bases, occurs in Marginellona, and in several genera within
Cystiscinae (Coovert, 1987: 19-22). This feature has not been reported
in Afrivoluta (Barnard, 1963; Liltved, 1985), but a similar cleft occurs
in the volutid subfamilies Scaphellinae (Clench & Turner, 1964: 135),
Athletinae (Woodward, 1900: 121) and Calliotectinae (Pace, 1902: 25).
The lateral lappets on the proximal portion of the head of Marginellona

14 NEMOURIA No. 37

Table 2. Survey of anatomical features reported to vary within the family
Marginellidae. Sources for data listed in Table 3.

1) Siphon: (0) to left of head; (1) fused to head.

2) Tentacles: (0) separate; (1) fused, with dorsal furrow.

3) Wing-like folds at bases of tentacles: (0) absent; (1) present.

4) Buccal caecum: (0) present; (1) absent.

5) Radula: (0) present; (1) absent.

6) Salivary glands: (0) ascinous; (1) tubular.

7) Accessory salivary gland: (0) absent; (1) single gland present.
8) Valve of Leiblein: (0) present; (1) absent.

9) Esophageal caecum: (0) absent; (1) present.

10) Gland of Leiblein (‘‘Poison Gland’’): (0) broad, sacculate, lacking
terminal bulb; (1) narrow, glandular, without terminal bulb; (2)
narrow, tubular, with terminal bulb.

11) Duct from gland of Leiblein: (0) entering esophagus posterior to
nerve ring; (1) bypassing valve of Leiblein.

12) Number of glandular structures joining pallial oviduct between
albumen gland and capsule gland: 1, 2, or 3.

13) Prostate gland opening to mantle cavity via: (0) slit; (1) tube.

14) Prostate gland: (0) along right wall of mantle cavity; (1) in penis.

15) Penis: (0) simple, with one lobe; (1) bilobed.

Table 3. Comparison of anatomical features of marginellids. Sources for
data footnoted. ? indicates that character state could not be inferred from
cited text or figure. Assignment of genera to subfamilies follows Coan
(1965). Species nomenclature updated according to Coovert (1989).

Characters from Table 1

123 4° 5 6 F899) 40%) WANE RIG) 14ts
Subfamily: Marginellinae
Fleming, 1828
Prunum marginata 10000 O° Oy P42 er rae |)
(Born 1778)
(as Marginella marginata)!
Marginella desjardini 0.0 Ocll 2 eh Ou? b2ui enn ne

(Marche-Marchad, 1957)!

September 30, 1991

Table 3. (continued)

Prunum martini (Petit, 1853) 0 0

(as Marginella fraterculus)
E. A. Smith, 1915?

Hyalina hyalina
(Thiele, 1913)?

Hyalina pallida
(Linné, 1758)4

Volvarina taeniolata
Morch, 1960°

Haloginella mustelina
(Angas, 1871)
(as Volvarina mustelina)®

Dentimargo cairoma
(Brookes, 1924)

(as Volvarinella cairoma)®
Mesoginella pygmaea
(Sowerby, 1846)°®
Subfamily: Cystiscinae

Stimpson, 1865

Granula sp.®

Subfamily: Marginelloninae

Coan, 1965

Marginellona gigas
(Martens)®

©

5
0

Harasewych & Kantor

6 7 8 9 10

]

O?12

Characters from Table 1

11 12
lz

15

1Graham, 1966; 2Marcus & Marcus, 1968; 3Eales, 1923; #Coan & Roth, 1976;
>Fretter, 1976; Ponder, 1970; 7Ponder, 1970, as Diliculum sp., see Coovert,

1989: 13; 8herein.

16 NEMOURIA No. 37

and Afrivoluta (Barnard, 1963) appear to be restricted to the subfamily
Marginelloninae within the Marginellidae, but are widespread throughout
the family Volutidae.

The presence of a buccal caecum or pouch, which contains the
buccal mass including odontophores and radula, with a sphincter at its
opening to the buccal cavity is a feature unique to marginellids and
toxoglossans (Fretter, 1976: 333). This structure may, however, be
reduced or lost in aradulate species. Marginellid radular morphology has
recently been reviewed by Coovert (1989), who defined seven radular
types. Although the loss of a radula is believed to be polyphyletic, all
aradulate taxa known to date are in the subfamily Marginellinae.

Salivary glands within the family Marginellidae may be ascinous or
tubular. Ascinous salivary glands appear to be uncommon and restricted
to the subfamily Marginellinae. A single, medial accessory salivary gland
has been reported in several marginelline and the only cystiscine taxon
studied thus far.

Within the family Marginellidae, the reduction and loss of the valve
of Leiblein appears to be correlated with the modification of the gland
of Leiblein to form a long, tubular “‘poison gland”’ that empties, via a
separate duct, into the buccal cavity. Marginellona gigas, which has a very
large, ascinous gland of Leiblein that empties into the mid-esophagus
posterior to the nerve ring, also has a very large and muscular valve of
Leiblein. Progressive stages in the development of a duct from the gland
of Leiblein that bypasses the valve of Leiblein have been documented
by Ponder (1970; 1973: fig. 3, L-O). Those marginellid taxa that lack
a valve of Leiblein all have a “‘poison gland’’ with a separate duct that
empties into the buccal cavity. Two of three taxa with an esophageal
caecum posterior to the nerve ring lack a valve of Leiblein. The third
species, Prunum martini, was reported to have an “‘inconstant valve of
Leiblein’’ based on a slight swelling of the esophagus observed in 3 of
5 specimens dissected (Marcus & Marcus, 1968: 65, as Marginella
fraterculus).

The pallial oviduct of marginellids differs from that of most other
neogastropods (Ponder, 1973: fig. 6) in having two glandular structures
between the capsule and albumen glands. The male genital ducts agree
in general respect with those of most other neogastropods, but exhibit |
infrafamilial variation in: position and size of prostate gland (rear of |
mantle cavity vs. in penis), type of opening to pallial cavity (duct or slit), |
and penis morphology (simple vs. bilobed).

Anatomical features of Marginellona gigas, including the presence ©
of a buccal caecum, a uniserial radula with broad, comb-like rachidian —
teeth, a simple stomach with single, broad opening to the digestive gland, —

September 30, 1991 Harasewych & Kantor 17

a large rectal gland, and a pallial oviduct with separate ingesting gland
and seminal receptacle, further support the original assignment of this
taxon to the family Marginellidae. However, the relationship of the
Marginelloninae to the Marginellinae and Cystiscinae remains unclear.
Coovert (1989: 30) suggested that Cystiscinae is the most primitive
subfamily in Marginellidae, based on the occurrence of a triserial radula
in a species tentatively ascribed to the genus Cystiscus. Marginellona gigas
has an unspecialized alimentary system, containing a large gland of
Leiblein without terminal bulb, as well as a large valve of Leiblein. ‘These
are primitive neogastropod features previously unrecorded in the Margi-
nellidae. Marginellona gigas also lacks both the glandular duct that
bypasses the valve of Leiblein, and the single accessory salivary gland
that are present in Marginellinae and in the only species of Cystiscinae
to be studied to date. Thus, the morphology of the anterior alimentary
system suggests that Marginelloninae may be the most primitive subfamily
in Marginellidae.

Although members of the Marginellidae share features with the
Toxoglossa (buccal caecum, Fretter, 1976: 333), Volutomitridae (single
accessory Salivary gland, Ponder, 1973: 330) and Volutidae (furrowed
head with lateral lappets, Martens, 1904: 109; Thiele, 1929: 356), we
agree with Fretter, (1976: 335) that our knowledge of this family is, as
yet, too scanty to deduce its sister group.

ACKNOWLEDGMENTS

We thank Dr. Boris Sirenko, of the Zoological Institute, Leningrad,
for making the preserved material available for this study. The senior
author gratefully acknowledges grants from the A. N. Severtzov Institute
of Animal Evolutionary Morphology and Ecology, Academy of Sciences
of the U.S.S.R., and the Smithsonian Institution Research Opportunities
Fund for support of travel to the Zoological Institute, Leningrad.

The scanning electron micrographs were made by Ms. Susanne
Braden of the Scanning Electron Microscopy Laboratory, National
Museum of Natural History, Smithsonian Institution.

LITERATURE CITED
Barnard, K. H. 1963. The family position of Afnivoluta pringlei Tomlin. Journal
of Conchology, 25(5): 198-199.

Bieler, R. & K. J. Boss. 1989. Johannes Thiele and his contributions to zoology.
Part. 1. Biography and bibliography. Nemouria, 34: 1-30.

18 NEMOURIA No. 37

Clench, W. J. & R. D. Turner. 1964. The subfamilies Volutinae, Zidoninae,
Odontocymbiolinae, and Calliotectinae in the western Atlantic. Johnsonia,
4(43): 129-180.

Coan, E. 1965. A proposed reclassification of the family Marginellidae. The
Veliger, 7(3): 184-194.

Coan, E. V. & B. Roth. 1976. Status of the genus Hyalina Schumacher, 1817
(Mollusca: Gastropoda). Journal of Molluscan Studies, 42(2): 217-222.

Coovert, G. 1987. A literature review and summary of marginellid external
anatomy. Marginella Marginalia, 3(2-3): 8-25.

Coovert, G. 1989. A literature review and summary of published marginellid
radulae. Marginella Marginalia, 7(1-6): 1-37.

Cossmann, A. E. M. 1906. Marginellidae. Essais de Paléoconchologie Comparée,
3: 79-99.

Eales, N. B. 1923. Mollusca. Part V.-Anatomy of Gastropoda (except Nudibran-
chia). British Museum (Natural History), British Antarctic (Terra Nova’)
Expedition, 1910. Natural History Report. Zoology, 7: 1-45.

Fretter, V. 1976. The anatomy and feeding of the volutacean prosobranch
Volvarina taemolata Morch. Journal of Molluscan Studies, 42(3): 327-336.

Graham, A. 1966. The fore-gut of some marginellid and cancellariid prosobran-
chs. Studies in Tropical Oceanography, 4(1): 134-151.

Harasewych, M. G. & Y. I. Kantor. 1991. Tenebrincola frigida, a new genus and
species of abyssal volute from the northern Pacific Ocean (Gastropoda:
Volutidae). Nemouria, 38: 1-11.

Hedegaard, C. 1990. Shell structures of the Recent Archeogastropoda. Cand.
Sci. Thesis, University of Aarhus, Denmark, 2 vols., 154 pp., 78 pls.

Liltved, W. R. 1985. Afnvoluta pringlet Tomlin, 1947. The Strandloper, Bulletin
of the Conchological Society of Southern Africa, 214: 9.

Marcus, E. & E. Marcus. 1968. On the prosobranchs Ancilla dimidiata and
Marginella fraterculus. Proceedings of the Malacological Society of London,
38(1): 55-69.

Martens, E. von. 1904!. Die beschalten Gastropoden der deutschen Tiefsee-
Expedition 1898-1899. A. Systematisch-geographischer Teil. Wissenschaftli-
che Ergebnisse der deutschen Tiefsee- Expedition auf dem Dampfer
*“Valdivia’’, 7: 1-146, pls. 1-4.

Pace, S. 1902. On the anatomy and relationships of Voluta musica, Linn.; with |
notes upon certain other supposed members of the Volutidae. Proceedings _

of the Malacological Society of London, 5(1): 21-31, pl. 11.
Ponder, W. F. 1970. Some aspects of the morphology of four species of the

neogastropod family Marginellidae with a discussion of the evolution of the |

toxoglossan poison gland. Journal of the Malacological Society of Australia,
2(1): 55-81.

|

September 30, 1991 Harasewych & Kantor 19

Ponder, W. F. 1973. The origin and evolution of the Neogastropoda. Malacologia,
12(2): 295-338.

Rehder, H. A. 1967. A new genus and two new species in the families Volutidae
and Turbinellidae (Mollusca: Gastropoda) from the western Pacific. Pacific
Science, 21(2): 182-187.

Rehder, H. A. 1970. Supplementary comments on deep water Volutidae from
the South China Sea and South Africa. The Veliger, 12(4): 415-416, figs.
1-6.

Rokop, F. J. 1972. Notes on abyssal gastropods of the eastern Pacific, with
descriptions of three new species. The Veliger, 15(1): 15-19, figs. 1-15.

Thiele, J. 1904!. Die beschalten Gastropoden der deutschen Tiefsee-Expedition
1898-1899. B. Anatomisch-systematische Untersuchungen einiger Gastro-
poden. Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition auf
dem Dampfer ‘‘Valdivia’’, 7: 147-180, pls. 6-9.

Thiele, J. 1929. Handbuch der systematischen Weichtierkunde, 1(1): 1-376, 470
figs. Gustav Fischer, Jena.

Tomlin, J. R. le B. 1917. A systematic list of the Marginellidae. Proceedings of
the Malacological Society of London, 12(5/6): 242-306.

Wagner, R. and R. T. Abbott, 1990. Standard Catalog of Shells, Supplement
4. World size records, 1990. American Malacologists, Inc., Melbourne,
Florida. ii + pp. 80-001-80-080.

Weaver, C. S. and J. E. duPont. 1970. Living Volutes. Delaware Museum of
Natural History, Greenville. xv + 375 p.

Wenz, W. 1943. Gastropoda. In: O.H. Schindewolf (ed.), Handbuch der
Paléozoologie. Band 6. Gebriider Borntraeger, Berlin. Teil 6: 1201-1506.

Woodward, M. F. 1900. Note on the anatomy of Voluta ancilla (Sol.), Neptuneopsts
gilchnsti, Sby. and Volutolithes abyssicola (Ad. & Rve). Proceedings of the
Malacological Society of London, 4(3): 117-125, pl. 10.

FOOTNOTE

1_Although the title page is dated 1903 and many earlier authors, including
Thiele, had cited this date, the work was published in January, 1904 (see Bieler
- & Boss, 1989: 12).

The manuscript was reviewed by Mr. Gary A. Coovert (Dayton Museum of
Natural History, OH) and Dr. Gary D. Rosenberg (Academy of Natural Sciences
of Philadelphia, PA).

Founding Editor Managing Editor
JOHN E. DU PONT RUDIGER BIELER

Consulting Editors
WILLIAM K. EMERSON, American Museum of Natural History, New
York, NY.
M. G. HARASEWYCH, National Museum of Natural History, Smithsonian
Institution, Washington, DC.
KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh, |
PA. |
RICHARD W. ‘THORINGTON, JR., National Museum of Natural History, |
Smithsonian Institution, Washington, DC.

Published by the

DELAWARE MUSEUM OF NATURAL HISTORY
4840 Kennett Pike, Wilmington,
Delaware 19807, U.S.A.

|
Blo
ae

ir NEMOURIA

Occasional Papers of the Delaware Museum of Natural History

NUMBER 38 SEPTEMBER 30, 1991

TENEBRINCOLA FRIGIDA, A NEW GENUS AND
SPECIES OF ABYSSAL VOLUTE FROM
THE NORTHERN PACIFIC OCEAN
(GASTROPODA: VOLUTIDAE)

M. G. Harasewych! and Yuri-I. Kantor?

ABSTRACT. TJenebrincola frigida, a new genus and species of volute, is described
on the basis of a specimen taken from the Aleutian Trench at a depth of 5020
meters. It differs from its only congener, 7: cukri (Rokop, 1972), in having a
broader shell with a distinct shoulder. Tenebrincola is included in the subfamily
Fulgorariinae on the basis of the morphology of its radula, operculum, accessory
salivary glands, and male reproductive system. The eastern Australian genus
Ericusa is tentatively considered to be the sister group, based on conchological
similarity.

Key words: Volutidae, Fulgorariinae, Tenebrincola, abyssal, anatomy, sys-
tematics.

INTRODUCTION

In the course of examining the collections of the Institute of
Oceanology, Academy of Sciences of the U.S.S.R., in Moscow, a single
specimen of an unusual, operculate volutid, collected by the R/V Vityaz
in the Aleutian Trench, was discovered. This specimen represents a new
species that most closely resembles Sigaluta cukri Rokop, 1972, which
occurs in abyssal depths off Baja California. Because the genus Sigaluta
Rehder, 1967, recently has been shown to be a synonym of Marginellona
Martens, 1904, of the family Marginellidae (Harasewych & Kantor,

1Department of Invertebrate Zoology, National Museum Serial Publication
of Natural History, Smithsonian Institution, Washington, ISSN 0085-3887
DC 20560, U.S.A.

2A. N. Severtzov Institute of Animal Evolutionary

Morphology and Ecology, Academy of Sciences of the U.S.S.R.,

Lenin Avenue, 33, Moscow 117071, U.S.S.R.

2 NEMOURIA No. 38

1991), a new genus is proposed to include both abyssal species.
Tenebrincola is referable to the subfamily Fulgorariinae, and represents
the first record of this subfamily in the abyssal fauna.

SYSTEMATICS

Family Volutidae Rafinesque, 1815
Subfamily Fulgorariinae Pilsbry and Olsson, 1954

Tenebrincola new genus

Type species: Tenebrincola frigida new species [described below].
Description: Shell to at least 68.4 mm, ovately to elongate fusiform,
extremely thin and fragile. Protoconch of about 2 inflated whorls, nucleus
deflected about 45° from shell axis, transition to teleoconch indistinct.
Teleoconch with up to 3 whorls. Outer lip smooth. Columella without
folds. Color ivory to white, internally and externally. Periostracum
smooth, extremely thin. Operculum small, corneous, ovate, with tapering,
terminal nucleus. Eyes absent. Siphonal appendages tapering, symmetri-
cal. Accessory salivary glands short, broad posteriorly. Radula uniserial,
with tricuspid rachidian teeth having broadly U-shaped attachment
region; lateral cusps much broader but slightly shorter than central cusp.
Pallial sperm duct closed. Penis with broad, lateral flap.

Etymology: L. tenebra—darkness + L. incola—inhabitant.

Tenebrincola frigida new species
Figures 1, 4-10

Description: Shell (Fig. 1) to at least 68.4 mm, extremely thin, fragile,
translucent, ovately fusiform, with low, blunt spire. Protoconch large,
bulbous, eroded, of about 2 whorls. Nucleus deflected from shell axis
by 43°. Transition to teleoconch indistinct. Teleoconch with about 3
convex, weakly shouldered whorls. Suture adpressed. Surface sculpture
limited to very fine growth striae. Aperture ovate, rounded anteriorly,
sharply tapered posteriorly. Outer lip smooth, thin, sinuous. Inner lip
smooth, parietal region not thickened. Columella lacking columellar or
siphonal folds. Periostracum (Fig. 4) very thin (= 1.5 wm), smooth,
continuous, imparting lustrous, ivory color to outer surface of shell.
Aperture white. Operculum (Fig. 7) small (0.2 Xaperture length),
corneous, narrowly ovate, with tapering terminal nucleus (abraded away
in holotype).

Ultrastructure: Shell thin (= 85 um), composed of three layers of

September 30, 1991 Harasewych & Kantor 3

orthogonally oriented, crossed-lamellar crystals (Fig. 5): outer layer 50
uum thick, crystal planes perpendicular to growing edge; middle layer 33
um thick, crystal planes parallel to growing edge; inner layer 2.5 um
thick, crystal planes perpendicular to growing edge.

Anatomy: [The soft tissues of the single, male specimen were very
friable. The visceral mass and portions of the mantle cavity disintegrated
during extraction from shell.] Animal uniformly rust-colored. Foot *4
length of shell aperture, broad (L/W = 2.3), with small anterior lateral
expansions delimiting propodial mucous gland, rounded posteriorly.
Siphon (Fig. 8, S) short, stout, with paired, symmetrical, tapering
siphonal appendages (Fig. 8, SA) on either side of siphon near base.
Head broad, with long, narrow, tapering tentacles (Fig. 8, T) on either
side of thin, medially furrowed hood (Fig. 8, MF) extending over
rhynchostome. Outer edges of tentacles with short, narrow, wing-like
lobes (Fig. 8, W). Eyes absent. Mantle cavity broad, shallow, with large,
dark osphradium (L/W = 2.8); ctenidium as wide and twice as long as
osphradium; hypobranchial gland large, transversely pleated; rectum
broad with conspicuous rectal gland along anterior half of dorsal surface.
Proboscis short, broad, pleurombolic, with thick, muscular walls. Probos-
cis retractor muscles thick, paired, extending from lateral mid-points of

Figure 1. Tenebrincola frigida, new species. Holotype, ZMUM N Lc 16252,
Aleutian Trench, SW of Agattu Island, Near Islands, 51°30’06”N, 172°04'30”E,
in 5020 m. 0.8 X.

4 NEMOURIA No. 38

Figure 2. Tenebrincola cukri (Rokop, 1972). Holotype, USNM 701261, WSW
of Cortes Bank, approximately 225 miles W. of Ensenada, Baja California,
Mexico, 32°05’N, 120°29'W to 32°03'N, 120°30'W, in 3775-3789 m. 0.8 X.

Figure 3. Ericusa sericata Thornley, 1951. USNM 845887, off Cape Moreton,
Queensland, Australia, in 180 m. 0.55 X.

September 30, 1991 Harasewych & Kantor 5

Figures 4-7. Tenebrincola frigida, new species. 4. Periostracum on shell fragment
from dorsal surface of body whorl (see Fig. 1). Scale bar = 30 um. 5. Shell
ultrastructure (same fragment as in Fig. 4). Fracture surface parallel to growing
edge, inner surface at bottom of photograph, periostracum at top. Scale bar =
30 um. 6. Lateral (55°) and dorsal views of radula. Scale bar = 100 um. 7.
Inner and outer surfaces of operculum. 2.6 X.

6 NEMOURIA No. 38

Figures 8-10. Tenebrincola frgida, new species. Anatomical features. 8. Dorsal
view of head, with siphon reflected to left. 9. Anterior portion of alimentary
system. 10. Penis, dorsal view and transverse section at mid-length. Scale bar =
1 cm for all drawings.

ASG, accessory salivary gland; DASG, duct of accessory salivary gland; DSG,
duct of salivary gland; GL, gland of Leiblein; LF, lateral flap; MF, medial furrow;
NR, nerve ring; PE, posterior esophagus; PAP, papilla; PD, penial duct; RS,
radular sac; S, siphon; SA, siphonal appendage; SG, salivary gland; T, tentacle;
VD, vas deferens; VL, valve of Leiblein; W, wing-like projection.

September 30, 1991 Harasewych & Kantor i

inner proboscis wall to walls of cephalic hemocoel. Buccal mass muscular.
Anterior esophagus broad, thick-walled. Radula (Fig. 5) short (4.2 mm),
uniserial, with 44 teeth; teeth tricuspid, with narrow, anteriorly recurved
basal plates; lateral cusps broadly triangular, thickened along lateral
edges, slightly shorter than narrow, flexible central cusp. Accessory
salivary glands (Fig. 9, ASG) short, tubular, distally expanded, lying
dorsal and anterior to salivary glands and entering buccal cavity by
separate ducts (Fig. 9, DASG). Salivary glands (Fig. 9, SG) small,
acinous, with ducts running laterally along esophagus before becoming
embedded in the walls at rear of retracted proboscis. Valve of Leiblein
(Fig. 9, VL) large, inflated. Gland of Leiblein (Fig. 9, GL) long, highly
convoluted, tubular, as wide as posterior esophagus (Fig. 9, PE).
[Stomach and intestine not preserved in dissected specimen.] Rectum
large, inflated, distally tapered, with green rectal gland lining anterior
half of dorsal surface. Anus with long, ventral papilla. Penis (Fig. 10)
as broad as long, dorsoventrally flattened, with short, terminal papilla
(Fig. 10, PAP) and thin lateral flap (Fig. 10, LF). Vas deferens (Fig.
10, VD) leading to base of penis and proximal 2/3 of penial duct closed
(Fig. 10, PD). Distal 1/3 of penis with penial duct running along ventral
surface forming groove extending to tip of papilla. Nerve ring (Fig. 9,
NR) highly concentrated, of Type 2 (Ponder, 1970: 159).

Type Locality: Aleutian Trench, SW of Agattu Island, Near Islands,
51°30’06"N, 172°04'30"E, in 5020 m. R/V Vityaz sta. 3359, Sigsbee
trawl, 8 June 1955.

Material examined: Holotype, Zoological Museum, University of
Moscow (ZMUM) N Lc 16252, 68.4 mm.

Distribution: This species is known only from the type locality.

Ecology: Although this species lives below both the calcite and aragonite
compensation depths (Morse & Berner, 1979), there is no evidence of
pitting or dissolution on the shell surface. Regions of the protoconch are
eroded, possibly by abrasion. Rectal contents included polychaete
spicules, sand, and foraminiferans.

Etymology: L. frigidus—cold.

Comparative remarks: This species is distinguished from Tenebrincola
cukn (Rokop, 1972), its only congener, by its proportionally broader
Shell, by the presence of a weak shoulder on the body whorl, and by the
slightly concave region between the shoulder and suture. Because T.
cukri is known only from a single specimen lacking soft parts, compari-
sons of radula or anatomical features cannot be made.

8 NEMOURIA No. 38

DISCUSSION

Although Stuardo and Villarroel (1974: 146) regarded the radular
morphology of the volutid subfamilies Zidoninae and Fulgorariinae to
be convergent, the three-dimensional topology of radular teeth, elucidated
by scanning electron microscopy, clearly distinguishes these two subfami-
lies. Radular teeth of Zidoninae are generally flat or convex on their
dorsal surface, with central cusps that are longer and as broad or broader
than the lateral cusps (Harasewych, 1991). Those members of Fulgo-
rariinae examined to date (Harasewych, 1991) are characterized by
rachidian teeth that have strongly curved basal plates and lateral cusps
that are thickest along the lateral edges and broader and generally longer
than the central cusps, forming a trough along the mid-line of the radular
ribbon. The radula of Tenebrincola frigida is of the fulgorariine type, but
is unusual in having very thin and somewhat flexible central cusps that
are slightly longer than the lateral cusps.

Comparatively few of the ten Recent subfamilies of Volutidae have
been studied anatomically in any detail. The best known are the
Zidoninae and Odontocymbiolinae (Clench & Turner, 1964; Ponder,
1970; Harasewych, 1987; Leal & Bouchet, 1989). The Volutinae,
Scaphellinae, Calliotectinae, Lyriinae and Athletinae are less well known
(Fischer, 1867; Woodward, 1900; Pace, 1902; Clench & Turner, 1964),
while knowledge of the remaining subfamilies is largely limited to radular
or gross external morphology (summarized in Weaver & duPont, 1970).

The presence of an operculum, short accessory salivary glands that
are not wound around the salivary glands, and a closed vas deferens and
penial duct prevent inclusion of Jenebrincola in the Zidoninae, but support
its assignment to Fulgorariinae.

The shallow, medial furrow in the head of Tenebrincola frigida is
regarded as a plesiomorphic volutid character that is shared with members
of the subfamilies Calliotectinae, Athletinae and Scaphellinae. The long
tentacles, absence of functional eyes, thin shell, and lack of columellar
or siphonal folds are all convergently derived adaptations for abyssal life.

The protoconch and teleoconch morphology of both species of
Tenebrincola most closely resembles that of Ericusa sericata Thornley,
1951 (Fig. 3), from 80 to 200 m off eastern Australia. Thus, the genus
Encusa H. & A. Adams, 1858, is tentatively considered to be the sister
group.

Of the abyssal volutes known to date (List 1) all but Arctomelon
benthalis are members of genera that are endemic to abyssal depths.
Tenebrincola is the first record of the subfamily Fulgorariinae in the
abyssal fauna. Tenebrincola frigida is the deepest dwelling volutid yet
discovered, and the only volutid known to occur in a trench.

September 30, 1991 Harasewych & Kantor 9

List 1. Members of the family Volutidae reported from abyssal (2 2000
m) depths. Subfamily assignment in square brackets [ ]. Geographic and
bathymetric distributions based on references in {}.

Arctomelon benthalis (Dall, 1896) [Zidoninae] 3058 m. Peru Basin.
‘Clarke, 1962: 27}.

Guivillea alabastrina (Watson, 1882) [Zidoninae] 2688-4063 m. Southern
Ocean, from Drake Passage east to Croizet Basin, and north to
Mozambique Channel. {Knudsen, 1973; Dell, 1990: 220-221}.

Tractohra sparta Dall, 1896 [Odontocymbiolinae] 2990-4082 m. Peru,
Guatemala and Central Pacific Basins. {Rokop, 1972}.

Tractolira germonae Harasewych, 1987 [Odontocymbiolinae] 1976-3944
m. South Antilles and Atlantic-Indian-Antarctic Basins. {Dell, 1990:
217}.

Tractolira tenebrosa Leal & Bouchet, 1989 [Odontocymbiolinae] 2370-
3270 m. Brazil Basin. {Leal & Bouchet, 1989: 6-7}.

Tractohra sp. [Odontocymbiolinae] 1772-3596 m. Ross Sea, Pacific
Antarctic Basin. {Dell, 1990: 217}.

Tenebrincola cukn (Rokop, 1972) [Fulgorariinae] 3775-3789 m. Central
Pacific Basin. {Rokop, 1972: 16-17}.

Tenebrincola fngida Harasewych & Kantor, 1991 [Fulgorariinae] 5020
m. Aleutian Trench. {herein}.

10 NEMOURIA No. 38

ACKNOWLEDGMENTS

We thank Valentina Lus of the Institute of Oceanology, Academy
of Sciences of the U.S.S.R., Moscow, for making the material available
for study. The senior author gratefully acknowledges grants from the
Academy of Sciences, U.S.S.R. and the Smithsonian Institution Research
Opportunities Fund for support of travel to the Institute of Oceanology,
Moscow. Susanne Braden provided valuable assistance with the SEM.

LITERATURE CITED

Clarke, A. H., Jr. 1962. Annotated list and bibliography of the abyssal marine
molluscs of the world. National Museum of Canada Bulletin, 181: 1-114.

Clench, W. J. and R. D. Turner. 1964. The subfamilies Volutinae, Zidoninae,
Odontocymbiolinae and Calliotectinae in the western Atlantic. Johnsonia,
4(43): 129-180.

Dell, R. K. 1990. Antarctic mollusca, with special reference to the fauna of the
Ross Sea. The Royal Society of New Zealand, Bulletin 27. iv + 311 pp.

Fischer, P. 1867. Sur l’anatomie des Lyna. Journal de Conchyliologie, 15(4):
349-356, pl. 13.

Harasewych, M. G. 1987. Tractolira germonae, a new abyssal Antarctic volute.
The Nautilus, 101(1): 3-8.

— 1991. The bathyal and abyssal volutes of the eastern Pacific. Nemouria (in
press).

Harasewych, M. G. and Y. I. Kantor. 1991. Rediscovery of Marginellona gigas
(Martens, 1904), with notes on the anatomy and systematic position of the
subfamily Marginelloninae (Gastropoda: Marginellidae). Nemouria, 37:
1-19.

Knudsen, J. 1973. Guivillea alabastrina (Watson, 1882), an abyssal volutid.
(Gastropoda: Mollusca). Galathea Report, 12: 127-131, pl.18, figs.1-5.

Leal, J. H. and P. Bouchet. 1989. New deep-water Volutidae from off
southeastern Brazil (Mollusca: Gastropoda). The Nautilus, 103(1): 1-12.

Morse, J. W. and R. C. Berner. 1979. Chemistry of calcium carbonate in the
deep oceans. Jn: Jenne, E. A. (ed.). Chemical modeling in aqueous systems.
American Chemical Society Symposium Series, 93: 499-535.

Pace, S. 1902. On the anatomy and relationships of Voluta musica Linn.; with
notes upon certain other supposed members of the Volutidae. Proceedings
of the Malacological Society of London, 5(1): 21-31, pl.11.

Ponder, W. F. 1970. The morphology of Alcithoe arabica (Gastropoda: Volutidae).
Malacological Review, 3: 127-165.

Rokop, F. J. 1972. Notes on abyssal gastropods of the eastern Pacific, with
descriptions of three new species. The Veliger, 15(1): 15-19, figs. 1-15.

September 30, 1991 Harasewych & Kantor 11

Stuardo, J. and M. Villarroel. 1974. On some living and fossil volutes referred
to Miomelon Dall, 1907 and Proscaphella von Ihering, 1907. The Veliger,
17(2): 139-155, figs. 1-21.

Weaver, C. S. and J. E. duPont. 1970. Living Volutes. Delaware Museum of
Natural History, Greenville. xv + 375 pp.

Woodward, M. F. 1900. Note on the anatomy of Voluta ancilla (Sol.), Neptuneopsts
gilchristt (Sby.) and Volutilithes abyssicola (Ad. & Rve.). Proceedings of the
Malacological Society of London, 4(3): 117-125, pl. 10.

The manuscript was reviewed by Dr. Ruth D. Turner (Museum of Comparative
Zoology, Cambridge, MA) and an anonymous reviewer.

Founding Editor Managing Editor
JOHN E. DU PONT RUDIGER BIELER

Consulting Editors

WILLIAM K. EMERSON, American Museum of Natural History, New
York, NY.

M. G. HARASEWYCH, National Museum of Natural History, Smithsonian
Institution, Washington, DC.

KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh,
PA.

RICHARD W. THORINGTON, JR.,; National Museum of Natural History,
Smithsonian Institution, Washington, DC.

Published by the

DELAWARE MUSEUM OF NATURAL HISTORY
4840 Kennett Pike, Wilmington,
Delaware 19807, U.S.A.

NEMOURIA

Occasional Papers of the Delaware Museum of Natural History

NUMBER 39 SEPTEMBER 30, 1991

JOHANNES THIELE AND HIS CONTRIBUTIONS
TO ZOOLOGY.
PART 2. GENUS-GROUP NAMES (MOLLUSCA)

Kenneth J. Boss! and Riidiger Bieler?

ABSTRACT. This is the second part of a series on the German zoologist Johannes
Thiele (1860-1935) and comprises a critical listing of the genus-group taxa which
he described as new to malacology. Each of these names is accompanied by
author and bibliographic references, original status, type-species with its original
binominal spelling and bibliographic source and some data on subsequent
taxonomic placements. Thiele introduced a total of 291 such names in the Phylum
Mollusca, distributed as follows: 11 Aplacophora; 39 Polyplacophora; 200
Gastropoda (138 Prosobranchia; 20 Opisthobranchia and 42 Pulmonata); 31
Bivalvia; 10 Cephalopoda; there were no new scaphopod or monoplacophoran
names. Of these, later authors recognized as valid 85 at the generic level, 110
at the subgeneric level; 71 are considered to be synonyms, and the remaining
25 are unjustified emendations or errors.

INTRODUCTION

As part of a series on the scientific contributions of Johannes Thiele,
the eminent German zoologist, we provide here an alphabetical listing
and analysis of all the genus-group taxa introduced by him in his
publications on mollusks as delineated by Bieler & Boss (1989). A total
of 291 names is included in the following format: (1) genus-group name;
(2) author(s); (3) year of publication; (4) condensed bibliographic
reference; (5) original status as given by Thiele; (6) subsequent status

1Museum of Comparative Zoology, Harvard University, Serial Publication
Cambridge, Massachussetts 02138, U.S.A. ISSN 0085-3887
2Delaware Museum of Natural History. Current address:

Field Museum of Natural History, Roosevelt Road at Lake

Shore Drive, Chicago, Illinois 60605, U.S.A.

NEMOURIA No. 39

Portrait of Johannes Thiele (undated, but photograph
probably taken shortly before his seventieth birthday
in 1930) and sample of his distinctive handwriting

on specimen label.

Zoolog. Museum Berlin.
kita
wend fo &
Niue tid Baral
Mal. 73
B49 74

September 30, 1991 Boss & Bieler 3

as assigned by Thiele himself or other authors; (7) method of designation
of the type-species; (8) name of the type-species as originally given by
Thiele; (9) the exact or correct binominal combination of the type-species
with its original author and page and other appropriate citations [Thiele’s
own species names of mollusks will be covered in future parts of this
series]; (10) family-level placement (a family name without brackets
indicates an original placement at the time of the original description;
names enclosed by brackets are subsequent placements either by Thiele
or other authors); and (11) class and/or other higher category.

For each taxon we have paid special attention to the type-species:
its original binomen is given along with a reference to its original
introduction into the literature and, in most cases, a citation of its first
illustration or figure. In addition, we provide data on the subsequent
hierarchical placement of the genus-group taxon; reference here is given
to certain classic authoritative texts (e.g., Wenz, Zilch, etc.), all of which
are listed in the bibliography.

NOTES ON RANKING OF TAXA

Usually, Thiele clearly indicated that he was establishing a new
genus-group taxon and introduced his new names as genera, subgenera,
or ‘sections.’ In some cases he omitted a rank and/or just referred to a
new “‘Gruppe”’ or “‘Untergruppe.”’ According to the International Code
of Zoological Nomenclature (ICZN, 1985), Article 10(e), the term
‘section’ is deemed to be a subgeneric name. Thiele himself often
obfuscated the problem of ranking when the sectional or subgeneric
name, instead of the generic name, was used in figure captions.
Nevertheless, we have retained his original ranking to illustrate the
intended position in relation to genus and subgenus.

THIELE’S TYPE CONCEPT

Thiele’s concept of the type-species and its designation is somewhat
at variance with the present, more rigorous regulations of the Interna-
tional Code of Zoological Nomenclature (ICZN, 1961, 1985; Article
68), wherein the listing of the first of several species in a new genus-group
taxon does not constitute a type designation. Part of the difficulty in the
interpretation of type-species for Thiele’s genus-group taxa stems from
his German usage, namely “‘typische Art’? for both ‘“‘typical species’”’ in
the sense of an example, as well as “‘type-species” in the nomenclatural
sense.

In his early publications such as Das Gebiss, Thiele (1893e: 376,

4 NEMOURIA No. 39

see also 372) sometimes considered the first species mentioned under a
generic name to be the type-species: ‘“‘Ausser der typischen Art, die ich
wie gewOhnlich zuerst beschreibe . . .’’. At other times, he apparently
cited a particular species as ‘‘a good example.” Further, he did not
necessarily restrict the example to a single species, but sometimes even
mentioned two typical species such as in the case of his new genus
Radsiella (1893e: 368-369): “‘Als typische Vertreter der Gattung sehe ich
die beiden ersten Arten ... an.” Such departures from current
requirements and usage are occasionally still true for his later works
(e.g., 1925b: 268): “‘von den typischen Arten.. .”’.

The more exacting concepts which we now employ with regard to
type-species, their designation and the like had yet to be universally
adopted by the community of systematists in Thiele’s time. Also, the
formal publication of the International Code of Zoological Nomenclature
in 1961 contained regulations (especially Articles 68 and 69), concerning
the boundary year 1931; this particularly affected taxa established in the
Handbuch der systematischen Weichtierkunde which was published during
this time; these names had to be treated differently with reference to
“original designation.”’

In our treatment of Thiele’s genus-group names, we utilized the
following interpretations: If Thiele explicitly used the word ‘“‘type’’ or its
_ equivalents (e.g., Ischnochiton: ‘“Typus einer eigenen Untergattung. . .”’;
Lissarca: “‘typische Art ist .. .’’), we credited this as type-species by
original designation. When Thiele positively included only one species
in a new genus-group taxon, this was treated as “‘type by monotypy.”’
We did not acknowledge Thiele with an original designation if he used
such phraseology as: “I place this in a new genus’’; “I introduce this for

. .’; rather, if the higher taxon was represented only by a single species,
then we accordingly considered the designation, as noted above, to be
one of monotypy; if two or more species were included, a subsequent
designation had to be found.

SOME PROBLEMS AND INCONSISTENCIES

Certain problems arise in the dating of a taxon, ascertaining the
method of designation of the type-species and attributing authorship to
the name. Thiele occasionally erred, and some of his mistakes were
simply matters of orthography. Others were occasioned by his lack, at
the time he worked, of modern published resources on animal taxa such
as Sherborn’s Index Animalum and Neave’s Nomenclator Zoologicus.
Additional problems were caused by our current application of rules and
interpretations not in use during Thiele’s career, and still more were

September 30, 1991 Boss & Bieler 5

associated with his having numerous papers in press simultaneously in
which new names might be repeatedly used. The following are examples
of some of the problems encountered.

Mistakes in spelling: This includes misspellings of names introduced
by other authors which have or could have caused problems; an example
is Thiele’s erroneous ‘‘Cleovitrea’’ for Iredale’s Cleotrivia.

Further, there is difficulty in distinguishing between misspellings
with no nomenclatural standing (indicated in our listing by placing the
name in quotation marks [*‘. . .”]) and deliberate, but unjustified
emendations. This may be compounded with Thiele’s usage of another
author’s taxon (usually without his naming the other author) in a
broadened, restricted, different or incorrect sense, a so-called “taxon
sensu Thiele.’? With regard to these, several genus-group names of earlier
authors have been later attributed to him; these were usually Thiele’s
misconception, or were subsequently perceived as his misconception of
the taxon of the original author.

Thus, we have Thiele’s misconception of Beanella Dall 1882,
erroneously referred to (by some authors) as Beanella Thiele rather than
more precisely as ‘“‘Beanella ‘Dall’ Thiele iz Troschel & Thiele 1893, non
Dall 1882.’ These so-called misconstrued taxa of Thiele do not have
type-species, although frequently they have had them designated by
subsequent authors. We, of course, cannot list all genus-group taxa falling
into this category. However, we have included a few of these, e.g.,
Artocaelatura ‘Germain’ Thiele 1931, non Germain 1921; Beanella ‘Dall’
Thiele 7n Troschel & Thiele 1893, non Dall 1882; Clathropleura “Tiberi’
Thiele iz Troschel & Thiele 1893, non Tiberi 1877; and Silicula ‘Jeffreys’
Thiele 1934, non Jeffreys 1879.

New names were not always clearly marked as such by Thiele. In
the case of his larger contributions such as the Handbuch der systematischen
Weichtierkunde, some are not properly indicated in the index.

Original authorship was sometimes confused by Thiele: he claimed
taxa of others (e.g., Pyrunculus); he credited authors of manuscript names
(e.g, Arcopella); and he attributed authorship to workers whose publica-
tions were later rejected (e.g., Chemnitz).

Thiele occasionally cited the wrong dates for some of his own taxa
(e.g., 1921 instead of 1924a) or he described taxa twice in subsequent
or “‘simultaneous”’ publications (e.g., Jonicina). Some of these errors can
be attributed either to having several descriptions of the same new taxa
in press simultaneously or to having articles in press for many years. In
a letter to an unnamed colleague at the British Museum (Natural
History), dated 14 February 1923, he complained of a delay in
publication: “Scientific work in Germany has now become very difficult

6 NEMOURIA No. 39

and completed manuscripts usually can be published only after a longer
period. During the war [1914-1918] I had sorted out and worked on
many gastropods from the material of the German Deep-Sea Expedition,
but the work which was completed in the fall of 1918 has not yet been
published’”’ [our translation]. Thiele here referred to one of his major
works, the Gastropoda der Deutschen Tiefsee-Expedition, which was finally
published in 1925.

In having published so many papers, especially those in disparate
and obscure journals, Thiele sometimes apparently forgot taxa (e.g.,
Prochiton) or subsequently decided to correct or emend in odd or
inappropriate places (e.g., footnote in 1890b, index of 1931c).

ASSESSMENT OF THE TAXA

Probably the most important result of delineating all of Thiele’s
genus-group taxa is that an evaluation of his contribution can be more
accurately made based on the work of later authors. As we remarked
earlier, Thiele was one of the greatest researchers in malacology in the
early twentieth century (Bieler & Boss, 1989); however, even he was
aware that some of the genus-group names he introduced were unneces-
sary emendations or outright synonyms.

We assessed the subsequent status of Thiele’s genus-group names
on their treatment in several authoritative works in malacology. For each
class or subclass, we utilized the following works: Aplacophora (Boss,
1982; Salvini-Plawen, 1978); Polyplacophora (A. G. Smith, 1960; Belle,
1983; Kaas & Belle, 1980; 1985a and b; 1987; 1990); Gastropoda
Prosobranchia (Wenz, 1938-1944); Pulmonata (Zilch, 1959-1960); Opis-
thobranchia (Odhner, 1968; Pruvot-Fol, 1954); Bivalvia (Chavan, Haas,
Hertlein, Keen, or Newell in Moore, 1969; Vokes, 1980); Cephalopoda
(Nesis, 1987). We additionally referred to other sources such as Vaught
(1989).

TABULATION OF THIELE’S GENUS-GROUP NAMES

MOLLUSCA (Grand total: 291)

APLACOPHORA: Total: 11 names
11 Genus-level

POLYPLACOPHORA: Total: 39 names
1 Genus-level
3 Subgenus-level
30 Synonyms

September 30, 1991 Boss & Bieler (|

2 Unjustified emendations
2 Errors
1 Unclear

GASTROPODA: Total: 200 names

Prosobranchia: Total: 138 names
44 Genus-level
74 Subgenus-level
15 Synonyms
5 Errors
Opisthobranchia: Total: 20 names
8 Genus-level
6 Subgenus-level
5 Synonyms
1 Emendation
Pulmonata: Total: 42 names
15 Genus-level
17 Subgenus-level
10 Synonyms

BIVALVIA: ‘Total: 31 names
4 Genus-level
10 Subgenus-level
10 Synonyms
3 Unjustified emendations
3 Errors
1 Unclear

CEPHALOPODA: Total: 10 names
2 Genus-level
6 Unjustified emendations
1 Synonym
1 Error

Thiele introduced a total of 291 genus-group names; of these, 85
were recognized subsequently at the generic, and 110 at the subgeneric,
level; 71 names are now considered synonyms and 25 were unjustified
emendations or errors. Despite the great contributions made by Thiele,
almost one third of his genus-group names were superfluous. Considering
the current annual increase in the establishment of new molluscan
generic-taxa (Boss, 1989), we hope this analysis of the fate of the names
of someone of such eminent stature as Thiele will constitute a warning
to contemporary and future workers that not only are many new names
probably unnecessary synonyms but they also burden the literature,
obfuscate reality and seriously encumber one of the foremost roles of

8 NEMOURIA No. 39

our system of classification, namely information retrieval.

With more than 15,000 genus-group names now available for Recent
mollusks (Vaught, 1989) and with probably fewer than 60,000 Recent
species in the Phylum Mollusca (Boss, 1971) there are already clearly
too many genus-group names. Such an overburden of our taxonomy is
unfortunately all too well documented: Ponder and Warén (1988) list
nearly 1,000 family-group names for only part of the Prosobranchia.

ALPHABETICAL LISTING OF FAMILIES WITH
TREATED GENUS-GROUP NAMES
(reflecting original and subsequent family-level placements):

APLACOPHORA

Acanthomeniidae: Acanthomenta

Amphimentiidae: Amphimenia

Chaetodermatidae: Metachaetoderma, Prochaetoderma

Dondersiidae: Heathia

Hemimeniidae: Archaeomenia

Lepidomentiidae: Acanthomenia, Heathia, Notomenia, Phyllomenia,
Sandalomenia

Limifossoridae: Metachaetoderma

Neomeniidae: Archaeomenia

Pararhopaliidae: Metamenia

Phylomeniidae: Phyllomenia

Prochaetodermatidae: Prochaetoderma

Proneomeniidae: Amphimenia, Metamenia, Pruvotia

Rhopalomeniidae: Pruvotia

POLYPLACOPHORA
Acanthochitonidae: Acanthochiton, Aristochiton, Mecynoplax,
Thaumastochiton

Callistoplacidae: Notochiton

Callochitonidae: Icoplax, Mopaliella

Chaetopleuridae: Calloplax, Helioradsia

Chitonidae: Adriella, Amaurochiton, Anthochiton, Chondroplax, Clathro-
pleura, Diochiton, Georgus, Helioradsia, Icoplax, Leptopleura,
Lophyriscus, Lophyropsis, Mecynoplax, Mopahopsis, Onithoplax,
Poeciloplax, Prochiton, Radsiella, Rhodoplax, Rhopalopleura,
Rhyssoplax, ‘‘Slerochiton,’’ Stereoplax, Sypharochiton, Toniciella,
Toniciopsis, Triboplax

Cryptoplacidae: Acanthochiton, Anistochiton, Thaumastochiton

Ischnochitonidae: Adriella, Anthochiton, Beanella, Calloplax, Chondropleura,

September 30, 1991 Boss & Bieler 9

Helioradsia, Icoplax, Lepidochiton, Lepidopleuroides, Lep-
topleura, Lophyriscus, Lophyropsis, Mopaliella, Mopalhop-
sis, Notochiton, Radsiella, Rhodoplax, Stereoplax, ‘‘Toni-
ciella,’’ Tonicina

Lepidochitonidae: Beanella, Lepidochiton, Mopattella

Lepidopleuridae: Parachiton

Leptochitonidae: Beanella, Lophyropsis, Notochiton, Parachiton

BIVALVIA

Arcidae: Noetiella, Paranoetia

Cardiidae: Microcardium

Cyamiidae: Ptychocardia

Dreissenidae: Sinomytilus

Ledidae: Silicula

Limopsidae: Hochstetterina, Limopsilla, Lissarcula

Mactridae: Electromactra

Mutelidae: fheringella

Mytilidae: Sinomytilus

Noetiidae: Paranoetia, Noetiella

Nuculidae: Brevinucula, Lionucula

Ostreidae: Dioeciostraea, Monoeciostraea

Pectinidae: Adamussium, Arctinula

Philobryidae: Hochstetterina

Semelidae: ‘‘Punigapia’’

Tellinidae: Arcopella, Clathrotellina, Hemimetis, Pistris, Striotellina, Tellinan-
gulus

Unionidae: faronia, Luteacarnea, Plagiolopsis, Pustulosa, Venustaconcha

Verticordiidae: Acreuciroa, ‘‘Lyonstela”’

Vesicomyidae: Ptychocardia

GASTROPODA
Prosobranchia

Aclidae: Trochaclis

Aclididae: ‘‘Pherusiana’’

Adeorbidae: Mérchiella [Moerchiella], Mérchinella [Moerchinella]

Amphiperatidae: see Cypraeidae

Assimineidae: Austropilula, Conacmella, Crossilla, Leucostele, Macrassiminea,
Metassiminea, Nanivitrea, Oriella, Ovassiminea, Pseudas-
siminea, Pseudocyclotus, Rupacilla, Sculptassiminea, Suterilla,
Turbacmella

Bithyniidae: Emmericiopsis

10 NEMOURIA INoir39

Buccinidae: Appisania, Bathydomus, Buccinella, Chlanidotella, Eosipho,
Ficulina, Fusinella, Macroniscus, Meteuthnia, Notoficula, Pro-
buccinum, Proneptunea, Prosipho

Bulimidae: Emmericiopsis

Bursidae: Bufonariella

Cancellariidae: Mericella

Capulidae: ‘‘Amathinoidas’’

Cerithiidae: Cerithiopsilla, Eumetula, Ischnocerithium, Proclava

Cerithiopsidae: Cerithiopsilla, Eumetula, Proseila

Cingulopsidae: Eatonina, Eatoniopsis, Watsonella

Columbellidae: Cilara, Metanachis, Paratilia, Pseudanachts

Cominellidae: Meteuthria

Conidae: Acrobela, Anticlinura, Benthomangelia, Citharella, Clinuropsis,

Conasprella, Fusiturris, Micropleurotoma, Ptychobela, Ptychosyrinx,
Strioconus, Typhlosyrinx

Cyclostomidae: Pseudocyclotus

Cyclostrematidae: Bothropoma, Chunula

Cymatiidae: Charoniella

Cypraeidae: ‘“‘Cleovitrea,’’? Pediculariella, Primovula, Procalpurnus

Epitoniidae: Chuniscala, Cirsotremopsis, Opalhopsts

Fossaridae: Fossarella, Iphitella

Hydrobiidae: Caspiella, Emmericiopsis, Hydrococcus, Indopyrgus, Lithoglyphop-

sis, Pachydrobtella, Probythinella, Walkerilla

Hydrococcidae: Hydrococcus

Lacunidae: Carinolacuna

Lepetellidae: Cocculinella

Littorinidae: Iphitella, Lissoconchus

Lyocyclidae: Lyocyclus

Melaniidae: Cubaedomus, Pseudocleopatra

Micromelaniidae: Caspiella

Muricidae: Drupella, Gracilimurex

Nassarlidae: Naytiopsis, Phicarcularia, Profundinassa, Pronentula

Nassidae: see Nassariidae

Naticidae: Frovina, Prolacuna, Sublacuna

Olividae: Gracilancilla, Olivellopsis, Parvoliva, Zemiropsts

Ovulidae: Primovula, Procalpurnus

Patellidae: Patellidea, Patellona, Patellopsis, Patinastra

Pediculariidae: Pediculariella

Planaxidae: Proplanaxis, Supplanaxts

Pleurotomidae: see Turridae

Pomatiasidae: Pseudotopoma

Potamididae: Batillariella, Cerithideopsilla, Cerithideopsis, ‘‘Zeacumanthus”’

September 30, 1991 Boss & Bieler 11

Pyramidellidae: Bacteridium, Ebalina, Euparthenia, Oopyramis, Stylopyramis

Ranellidae: Charoniella

Rissoellidae: Feffreysiella, Feffreysilla, Feffreysina, fFeffreysiopsis

Rissoidae: Alcidiella, Boogina, Eatonina, Eatoniopsis, Watsonella

Scalidae: see Epitoniidae

Skeneopsidae: Microdiscula

Stomatiidae: Pseudostomatella

Strombidae: Aliger, Rostellariella

Thiaridae: Cubaedomus, Pseudocleopatra

Tornidae: Mérchinella [Moerchinella]

Trichotropidae: Lyocyclus

Triphoridae: Euthymella

Trochaclididae: Trochaclis

Trochidae: Callumbonella, Calotropis, Margarella, Margaritella, Margaritop-

sis, Morchiella [Moerchiella], Nanula, Prothalotia, Pseudosto-
matella, Umbotrochus

Turbinidae: Bothropoma, Subninella

Turridae: Acrobela, Anticlinura, Benthomangelia, Citharella, Clinuropsis,
Fusiturris, Maicropleurotoma, Ptychobela, Ptychosyrinx, Typhlosy-
rinx

Turritellidae: Mesaliopsis, Protomella

Vermetidae: Segmentella

Vitrinellidae: Microdiscula

Viviparidae: Callinina

Volutidae: Harpovoluta

Opisthobranchia
Actaeonidae: Metactaeon, Neactaeonina, Pseudactaeon
Atyidae: Haminella

Cadlinidae: Cadlinella
Charcotiidae: Pseudotritonia
Coryphellidae: Himatina
Diaphanidae: Odostomiopsis
Dorididae: Cadlinella
Dotonidae: Jduliella
Echinochilidae: Cadlinella
Flabellinidae: Himatina
Goniaeolididae: Goniaeolis
Haminoeidae: Haminella
Hedylidae: Hedylopsis, Parhedyle
Hedylopsidae: Hedylopsis
Iduliidae: Iduliella

12 NEMOURIA No. 39

Lamellidoridae: Prodoridunculus
Microhedylidae: Parhedyle

Notaeolidiidae: Pseudotritonia
Notodiaphanidae: Diaphanella, Notodiaphana
Onchidoridae: Prodoridunculus

Oxynoidae: Lophopleura

Philinidae: Lophopleura

Pleurobranchidae: Pleurobranchella
Polyceridae: Prodoridunculus

Retusidae: Pyrunculus, Semiretusa, Utriculastra
Scaphandridae: Sphaerocylichna

Pulmonata
Ariophantidae: Ariocaelatura, Conibycus, Durgellina, Hemiglyptopsis, Parasi-
tala, Rhinocochlis, Sitalina, Sitalinopsis, Striokaliella

Bulimulidae: Macrodontopsis, Scansicochlea

Camaenidae: Baudinella

Clausiliidae: Proreinia, Roseniella

Ellobiidae: Cassidulella, Cassidulina

Endodontidae: Garrettina, Nesodiscus, Notodiscus, Paryphantopsis

Enidae: Paracerastus, Rhachidina, Zebrinops

Euconulidae: Conibycus, Durgellina, Parasitala, Sitalina

Haplotrematidae: Proselenites

Helicarionidae: Ariocaelatura, Hemiglyptopsis, Sitalinopsts

Lymnaeidae: Pseudisidora

Odondostomidae: Macrodontopsis

Orculidae: Pagodinella

Planorbidae: Afroplanorbis

Pleurodontidae: Baudinella

Polygyridae: Daedalocheila

Streptaxidae: Afristreptaxis, Augustula, Macrogonaxis, Mirellia, Pauciden-
tella, Pseudogonaxts

Subulinidae: Ochrodermatina, Striosubulina

Systrophiidae: Microhappia, Prohappia, Proselenites

Urocoptidae: Spiraxilla

Vaginulidae: Novovaginula

Vertiginidae: Pagodinella

Vitrinidae: Arabivitrina

Zonitidae: Geodiaphana

CEPHALOPODA
Alluroteuthidae: Parateuthis

September 30, 1991 Boss & Bieler 13

Gonatidae: ‘‘Berryiteuthis”’

Psychroteuthidae: Psychroteuthis

Sepiidae: Acanthosepia, Ascarosepia, Doratosepia, Lophosepia, Rhombosepia,
Spathidosepia

Vampyroteuthidae: Hymenoteuthts

ALPHABETICAL LISTING OF GENUS-GROUP NAMES:

Acanthochiton Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 14; emenda-
tion pro Acanthochitona Gray 1821, as genus; A. G. Smith 1960:
169 credited the spelling to Iredale 1915 and listed the name as a
‘nomen vanum.’? Cryptoplacidae [Acanthochitonidae, teste A. G.
Smith 1960: 169]. Polyplacophora.

Acanthomenia Thiele 1913d.
Deutsche Stidpolar-Expedition 1901-1903, 14(6)(1): 61; as new
genus. Type-species by monotypy: Acanthomenia gaussiana Thiele
1913. Lepidomeniidae [Acanthomeniidae, teste Boss 1982: 952].
Aplacophora.

Acanthosepia Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 954; emenda-
tion pro Acanthosepion Rochebrune 1884; as section of Sepza
Linnaeus 1758. Sepiidae. Cephalopoda.

Acreuciroa Thiele in Thiele & Jaeckel 1931b.

Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition . . .
*Valdivia’’, 21(1): 250 (92); as new section of Euciroa Dall 1881
[subgenus of Euciroa, teste Keen in Moore 1969: N857]. Type-
species by monotypy: Euciroa (Acreuciroa) rostrata Thiele in Thiele
& Jaeckel 1931b. Subsequently attributed to Jaeckel & Thiele,
Thiele & Jaeckel, and to Thiele alone by various authors. Verticor-
diidae. Bivalvia.

Acrobela Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
“Valdivia”, 17(2): 238 [204]; as new subgenus of Bela Gray 1847
[genus, teste Thiele 1929: 362]. Type-species by original designa-
uon: Bela (Acrobela) optima Thiele 1925. Pleurotomidae [Conidae,
teste Thiele 1929: 362; Turridae, teste Wenz 1943: 1413].
Gastropoda: Prosobranchia.

Adamussium Thiele 1934.

14 NEMOURIA No. 39

Handbuch der systematischen Weichtierkunde, 2(3): 807; as new
genus. Type-species by monotypy: Adamussium colbecki (E. Smith)
[Pecten colbecki E. A. Smith 1902: 212, pl. 25 fig. 11]. Pectinidae.
Bivalvia.

Adriella Thiele in Troschel & Thiele 1893e.

Das Gebiss der Schnecken, 2(8): 391; as new genus [= Trachyder-
mon Carpenter 1864, teste Thiele 1909d: 4; = Lepidochitona Gray
1821, s.s., teste Thiele 1929: 8]. Type-species by monotypy: Adriella
variegata Philippi [Chiton variegatus Philippi 1836, p. 107; 1844:
83, pl. 19 fig. 13; non Réding 1798, nec de Blainville 1825, nec
Leach 1852; = Lepitochitona cinerea (Linnaeus 1767), teste Kaas &
Belle 1980: 138]. Chitonidae [Ischnochitonidae, teste A. G. Smith
1960: 156, Belle 1983: 96]. Polyplacophora.

Afristreptaxis Thiele 1932a.
Archiv fir Molluskenkunde, 64(1): 12; as new genus [= Eustreptaxis
L. Pfeiffer 1878, teste Zilch 1960: 559]. Type-species by original
designation: wosseler1 (Thiele) [Streptaxis vosseleri Thiele 1911].
Streptaxidae. Gastropoda: Pulmonata.

Afroplanorbis Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 480; as new
section of Planorbis O. F. Miller 1774. Type-species by original
designation: sudanicus Martens [Planorbis sudanicus Martens 1870:
35]. Planorbidae. Gastropoda: Pulmonata.

Alcidiella Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 163; nomen
novum pro Alcidia Monterosato 1890, non Westwood 1879, nec
Bourguignat 1889 [Thiele apparently renamed this without knowing
of the emendation to Alcidiella by Cossmann (1921)]. Rissoidae.
Gastropoda: Prosobranchia.

Aliger Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 254; as new
section of Strombus (Strombus) Linnaeus 1758. Type-species by
monotypy: Strombus (Aliger) gallus Linnaeus [Strombus gallus Lin-
naeus 1758: 743]. Strombidae. Gastropoda: Prosobranchia.

‘“Amathinoidas’’ [1929].
Handbuch der systematischen Weichtierkunde, 1(1): 246; error pro
Amanthinoides Sacco, 1896; as subgenus of Thyca H. & A. Adams
1854. Capulidae. Gastropoda: Prosobranchia.

September 30, 1991 Boss & Bieler 15

Amaurochiton Thiele in Troschel & Thiele 1893e.

Das Gebiss der Schnecken, 2(8): 362; as new genus [= Chiton
Linnaeus 1758, s.s., teste Thiele 1909d: 2, and Belle 1983: 123;
= section of Chiton s.s., teste Thiele 1929: 19; subgenus of Chiton,
teste A. G. Smith 1960: 166]. Type-species by subsequent designa-
tion (Kobelt 1894: 26): Chiton olivaceus [Frembly 1827: 199,
Suppl. pl. 16 fig. 4], non Spengler 1797, [= striatus Barnes 1824,
non Brusina 1870; = magnificus Deshayes 1822, = latus G. B.
Sowerby I 1825, non Lowe 1825, nec Leach 1852, = murray
Haddon 1886, teste Kaas & Belle 1980: 125]. Chitonidae. Polyplaco-
phora.

Amphimenia Thiele 1894b.
Zeitschrift fiir wissenschaftliche Zoologie, 58(2): 244, 272; as new
subgenus of Proneomenia Hubrecht 1880 [genus, teste Thiele 1913e:
x, 39]. Type-species by monotypy: Proneomenia neapolitana ‘Vhiele
1894. [Proneomentidae, teste Thiele, 1913e: x, 39; Amphimentidae,
teste Boss 1982: 952]. Aplacophora.

Anthochiton Thiele in Troschel & Thiele 1893e.

Das Gebiss der Schnecken, 2(8): 377; as new genus [= Chiton
(Clathropleura), teste Thiele 1909d: 3; = section of Chiton (Rhys-
soplax) Thiele 1893, teste Thiele 1929: 20; = Chiton (Rhyssoplax),
teste A. G. Smith 1960: 165, Belle 1983: 124]. Type-species by
monotypy: Anthochiton tulipa Quoy & Gaimard [Chiton tulipa Quoy
& Gaimard 1835: 389, pl. 74 figs. 35, 36 = C. tulipa alfredensis
Ashby 1828, teste Kaas & Belle 1980: 135]. Chitonidae [Ischnochi-
tonidae, teste Belle 1983: 124]. Polyplacophora.

Anticlinura Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 1002; nomen
novum pro Clinuropsis Thiele 1929 (q.v.), non Clinuropsis E. Vincent
1913; [subgenus of Pleurotomella, teste Wenz 1943: 1460]. Conidae
[Turridae, teste Wenz 1943: 1460]. Gastropoda: Prosobranchia.

Appisania Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 314; as new
section of Metula H. & A. Adams 1853 [subgenus of Metula, teste
Wenz 1941: 1195]. Type-species by monotypy: Metula (Appisania)
montrouziert (Crosse) [Pisania montrouzieri Crosse 1862: 251, pl.
10 fig. 5 [sic, 7]]. Buccinidae. Gastropoda: Prosobranchia.

Arabivitrina Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 600; as new

16 NEMOURIA No. 39

section of Vitrina Draparnaud 1801 [subgenus of Phenacolimax, teste
Zilch 1959: 238]. Type-species by monotypy: Vitrina (Arabivitrina)
arabica Thiele [Vitrina arabica Thiele 1910]. Vitrinidae. Gastropoda:
Pulmonata.

Archaeomenia Thiele 1906c.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
“Valdivia’’, 9(2): 315 ff. [1 ff.]; as new genus. Type-species by
monotypy: Archaeomenia prisca Thiele 1906. [Neomentidae, teste
Thiele 1913e: x, 27; Hemimeniidae, teste Boss 1982: 950].
Aplacophora.

Arcopella Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 914; as new
section of Arcopagia (Arcopagia) T. Brown 1827 [listed by Thiele
as “‘Sectio Arcopella Monterosato in sched.”; subgenus of Tellina
Linnaeus 1758, teste Keen im Moore, 1969: N614]. Type-species
by monotypy: Arcopagia (Arcopella) balaustina (Linné) [Tellina
balaustina Linnaeus, 1758: 676]. Tellinidae. Bivalvia.

Arctinula Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 806; as new
section of Propeamussium (Palliolum) Monterosato 1884 [synonym
of Pecten (Delectopecten) Stewart 1930, teste Hertlein in Moore
1969: N354]. Type-species by monotypy: Propeamussium (Arctinula)
groenlandicum [sic] (Sowerby) [Pecten greenlandicus G. B. Sowerby
II 1842: 57, pl. 13 fig. 40]. Pectinidae. Bivalvia.

Ariocaelatura ‘‘Germain’’ Thiele 1931c.

Handbuch der systematischen Weichtierkunde, 1(2): 615; non
Germain 1921 [Zilch (1959: 307) listed Pseudocaelatura Wenz 1947,
with Ariocaelatura Thiele 1931, non Germain 1921, as a synonym;
Zilch (1960: 730) noted that Pseudocaelatura Wenz 1947 was
preoccupied by Pseudocaelatura Germain 1921, and replaced the
name by Dancea; subgenus of Dancea Zilch 1960, teste Vaught
1989: 96]. See Introduction, p. 5. Ariophantidae [Helicarionidae,
teste Zilch 1959: 307]. Gastropoda: Pulmonata.

Aristochiton Thiele 1909d.
Zoologica, 22(56/1): 35; as new genus [= Craspedochiton Shuttle-
worth 1853, teste A. G. Smith 1960: 169; = Notoplax H. Adams
1861, teste Kaas & Belle 1980: 60]. Type-species by monotypy:
Aristochiton hirtus Thiele 1909. [Cryptoplacidae, teste Thiele 1909i:
116; Acanthochitonidae, teste A. G. Smith 1960: 169, Belle 1983:

September 30, 1991 Boss & Bieler 17

144]. Polyplacophora.

Ascarosepia Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 954; emenda-
tion pro Ascarosepion Rochebrune 1884; as section of Sepia Lin-
naeus, 1758. Sepiidae. Cephalopoda.

Augustula Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 727; as new
genus. Type-species by monotypy: Imperturbatia braueri Martens
[Streptaxis (Imperturbatia) braueri Martens in Martens & Wiegmann
1898: 13, pl. 2 fig. 8]. Streptaxidae. Gastropoda: Pulmonata.

Austropilula Thiele 1927a.
Zoologische Jahrbticher, 53(1/3): 122, 126; as new section of
Assiminea Fleming 1828 [subgenus of Assiminea, teste Wenz 1939:
634]. Type-species by original designation: Assiminea beddomeana
Nevill [G. Nevill 1880: 163; 1881a: 158, pl. 7 fig. 3]. Assimineidae.
Gastropoda: Prosobranchia.

Bacteridium Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 236; as new
subgenus of Eulimella Gray 1847 [genus, teste Wenz 1940: 867].
‘Type-species by monotypy: Eulimella (Bacteridium) praeclara Thiele
[Eulimella praeclara Thiele 1925]. Pyramidellidae. Gastropoda:
Prosobranchia.

Bathydomus Thiele 1912b.
Deutsche Siidpolar-Expedition 1901-1903, 13(5): 247; as new
genus [section of Chlanidota Martens 1878, teste Thiele 1929: 315;
subgenus of Chianidota, teste Wenz 1943: 1201]. Type-species by
monotypy: Bathydomus obtectus Thiele 1912. [Buccinidae, teste
Thiele 1929: 315]. Gastropoda: Prosobranchia.

Batillariella Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 208; as new
subgenus of Batillaria Benson 1842. Type-species by monotypy:
Batillaria (Batillariella) estuarina (Tate) [Bittium estuarinum Tate
1893: 190, pl. 5 fig. 12]. Potamididae. Gastropoda: Prosobranchia.

Baudinella Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 685; as new
section of Angasella A. Adams 1863 [subgenus of Angasella, teste
Zilch 1960: 619]. Type-species by monotypy: Angasella (Baudinella)
baudinensis (E. Smith) [Helix (Gonostoma) baudinensis E. A. Smith

18 NEMOURIA No. 39

1893: 97]. Pleurodontidae [Camaenidae, teste Zilch 1960: 619].
Gastropoda: Pulmonata.

Beanella “‘Dall’’ Thiele 7x Troschel & Thiele 1893e.

Das Gebiss der Schnecken, 2(8): 388; non Dall 1882 [as synonym,
listed as Beanella Dall 1881, nomen nudum, of Middendorffia Dall
1882; Boss, Rosewater & Ruhoff 1968, listed Beanella Dall as of
1882, a new name for Beania ‘Carpenter’? Dall 1879; Beanella
Thiele = Ischnochiton Gray 1847, teste A. G. Smith 1960: 155; =
Leptochiton Gray 1847, s.s., teste Belle 1983: 52, with rissoz
Payraudeau 1826 [= Chiton algesirensis Capellini, 1859] as type
species by subsequent designation]. [Lepidochitonidae, teste Thiele
1929: 9; Ischnochitonidae, teste A. G. Smith 1960: 155; Leptochi-
tonidae, teste Kaas & Belle 1985: 36]. See Introduction, p. 5.
Polyplacophora.

Benthomangelia Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
‘Valdivia’, 17(2): 208 [174], 220, 225; as new section of Mangelia
Risso 1826 [subgenus of Cythara Schumacher 1817, teste Thiele
1929: 367]. Type-species by original designation: Mangelia tro-
phonoidea (Schepman) [Surcula trophonoidea Schepman 1911 [1913]:
426, pl. 28 fig. 3]. Pleurotomidae [Conidae, teste Thiele 1929:
367; Turridae, teste Wenz 1943: 1437]. Gastropoda: Prosobranchia.

‘‘Berryiteuthis’’ [1926b].
Handbuch der Zoologie, 5(3): 252; error pro Berryteuthis Naef
1923. Gonatidae. Cephalopoda.

Boogina Thiele 1913a.
Zentralblatt fiir Zoologie, 2(3): 86; nomen novum pro Watsonella
Thiele 1912 (g.v.), non Grabau 1900. Rissoidae. Gastropoda:
Prosobranchia.

Bothropoma Thiele 1924a.
Mitteilungen aus dem Zoologischen Museum in Berlin, 11(1): 63,
71; as new genus. Type-species by monotypy: Bothropoma isseli
Thiele 1924. Turbinidae [Cyclostrematidae, teste Thiele 1925d:
76]. Gastropoda: Prosobranchia.

Brevinucula Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 786; as new
subgenus of Nucula Lamarck 1799 [genus, teste Keen 22 Moore
1969: N231]. Type-species by monotypy: Nucula (Brevinucula)

September 30, 1991 Boss & Bieler 19

guineensis Thiele [Nucula guineensis Thiele & Jaeckel 1931].
Nuculidae. Bivalvia.

Buccinella Thiele 1912b.
Deutsche Stidpolar-Expedition 1901-1903, 13(2): 244, 246; as new
genus [non Buccinella Perry 1811; Buccinella Thiele replaced by
Fusinella Thiele 1917, (g.v.)]. Type-species by original designation:
Buccinella jucunda Thiele 1912. Buccinidae. Gastropoda: Prosobran-
chia.

Bufonariella Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 284; “‘nomen
novum”’ [error pro “‘new section’’?]; listed as section [subgenus of
Bursa Réding 1798, teste Wenz 1941: 1072]. Type-species by
monotypy: Bursa (Bufonariella) scrobiculator (Linnaeus) [Murex
scrobiculator Linnaeus 1758]. Bursidae. Gastropoda: Prosobran-
chia.

Cadlinella Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 431; as new
genus. Type-species by monotypy: Cadlinella ornatissima (Risbec)
[Cadlina ornatissima Risbec 1928: 163, text-fig. 47, pl. 8 fig. 4].
Dorididae [Echinochilidae, teste Odhner in Franc, 1968: 866;
Cadlinidae, teste Vaught 1989: 89]. Gastropoda: Opisthobranchia.

Callinina Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 747; nomen
novum pro Callina Hannibal 1912 [as listed by Thiele 1929: 114],
non Lowe 1855; subgenus of Viviparus Montfort 1810, teste Wenz
1939: 490]. See also Tomlin 1936: 136. Viviparidae. Gastropoda:
Prosobranchia.

Calloplax Thiele 1909d.
Zoologica, 22(56/1): 19; as new genus [subgenus of Callistochiton
Dall 1879, teste Belle 1983: 82]. Type-species by monotypy: Chiton
janetrensis Gray 1828 [p. 6, pl. 3 fig. 8]. [Ischnochitonidae, teste
Thiele 19091: 116; Chaetopleuridae, teste A. G. Smith 1960: 162].
Polyplacophora.

Callumbonella Thiele 1924a.
Mitteilungen aus dem Zoologischen Museum in Berlin, 11(1): 58,
69; nomen novum pro Umbotrochus (Monterosato MS) Thiele 1924
(q.v.), non Perner 1904. Trochidae. Gastropoda: Prosobranchia.

Calotropis Thiele 1929.

20

NEMOURIA No. 39

Handbuch der systematischen Weichtierkunde, 1(1): 49; nomen
novum pro Calliotropis Oliver 1926, non Seguenza, 1903; as section
of Calliostoma Swainson 1840, s.s. Trochidae. Gastropoda: Proso-
branchia.

Carinolacuna Thiele 1929.

Handbuch der systematischen Weichtierkunde, 1(1): 122; as new
subgenus of Lacuna Turton 1827 [genus, teste Wenz 1939: 513].
Type-species by monotypy: Lacuna (Carinolacuna) carinifera A.
Adams [Lacuna carinifera A. Adams 1851 [1853]: 225]. Lacunidae.
Gastropoda: Prosobranchia.

Caspiella Thiele 1928c.

Zoologische Jahrbticher, 55(5-6): 353, 381; as new genus. Type-
species by original designation: Rissoa conus Eichwald [1838: 155].
Micromelaniidae or Hydrobiidae [?Micromelaniidae, teste Thiele
1929: 159; Micromelaniidae, teste Wenz 1939: 606]. Gastropoda:
Prosobranchia.

Cassidulella Thiele 1934.

Handbuch der systematischen Weichtierkunde, 2(3): 1004; nomen
novum pro Cassidulina Thiele 1931 (g.v.); non Cassidulina Orbigny
1826; non Cassidulella Strand 1928, which was replaced by
Cassidulta Strand 1942 [= Cassidulta Strand 1942, teste Zilch 1959:
76]. Ellobiidae. Gastropoda: Pulmonata.

Cassidulina Thiele 193lc.

Handbuch der systematischen Weichtierkunde, 1(2): 467; as new
subgenus of Cassidula Férrussac 1821 [non Cassidulina Orbigny
1826; Cassidulina Thiele replaced by Cassidulella Thiele 1934 (q.v.);
= Cassidulta Strand 1942, teste Zilch 1959: 76]. Type-species by
monotypy: Cassidula (Cassidulina) doliolum (Petit) [Auricula doliolum
Petit de la Saussaye, 1843: 201]. Ellobiidae. Gastropoda: Pul-
monata.

Cerithideopsilla Thiele 1929.

Handbuch der systematischen Weichtierkunde, 1(1): 206; as new
section of Cerithidea (Cerithideopsis) Thiele 1929 (q.v.) [subgenus
of Cerithidea Swainson 1840, teste Wenz 1940: 742]. Type-species
by monotypy: Cerithidea (Cerithideopsilla) fluviatis (Potiez & Michaud)
[Cerithium fluviatile Potiez & Michaud 1838: 363, pl. 31 figs. 19,
20; = Murex cingulata Gmelin 1791, teste Wenz 1940: 742].
Potamididae. Gastropoda: Prosobranchia.

Cerithideopsis Thiele 1929.

September 30, 1991 Boss & Bieler 21

Handbuch der systematischen Weichtierkunde, 1(1): 206; as new
subgenus and section of Cerithidea Swainson 1840 [subgenus of
Cerithidea, teste Wenz 1940: 742]. Type-species by monotypy:
Cerithidea (Cerithideopsis) iostoma (Pfeiffer) [Potamides iostomus L.
Pfeiffer 1839: 357]. Potamididae. Gastropoda: Prosobranchia.

Cerithiopsilla Thiele 1912b.
Deutsche Stidpolar-Expedition 1901-1903, 13(2): 202, 204; as new
subgenus of Cerithiopsis Forbes & Hanley 1849, or as new genus
[subgenus of Cerithiopsis, teste Thiele 1925b: 121 [87]]. Type-
species by original designation: Cernthiopsilla cincta Thiele 1912.
Cerithiopsidae [Cerithiidae, teste Powell 1960: 140]. Gastropoda:
Prosobranchia.

Charoniella Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 283; as new
section of Charonia Gistel 1848; non Charoniella Powell & Bartrum
1929; [synonym of Negyrina Iredale 1929, teste Wenz 1941: 1066].
‘Type-species by monotypy: Charonia (Charoniella) subdistorta (La-
marck) [Triton subdistortum Lamarck 1822: 186]. Cymatiidae [=
Ranellidae]. Gastropoda: Prosobranchia.

Chlanidotella Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 317; as new
subgenus of Thalassoplanes Dall 1908. Type-species by monotypy:
Thalassoplanes (Chlanidotella) modesta (Martens) [Cominella modesta
Martens 1885: 91]. Buccinidae. Gastropoda: Prosobranchia.

Chondroplax Thiele in Troschel & Thiele 1893e.
Das Gebiss der Schnecken, 2(8): 363; as new genus [section of
Chiton, teste Thiele 1909d: 2, 5; = Chiton s.s. Linnaeus 1758, teste
A. G. Smith 1960: 165; Belle 1983: 123]. Type-species by
subsequent designation (Kobelt 1894: 26): Chiton granosus [Frembly
1827: 200, suppl. pl. 17 fig. 1]. Chitonidae. Polyplacophora.

Chondropleura Thiele 1906d.

Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
“Valdivia”, 9(2): 334; as new subgenus of Ischnochiton Gray 1847
[= Ischnochiton (Stenosemus) Middendorff 1847, teste Belle 1980:
69; Kaas & Belle 1990: 60]. Type-species by subsequent designation
(A. G. Smith 1960: 156): Lophyrus exaratus G. O. Sars 1878 [p.
113, pl. 3 fig. 1]. [Ischnochitonidae, teste Thiele 1929: 18].
Polyplacophora.

Chuniscala Thiele 1928d.

Ze NEMOURIA No. 39

Zeitschrift fiir wissenschaftliche Zoologie, 132: 93; new subgroup
of Scala (Klein 1753) Bruguiére 1792 [= Epitonium Réding 1798]
[section of Scala (Cirsotrema) Morch 1852, teste Thiele 1929: 222;
subgenus of Cursotrema, teste Wenz 1940: 798]. Type-species by
original designation: Scala agulhasensis Thiele 1925. Scalidae [=
Epitoniidae]. Gastropoda: Prosobranchia.

Chunula Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition . .
*Valdivia’’, 17(2): 62 [28], 58 [24]; as new genus [nom. nud. in
Thiele 1924a: 60, 71]. Type-species by original designation:
Chunula typica Thiele 1925. Cyclostrematidae. Gastropoda: Proso-
branchia.

Cilara Thiele 1924b.
Archiv fiir Molluskenkunde, 56(5): 202, 210; as new section of
Pyrene (Muitrella) Risso 1826 [subgenus of Mitrella, teste Wenz 1941:
1139]. Type-species by monotypy: Buccinum secalinum Philippi
[1846: 53]. Columbellidae. Gastropoda: Prosobranchia.

Cirsotremopsis Thiele 1928d.

Zeitschrift fiir wissenschaftliche Zoologie, 132: 92; as new section
of Scala (Klein 1753) Bruguiére 1792 [= Epitonium Roding 1798]
[sect. of Scala (Cirsotrema) Mérch 1852, teste Thiele 1929: 222;
subgenus of Cursotrema, teste Wenz 1940: 795]. Type-species by
monotypy: Scalaria cochlea Sowerby [G. B. Sowerby II 1844: 103,
pl. 35 fig. 142]. Scalidae [= Epitoniidae]. Gastropoda: Prosobran-
chia.

Citharella Thiele 1929.

Handbuch der systematischen Weichtierkunde, 1(1): 366; probable
emendation pro Cyharella Monterosato 1875; as section of subgenus
Cythara Schumacher 1817 s.s. [Wenz 1943: 1432 listed this as
Cytharella, a subgenus of Cythara; the spelling Cyharella by Mon-
terosato was considered a lapsus calami by Wenz (1943: 1432)].
Conidae [Turridae, teste Wenz 1943: 1432]. Gastropoda: Proso-
branchia.

Clathropleura ‘Tiberi’? Thiele in Troschel & Thiele 1893e.
Das Gebiss der Schnecken, 2(8): 367; non Tiberi 1877 [Clathro-
pleura ‘Tiberi 1887 part. = Chiton (Rhyssoplax) Thiele 1893, teste
Thiele 1929: 20; Clathropleura Thiele 1893, non Tiberi 1877, =
Rhyssoplax, teste Belle: 90]. Type species by original designation:
Chiton siculus Gray 1828 [p. 5]. See Introduction, p. 5. Polyplaco-

September 30, 1991 Boss & Bieler 23

phora.

Clathrotellina Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 917; as new
subgenus of Merisca Dall 1900 [subgenus of Tellina Linnaeus 1758,
teste Keen 1969: N615]. Type-species by monotypy: Merisca
(Clathrotellina) pretiosa (Deshayes) [Tellina pretiosa Deshayes 1855:
360, non Eichwald 1830; = T: pretium Salisbury 1934]. Tellinidae.
Bivalvia.

*‘Cleovitrea’’ [1931c].
Handbuch der systematischen Weichtierkunde, 1(2): 739; error pro
Cleotrivia Iredale 1930 [corrected by Thiele 1934: 1010]. Cypraeidae
[Triviidae, teste Vaught 1989: 34]. Gastropoda: Prosobranchia.

Clinuropsis Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 372; as new
section of Pleurotomella Verrill 1873 [non Clnuropsis E. Vincent
1913; Clinuropsis Thiele replaced by Anticlinura Thiele 1934, (g.v.)].
Type-species by original designation: Clinura monochorda Dall
[1908: 214, 292, pl. 13 fig. 1]. Conidae [Turridae, teste Wenz
1943: 1460]. Gastropoda: Prosobranchia.

Cocculinella Thiele 1909g.
Systematisches Conchylien-Cabinet von Martini und Chemnitz,
2(1la): 21; as new genus. Type-species by monotypy: Acmaea
minutissima E. Smith 1904 [E. A. Smith 1904b: 4]. Lepetellidae.
Gastropoda: Prosobranchia.

Conacmella Thiele 1927a.
Zoologische Jahrbticher, 53: 123, 126; as new genus. Type-species
by original designation: Acmella vagans Pilsbry [(Pilsbry MS) Thiele
1927]. Assimineidae. Gastropoda: Prosobranchia.

Conasprella Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 373; as new
subgenus of Conus Linnaeus 1758 [subjective synonym of Hemiconus
Cossmann 1889, teste Wenz 1943: 1466]. Type-species by mono-
typy: Conus (Conasprella) cancellatus Bruguiére [Conus cancellatus
Hwass in Bruguiére 1792: 712-713]. Conidae. Gastropoda: Proso-
branchia.

Conibycus Thiele 1928a.
Zoologische Jahrbiicher, 55(1-3): 136; as new genus. Type-species
by original designation: Conibycus dahli Thiele 1928. Ariophantidae

24 NEMOURIA No. 39

f[Euconulidae, teste Zilch 1959: 282]. Gastropoda: Pulmonata.

Crossilla Thiele 1927a.
Zoologische Jahrbticher, 53(1-3): 123, 126; as new section of
Assiminea Fleming 1828 [subgenus of Assiminea, teste Wenz 1939:
633]. Type-species by original designation: Assiminea caledonica
(Crosse) [Hydrocena caledonica Crosse 1869: 24, pl. 2 fig. 4].
Assimineidae. Gastropoda: Prosobranchia.

Cubaedomus Thiele 1928c.
Zoologische Jahrbiicher, 55(5-6): 397, 401; as new subgenus of
Hemisinus Swainson 1840. Type-species by monotypy: Melania
brevis Orbigny [Melania brevis A. Orbigny im Sagra 1842: 10; non
I. Lea 1842]. Melaniidae [Thiaridae, teste Wenz 1939: 719].
Gastropoda: Prosobranchia.

Daedalocheila Thiele 1931.
Handbuch der systematischen Weichtierkunde, 1(2): 580; emenda-
tion pro Daedalochila Beck 1837; as subgenus of Polygyra Say 1818.
Polygyridae. Gastropoda: Pulmonata.

Diaphanella Thiele 1912b.

Deutsche Stidpolar-Expedition 1901-1903, 13(2): 277; as new
group [non Diaphanella Clessin 1880; Diaphanella Thiele replaced
by Notodiaphana Thiele 1917 (g.v.); non Diuaphanella P. Hesse
1916, replaced by Geodiaphana Thiele 1917 (qg.v.)]. Type-species
by monotypy: Bulla fragilis Vélain [1877: 128, pl. 4 fig. 31].
[Notodiaphanidae, teste Thiele 193lc: 383]. Gastropoda: Opis-
thobranchia.

Diochiton Thiele in Troschel & Thiele 1893e.
Das Gebiss der Schnecken, 2(8): 364; as new genus [= Chiton
Linnaeus 1758, s.s., teste Thiele 1909d: 2, A. G. Smith 1960: I65,
and Belle 1983: 123; = section of Chiton s.s., teste Thiele 1929:
20]. Type-species by monotypy: Diochiton albilineatus [sic] Sowerby
[Chiton albolineatus Broderip & G. B. Sowerby I 1829: 368].
Chitonidae. Polyplacophora.

Dioeciostraea Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 814; emenda-
tion pro Dioeciostrea Orton 1928, as synonym of Ostrea (Crassostrea)
Sacco 1897; Stenzel (1971: N1128) listed this as a nomen nullum
in the synonymy of Crassostrea. Ostreidae. Bivalvia.

Doratosepia Thiele 1934.

September 30, 1991 Boss & Bieler 25

Handbuch der systematischen Weichtierkunde, 2(3): 954; emenda-
tion pro Doratosepion Rochebrune 1884; as section of Sepia Linnaeus
1758. Sepiidae. Cephalopoda.

Drupella Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
‘Valdivia’, 17(2): 171 [137]; as new subgenus of Purpura Bruguiere
1789 [subgen. of Drupa Rodding 1798, teste Thiele 1929: 295].
Type-species by designation under plenary powers (ICZN Opinion
1154, 1980: 85): Purpura elata Blainville 1832: 207, pl. 11 fig. 1].
Muricidae. Gastropoda: Prosobranchia.

Durgellina Thiele 1928a.
Zoologische Jahrbiicher, 55(1-3): 135; as new genus. Type-species
by original designation: Durgellina vitrina Thiele 1928 [= Helix
calculosa ‘“‘Couthouy’? Gould 1852, teste Zilch 1959: 281]. Ario-
phantidae [Euconulidae, teste Zilch 1959: 281]. Gastropoda: Pul-
monata.

Eatonina Thiele 1912b.
Deutsche Siidpolar-Expedition 1901-1903, 13(2): 279; as new
subgenus of Eatoniella Dall 1876 [genus, teste Thiele 1929: 167].
Type-species by monotypy: Eatoniella (Eatonina) pusilla Thiele 1912.
[Rissoidae, teste Thiele 1925d: 80; Cingulopsidae, teste Ponder
1985: 100]. Gastropoda: Prosobranchia.

Eatoniopsis Thiele 1912b.
Deutsche Sidpolar-Expedition 1901-1903, 13(2): 237; as new
genus [= Skenella Pfeffer 1886, teste Ponder 1985: 100]. Type-
species by monotypy: Eatoniella paludinoides E. Smith [E. A. Smith
1902: 205, pl. 24 fig. 18]. [Rissoidae, teste Thiele 1925d: 80;
Cingulopsidae, teste Ponder 1985: 100]. Gastropoda: Prosobran-
chia.

Ebalina Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 236; as new
section of Eulimella (Ebala) Leach 1847 [subgenus of Ebala, teste
Wenz 1940: 866]. Type-species by monotypy: Eulimella (Ebalina)
monolirata (Folin) [Aclis monolirata de Folin in Fischer, de Folin
& Périer 1872 [1873]: 178, pl. 7 fig. 6]. Pyramidellidae. Gastro-
poda: Prosobranchia.

*‘Electromactra’’ [1934].
Handbuch der systematischen Weichtierkunde, 2(3): 901; error pro
Electomactra Iredale 1930, as section of Mactra (Mactrella) Gray

26 NEMOURIA No. 39

1853. Mactridae. Bivalvia.

Emmericiopsis Thiele 1928c.
Zoologische Jahrbiicher, 55(5-6): 357, 380; as new section of
Bythina (Gabbia) Tryon 1865 [subgenus of Bulimus Scopoli 1777,
teste Wenz 1939: 590]. Type-species by monotypy: Pachydrobia
lacustris Martens [Pachydrobia lacustris Martens 1897: 26, pl. 9 figs.
12-15]. Hydrobiidae [Bulimidae, teste Wenz 1939: 590; = Bithyntidae].
Gastropoda: Prosobranchia.

Eosipho Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 307; as new
subgenus of Szpho Bruguiére 1792 [genus, teste Wenz 1941: 1154].
Type-species by monotypy: Sipho (Eosipho) smithi (Schepman)
[Chrysodomus (Sipho) smithi Schepman 1911: 300, pl. 19 fig. 6, pl.
23 fig. 11]. Buccinidae. Gastropoda: Prosobranchia.

Eumetula Thiele 1912b.

Deutsche Siidpolar-Expedition 1901-1903, 13(2): 202, 205; as new
subgenus of Eumeta Morch 1868 [non Walker 1855] [synonym (p.
122 [88]) or probable synonym (p. 125 [91]) of Metaxia Montero-
sato 1884, teste Thiele 1925b; genus, teste Thiele 1929: 217].
‘Type-species by monotypy: Eumeta (Eumetula) dilecta Thiele 1912.
Cerithiopsidae [Cerithiidae, teste Powell 1960: 140]. Gastropoda:
Prosobranchia.

Euparthenia Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 233; nomen
novum pro Parthenia Lowe 1840 [part.], non Robineau-Desvoidy
1830; [subgenus of Kleinella A. Adams 1860, teste Wenz 1940:
850]. Pyramidellidae. Gastropoda: Prosobranchia.

Euthymella Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 219; nomen
novum pro Euthymia Jousseaume 1884, non Stal 1876 [subgenus
of Triphora Blainville 1828, teste Wenz 1940: 786]. Triphoridae.
Gastropoda: Prosobranchia.

Ficulina Thiele 1912b.
Deutsche Stidpolar-Expedition 1901-1903, 13(2): 271; as new
subgenus of Cominella Gray 1850 or as new genus [non Ficulina
Gray 1867; Ficulina Thiele replaced by Notoficula Thiele 1917,
(q.v.)|]. Type-species by monotypy: Chlanidota bouveti Thiele 1912.
[Buccinidae, teste Thiele 1929: 315]. Gastropoda: Prosobranchia.

September 30, 1991 Boss & Bieler 27

Fossarella Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
‘Valdivia’, 17(2): 96 [62]; as new genus. Type-species by original
designation: Fossarella pacifica Thiele 1925. Fossaridae. Gastropoda:
Prosobranchia.

Frovina Thiele 1912b.
Deutsche Stidpolar-Expedition 1901-1903, 13(2): 196; as new
genus. Type-species by monotypy: Frovina soror Thiele 1912.
[Naticidae, teste Thiele 1929: 259]. Gastropoda: Prosobranchia.

Fusinella Thiele 1917a.
Nachrichtsblatt der Deutschen Malakozologischen Gesellschaft,
49(1): 24; nomen novum pro Buccinella Thiele 1912 (g.v.), non
Perry 1811. Buccinidae. Gastropoda: Prosobranchia.

Fusiturris Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 361; as new
section of Zurris (Turris) O. F. Miller 1766 [sic] [subgenus of
Turris Réding 1798, teste Wenz 1943: 1404]. Type-species by
monotypy: Turns (Fusiturris) undatiruga (Bivona) [Pleurotoma unda-
tiruga Bivona in Philippi 1844: 171, pl. 26 fig. 13]. Conidae
[Turridae, teste Wenz 1943: 1404]. Gastropoda: Prosobranchia.

Garrettina Thiele 193l1c.
Handbuch der systematischen Weichtierkunde, 1(2): 572; nomen
novum pro Garrettia Cossmann 1900 non (Pease) Paetel 1873
[synonym of Libera Garrett 1881, teste Zilch 1959: 215; see also
Pilsbry 1932: 70]. Endodontidae. Gastropoda: Pulmonata.

Geodiaphana Thiele 1917a.
Nachrichtsblatt der Deutschen Malakozologischen Gesellschaft,
49(1): 23; nomen novum pro Diaphanella P. Hesse 1916, non
Clessin 1880, nec Thiele 1912 (¢.v.); = Hydatina Westerlund 1886,
non Schumacher 1817, nec Ehrenberg 1830 [= Mediterranea Clessin
1880, teste Thiele 1931c: 590; subgenus of Oxychilus Fitzinger
1833, teste Zilch 1959: 254]. [Zonitidae, teste Thiele 1931c: 590].
Gastropoda: Pulmonata.

Georgus Thiele in Troschel & Thiele 1893e.
Das Gebiss der Schnecken, 2(8): 366; as new genus [= Chiton
(Clathropleura), teste Thiele 1909d: 2, 5; = Chiton Linnaeus 1758,
s.s., teste A. G. Smith 1960: 165, Belle 1983: 123]. Type-species
by subsequent designation (Kobelt 1894: 26): Chiton rusticus
[Deshayes 1863: 39, pl. 4 figs. 1-3; = mauritianus Quoy & Gaimard

28 NEMOURIA No. 39

1835, teste Belle 1983: 123]; Thiele (1909d: 2, 5) subsequently
re-identified the material previously called Chiton rusticus as Chiton
(Clathropleura) angusticostatus Quoy & Gaimard 1835, resulting in
a misidentified type species in the sense of ICZN 1985, Art. 70].
Chitonidae. Polyplacophora.

Goniaeolis Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 450; emenda-
tion for Goniégolis M. Sars 1859 [Thiele (1931c: 450) also changed
Goniéolis M. Sars 1859 to Gonteolis}|. Goniaeolididae. Gastropoda:
Opisthobranchia.

Gracilancilla Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
*“Valdivia’’, 17(2): 188 [154]; as new subgenus of Ancilla Lamarck
1799. Type-species by original designation: Ancilla sumatrana Thiele
1925. Olividae. Gastropoda: Prosobranchia.

Gracilimurex Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 289; as new
subgenus of Murex Linnaeus 1758 [genus, teste Wenz 1941: 1087].
Type-species by monotypy: Murex (Gracilimurex) bicolor ‘Thiele
1929. Muricidae. Gastropoda: Prosobranchia.

Haminella Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
“Valdivia”, 17(2): 267 [233]; as new subgenus of Haminea Gray
1847 [subgenus of Haminaea [sic] Turton & Kingston 1830, teste
Zilch 1959: 42]. Type-species by original designation: Hamuinea
maltzani Thiele 1925. Atyidae [Hamineidae [sic, Haminoeidae],
teste Vaught 1989: 65]. Gastropoda: Opisthobranchia.

Harpovoluta Thiele 1912b.
Deutsche Stidpolar-Expedition 1901-1903, 13(2): 213; as new
genus. Type-species by original designation: Harpovoluta vanhéffeni
[vanhoeffeni] Thiele 1912. Volutidae. Gastropoda: Prosobranchia.

Heathia Thiele 1913e.
Das Tierreich, 38: ix, 17; as new genus. Type-species by monotypy:
Ichthyomenia porosa Heath 1911 [p. 48, figs.]. Lepidomeniidae
[Dondersiidae, teste Boss 1982: 949]. Aplacophora.

Hedylopsis Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 443; as new
genus. Type-species by monotypy: Hedylopsis spiculifera (Ko-

September 30, 1991 Boss & Bieler 29

walevsky) [Hedyle spiculifera Kowalevsky 1901b: 21-25, pl. 4 figs.
49-51, pl. 5 figs. 56-69]. Hedylidae [Hedylopsidae, teste Pruvot-Fol
1954: 165]. Gastropoda: Opisthobranchia.

Helioradsia Thiele in Troschel & Thiele 1893e.

Das Gebiss der Schnecken, 2(8): 385; as new genus [= Chaetopleura
s.s. Shuttleworth 1853, teste Belle 1983: 85; = Chaetopleura
(Palliochiton) Dall 1879, teste Kaas & Belle 1987: 103]. Type-
species by monotypy: Helioradsia gemma Carpenter [Chaetopleura
gemma ‘‘Carpenter’’ Dall 1879: 78, fig. 9]. Chitonidae [Chaeto-
pleuridae, teste Belle 1983: 85; Ischnochitonidae, teste Kaas &
Belle 1987: 103]. Polyplacophora.

Hemiglyptopsis Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 624; as new
subgenus of Antinous Robson 1914. Type-species by monotypy:
Antinous (Hemiglyptopsis) foullioyi (Guillou) [Helix foullioyi Le Guil-
lou 1845: 187]. Ariophantidae [Helicarionidae, teste Zilch 1959:
298]. Gastropoda: Pulmonata.

Hemimetis Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 915; as new
subgenus of Apolymetis Salisbury 1929 [subgenus of Tellina Linnaeus
1758, teste Keen in Moore 1969: N615]. Type-species by mono-
typy: Apolymetis (Hemimetis) plicata (Valenciennes): [Tellina plicata
Valenciennes in Lamarck 1827: 154; see also Lamarck 1798: pl.
287 fig. 3]. Tellinidae. Bivalvia.

HMimatina Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 453; nomen
novum pro Himatella Bergh 1891, non Zittel 1879 [= Coryphella
Gray 1850, teste Odhner im Franc 1968: 881]. Flabellinidae
[Coryphellidae, teste Odhner in Franc 1968: 881]. Gastropoda:
Opisthobranchia.

Hochstetterina Thiele 1934.

Handbuch der systematischen Weichtierkunde, 2(3): 796; [listed
by Thiele as “‘nom. nov.” pro Hochstetteria Vélain 1877 part.;
subgenus of Philobrya Carpenter 1872, teste Keen in Moore 1969:
N269]. Type-species by monotypy: Hochstetterina crenella (Vélain)
[Hochstetteria crenella Vélain, 1877: 131-132, pl. 5 figs. 5, 6].
Limopsidae [Philobryidae, teste Keen in Moore 1969: N270].
Bivalvia.

Hydrococcus Thiele 1928c.

30 NEMOURIA No. 39

Zoologische Jahrbticher, 55(5-6): 375, 380; as new genus. Type-
species by monotypy: Hydrococcus graniformis Thiele 1928, new
name for Paludina granum Menke 1843, non Say 1822 [= Rissoa
(Setia) braziert Tenison-Woods 1876 = Hydrococcus brazieri (Tenison-
Woods 1876), teste Ponder 1982: 65]. Hydrobiidae [Hydrococ-
cidae, teste Wenz, 1939: 588]. Gastropoda: Prosobranchia.

Hymenoteuthis Thiele 1916a.
Zoologischer Anzeiger, 48(1): 4; as mew genus or subgen. of
Vampyroteuthis Chun 1903 [genus, teste Thiele 1934: 984; subjec-
tive synonym of Vampyroteuthis, teste Nesis 1987: 276]. Type-
species by monotypy: Cirroteuthis macrope Berry 1910 [1911: 589}.
Vampyroteuthidae. Cephalopoda.

Icoplax Thiele in Troschel & Thiele 1893e.

Das Gebiss der Schnecken, 2(8): 392; as new genus [subgenus of
Callochiton Gray 1847, teste Thiele 1929: 9; = Callochiton Gray
1847, s.s., teste A. G. Smith 1960: I58, Belle 1983: 90].
Type-species by monotypy: Icoplax punicea Couthouy [Chiton puniceus
(Couthouy MS) Gould 1846: 143; = Trachydermon steinenu Pfeiffer
(sic, err. pro Pfeffer) 1886, teste Belle 1983: 90]. Chitonidae
[Callochitonidae, teste A. G. Smith 1960: 158; Ischnochitonidae,
teste Kaas & Belle 1985b: 11]. Polyplacophora.

Iduliella Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 449; nomen
novum pro Dotilla Bergh 1879, non Stimpson 1858. Iduliidae
[Dotonidae, teste Odhner im Franc 1968: 877]. Gastropoda:
Opisthobranchia.

Indopyrgus Thiele 1928c.
Zoologische Jahrbiicher, 55(5-6): 373, 378; as new subgenus of
Pomatopyrgus Stimpson 1865. Type-species by monotypy: Poma-
topyrgus (Indopyrgus) nevilli Thiele 1928. Hydrobiidae. Gastropoda:
Prosobranchia.

Iphitella Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
‘Valdivia’, 17(2): 93 [59]; nomen novum pro Iphitus Jeffreys 1883,
non Rafinesque 1815. Fossaridae [? Littorinidae, teste Thiele 1929:
127]. Gastropoda: Prosobranchia.

Ischnocerithium Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 212; as new
subgenus of Colina H. & A. Adams 1854. Type-species by

September 30, 1991 Boss & Bieler 31

monotypy: Colina (Ischnocerithium) rostrata (Sowerby) [Cerithium
rostratum G. B. Sowerby II 1855: 861, pl. 180, fig. 104].
Cerithiidae. Gastropoda: Prosobranchia.

Jaronia Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 818; emenda-
tion pro Jaronia Pallary 1924 [German spelling of “‘I‘‘]; as section
of Caelatura Conrad 1853. Unionidae. Bivalvia.

Jeffreysiella Thiele 1912b.
Deutsche Stidpolar-Expedition 1901-1903, 13(2): 239; as new
genus [subgen. of Rissoella, teste Thiele 1925b: 91 [57]]. Type-
species by monotypy: feffreysiella notabilis Thiele 1912. [Rissoellidae,
teste Thiele 1925b: 91 [57]]. Gastropoda: Prosobranchia.

Jeffreysilla Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
“Valdivia”, 17(2): 92 [58]; n. subgroup of Russoella Gray 1847
[subgenus of Rissoella, teste Thiele 1929: 179]. Type-species by
monotypy: Rissoella (feffreysilla) zebra Thiele 1925. Rissoellidae.
Gastropoda: Prosobranchia.

Jeffreysina Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
*“Valdivia’’, 17(2): 92 [58]; as new subgenus of Rissoella Gray 1847.
Type-species by subsequent designation (Wenz 1939: 649): Rissoella
Ceffreysina) globularis (Jeffreys) [feffreysia globularis Jeffreys MS.
Forbes & Hanley 1853: 268-269, pl. 133 fig. 5; teste Warén 1980:
24]. Rissoellidae. Gastropoda: Prosobranchia.

Jeffreysiopsis Thiele 1912b.
Deutsche Siidpolar-Expedition 1901-1903, 13(2): 276; as new
subgenus of Feffreysia Alder 1849 [Jeffreysiopsis Thiele = Rissoella
Gray 1847, s.s., teste Thiele 1925b: 91 [57]]. Type-species by
monotypy: Paludestrina duperrei Vélain [1877: 115, pl. 3 figs. 18,
19]. [Rissoellidae, teste Thiele 1925b: 91 [57]]. Gastropoda:
Prosobranchia.

Jheringella Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 841; emenda-
tion pro Iheringella Pilsbry 1893 [German spelling of ‘‘I‘‘]; as genus.
Mutelidae. Bivalvia.

Lepidochiton Thiele 1929.
Handbuch der systernatischen Weichtierkunde, 1(1): 8; emendation

32 NEMOURIA No. 39

pro Lepidochitona Gray 1821, non Lepidochiton Carpenter 1857; as
genus [= Lepitochitona, teste A. G. Smith 1960: 156]. Lepidochi-
tonidae [Ischnochitonidae, teste A. G. Smith 1960: 156]. Polyplaco-
phora.

Lepidopleuroides Thiele 1928b.
Fauna Arctica, 5(2): 564; as new subgenus of Ischnochiton Gray
1847 [= Ischnochiton (Stenosemus) Middendorff 1847, teste A. G.
Smith 1960: I55, Belle 1983: 69; Kaas & Belle 1990: 60].
‘Type-species by original designation: Ischnochiton (Lepidopleuroides)
albus (Linnaeus) [Chiton albus Linnaeus 1767: 1107]; see Kaas &
Belle 1980: 6, for synonyms. Ischnochitonidae. Polyplacophora.

Leptopleura Thiele in Troschel & Thiele 1893e.

Das Gebiss der Schnecken, 2(8): 381; as new genus [= Ischnochiton
Gray 1847, teste Thiele 1909d: 3; A. G. Smith 1960: I55; Belle
1983: 66; Kaas & Belle 1990: 78]. Type-species by monotypy:
Leptopleura catenulata Sowerby [Chiton catenulatus G. B. Sowerby
1832: 104]; material subsequently re-identified as Ischnochiton
fraternus Thiele 1909, by Thiele 19091: 77. Chitonidae [Ischnochi-
tonidae, teste A. G. Smith 1960: 155; Kaas & Belle 1990: 78].
Polyplacophora.

Leucostele Thiele 1927a.

Zoologische Jahrbticher, 53(1-3): 123, 126, 129; as new section of
Paludinella L. Pfeiffer 1841 [introduced as as new section, but
Thiele referred to Leucostele Semper on pp. 126, 129; listed by
Thiele 1929: 170 as “‘Leucostele (Semper) Thiele 1927"; subgenus
of Paludinella, teste Wenz, 1939: 633]. Type-species by original
designation: Paludinella (Leucostele) vitrea Thiele 1927. Assimineidae.
Gastropoda: Prosobranchia.

Limopsilla Thiele 1923.
Zoologischer Anzeiger, 55(11/13): 289; as new genus [= Limopsis
Sassi 1827, teste Newell 72 Moore 1969: N265]. Type-species by
monotypy: Limopsis pumilio E. Smith [E. A. Smith, 1904a: 43, pl.
3 figs. 27, 28]. Limopsinae [Limopsidae, teste Thiele 1926b: 186].
Bivalvia.

Lionucula Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 786; emenda-
tion pro Letonucula W. Quenstedt 1930; as subgenus of Nucula
Lamarck 1799. Nuculidae. Bivalvia.

Lissarcula Thiele 1923.

September 30, 1991 Boss & Bieler 33

Zoologischer Anzeiger, 55(11/13): 290; as new genus [?= Limopsis
Sassi, 1827, teste Newell 72 Moore 1969: N265]. Type-species by
original designation: Lissarcula australis ‘Thiele 1923. Limopsinae
[Limopsidae, teste Thiele 1926b: 186]. Bivalvia.

Lissoconchus Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 125; as new
subgenus of Cremnoconchus Blanford 1869. Type-species by mono-
typy: Cremnoconchus (Lissoconchus) conicus Blanford [Cremnoconchus
conicus W. T. Blanford 1870: 10, pl. 3 fig. 3]. Littorinidae.
Gastropoda: Prosobranchia.

Lithoglyphopsis Thiele 1928c.
Zoologische Jahrbticher, 55(5-6): 366, 379; as new genus. Type-
species by original designation: Lithoglypha modestus Gredler
[Lithoglyphus modestus Gredler 1887: 17]. Hydrobiidae. Gastropoda:
Prosobranchia.

Lophopleura Thiele 1912b.
Deutsche Stidpolar-Expedition 1901-1903, 13(2): 279; as new
subgenus of Lobiger Krohn 1847 [genus, teste Thiele 1925b: 281
[247]]. Type-species by monotypy: Lobiger (Lophopleura) capensts
Thiele 1912. [Philinidae, teste Thiele 1925b: 281 [247]; Oxynoidae,
teste Thiele 1931c: 412]. Gastropoda: Opisthobranchia.

Lophosepia Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 954; emenda-
tion of Lophosepion Rochebrune 1884; as section of Sepia Linnaeus
1758. Sepiidae. Cephalopoda.

Lophyriscus Thiele in Troschel & Thiele 1893e.

Das Gebiss der Schnecken, 2(8): 377; as new genus [= section of
Ischnochiton Gray 1847, teste Thiele 1909d: 3, 7; = Ischnochiton s:.s.
teste Thiele 1929: 17; Kaas & Belle 1990: 78]. Type-species by
subsequent designation (Kobelt 1894: 26): Chiton textilis [Gray
1828: 5, pl. 6 fig. 20]; material subsequently re-identified as C.
oniscus Krauss 1848, by Thiele 1909d: 3, 6; Belle 1983: 66]. See
Kaas & Belle (1980: 93, 131) for additional homonyms and
synonyms. Chitonidae [Ischnochitonidae, teste A. G. Smith 1960:
155; Kaas & Belle 1990: 78]. Polyplacophora.

Lophyropsis Thiele in Troschel & Thiele 1893e.
Das Gebiss der Schnecken, 2(8): 387; as new genus [= Lepidopleurus
Risso 1826, teste Thiele 1909d: 4; = Lepidopleurus (Leptochiton)
Gray 1847, teste Thiele 1929: 6; = subgenus of Ischnochiton Gray

34 NEMOURIA No. 39

1847, teste A. G. Smith 1960: 156; = Leptochiton Gray 1847, s.s.,
teste Belle 1983: 52]. Type-species by monotypy: Lophyropsis
imitatnx [sic] E. Smith [emendation pro Chiton (Ischnochiton)
imitator E. A. Smith 1881]; Thiele (1909d: 4, 7) subsequently
re-identified the material previously called Lophyropsis imitatnix as
Lepidopleurus medinae Plate 1899, resulting in a misidentified type
species in the sense of ICZN 1985, Art. 70]. See Kaas & Belle
(1980: 62) for homonyms of Chiton imitator. Chitonidae [Ischnochi-
tonidae, teste A. G. Smith 1960: 156; Leptochitonidae, teste Kaas
& Belle 1985a: 36]. Polyplacophora.

Luteacarnea Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 827; nomen
novum pro Striata Frierson 1927, non O. Boettger 1878; as
subgroup of Quadrula Rafinesque 1820. Unionidae. Bivalvia.

Lyocyclus Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
*“Valdivia’’, 17(2): 116 [82]; as new group [genus, teste Wenz 1940:
896]. Type-species by original designation: Lyocyclus solutus Thiele
1925. Lyocyclidae Thiele 1925 [Trichotropidae, teste Thiele 1929:
242]. Gastropoda: Prosobranchia.

~ “Tyonstela’’ [1912b].

Deutsche Stidpolar-Expedition 1901-1903, 13(5): 232; error pro
Lyonsiella Sars 1872. [Verticordiidae, teste Thiele 1934: 944].
Bivalvia.

Macrassiminea Thiele 1927a.
Zoologische Jahrbticher, 53(1-3): 122, 126; as new section of
Assiminea Fleming 1828 [subgenus of Assiminea, teste Wenz 1939:
632]. Type-species by original designation: Assiminea francesi (Gray)
[Turbo francesu Gray 1828: suppl. p. 18, pl. 6 fig. 28]. Assimineidae.
Gastropoda: Prosobranchia.

Macrodontopsts Thiele 1931c. |
Handbuch der systematischen Weichtierkunde, 1(2): 662; as new
section of Macrodontes Swainson 1840 [subgenus of Odontostomus
Beck 1837, teste Zilch 1960: 509]. Type-species by monotypy:
Macrodontes simplex Thiele 1906. Bulimulidae [Odondostomidae,
teste Zilch 1960: 509]. Gastropoda: Pulmonata.

Macrogonaxis Thiele 1932a.
Archiv fiir Molluskenkunde, 64(1): 11; as new section of Tayloria
(Pseudogonaxis) Thiele 1932 [subgenus of Tayloria Bourguignat

September 30, 1991 Boss & Bieler 35

1889, teste Zilch 1960: 558]. Type-species by original designation:
enneoides (Martens) [Streptaxis enneoides Martens 1878: 295, pl. 2
figs. 5, 6]. [Streptaxidae, teste Zilch 1960: 558]. Gastropoda:
Pulmonata.

Macroniscus Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 316; as new
subgenus of Macron H. & A. Adams 1853. Type-species by
monotypy: Macron (Macroniscus) lividus A. Adams [Pseudoliva livida
A. Adams 1855: 136]. Buccinidae. Gastropoda: Prosobranchia.

Margarella Thiele in Troschel & Thiele 1893e.
Das Gebiss der Schnecken, 2(8): 406; nomen novum pro Mar-
garitella Thiele 1891 (q.v.), non Meek & Hayden 1860, nec
Schmidt 1880 [subgenus of Margarites Gray 1847, teste Thiele
1912b: 189]. See also notes concerning Margarella/Margaritella in
Thiele 1906a: 13, 1912b: 188, and 1924a: 67. Trochidae. Gastro-
poda: Prosobranchia.

Margaritella Thiele in Troschel & Thiele 1891c.
Das Gebiss der Schnecken, 2(7): 259; as new genus, non Mar-
garitella Meek & Hayden 1860, nec Schmidt 1880 (Porifera)
[Margaritella Thiele replaced by Margarella Thiele 1893, (q.v.)].
Type-species by subsequent designation (Thiele 1924a: 67, for
replacement name Margarella): Margarita expansa Sowerby I 1838.
Trochidae. Gastropoda: Prosobranchia.

Margaritopsis Thiele 1906a.

Nachrichtsblatt der Deutschen Malakozologischen Gesellschaft,
38(1): 15; as new genus [section of Margarites Gray 1847, teste
Thiele 1924a: 50, 67; subgenus of Margarites, teste Wenz 1938:
270]. Type-species by subsequent designation (Thiele 1924a: 50,
67): Marganta frielei Krause [Margarita frielei Krause 1885: 263; =
M. argentata gigantea Leche 1878, teste Thiele 1924a: 50]. [Trochi-
dae, teste Thiele 1924a: 67]. Gastropoda: Prosobranchia.

Mecynoplax Thiele in Troschel & Thiele 1893e.
Das Gebiss der Schnecken, 2(8): 393; as new genus [= Acanthochites
(Notoplax) H. Adams 1861, teste Thiele 1909d: 4, 7; Belle 1983:
144; = Cryptoconchus (Notoplax) H. Adams 1861, s.s., teste Thiele
1929: 14; = Craspedochiton Shuttleworth 1853, teste A. G. Smith
1960: 169]. Type-species by monotypy: Mecynoplax acutirostrata
Reeve [Chiton acutirostratus Reeve 1847b: 25]. Thiele (1909d: 4,
40) subsequently re-identified the material previously called Mecyno-

36 NEMOURIA No. 39

plax acutirostrata as Acanthochites (Notoplax) hilgendorfi Thiele 1909,
resulting in a misidentified type species in the sense of ICZN 1985,
Art. 70]. Chitonidae [Acanthochitonidae, teste A. G. Smith 1960:
169]. Polyplacophora.

Mericella Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 352; as new
section of Cancellaria (Narona) H. & A. Adams 1854 [subgenus of
Narona, teste Wenz 1943: 1364]. Type-species by monotypy:
Cancellaria (Mericella) jucunda Thiele [Cancellaria (Merica) jucunda
Thiele, 1925]. Cancellariidae. Gastropoda: Prosobranchia.

Mesaliopsis Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 181; as new
section of Mesalia Gray 1842 [subgenus of Mesalia, teste Wenz
1939: 651]. Type-species by monotypy: Mesalia (Mesaliopsis) opalina
(Adams & Reeve) [Turritella opalina A. Adams & Reeve 1850: 48,
pl. 12 fig. 7]. Turritellidae. Gastropoda: Prosobranchia.

Metachaetoderma Thiele 1913e.
Das Tierreich, 38: ix, 12; as new genus. Type-species by monotypy:
Chaetoderma challengeri Nierstrasz 1903 [p. 359, pl. 35 figs. 1-26].
Chaetodermatidae [Limifossoridae, teste Boss 1982: 948]. Aplaco-
phora.

Metactaeon Thiele 193l1c.
Handbuch der systematischen Weichtierkunde, 1(2): 378; as new
subgenus of Actaeon Montfort 1810 or as new genus. Type-species
by monotypy: Actaeon (Metactaeon) aequatorialis Thiele 1925 [Ac-
taeon aequatorialis Thiele 1925]. Actaeonidae. Gastropoda: Opis-
thobranchia.

Metamenia Thiele 1913d.
Deutsche Stidpolar-Expedition 1901-1903, 14(1): 52; as new genus.
Type-species by monotypy: Metamenia intermedia Thiele 1913.
Proneomentidae [Pararhopaliidae, teste Boss 1982: 951]. Aplaco-
phora.

Metanachis Thiele 1924b.
Archiv ftir Molluskenkunde, 56(5): 207, 210; as new section of
Columbella Lamarck 1799 [subgenus of Pterygia Roding 1798, teste
Wenz 1941: 1150]. Type-species by original designation: Columbella
jaspidea Sowerby [G. B. Sowerby II 1844: 132, pl. 39 fig. 125].
Columbellidae. Gastropoda: Prosobranchia.

September 30, 1991 Boss & Bieler 37

Metassiminea Thiele 1927a.
Zoologische Jahrbticher, 53(1-3), 122, 126; as new section of
Assiminea Fleming 1828 [subgenus of Asszminea, teste Wenz 1939:
633]. Type-species by original designation: Assiminea philippinica
O. Boettger [1887: 195]. Assimineidae. Gastropoda: Prosobranchia.

Meteuthria Thiele 1912b.
Deutsche Siidpolar-Expedition 1901-1903, 13(2): 243; as new
genus [subgenus of Thalassoplanes Dall 1908, teste Thiele 1929:
317]. Type-species by original designation: Euthria (Glypteuthria)
martenst Strebel [1905: 630, pl. 21 fig. 13]. Buccinidae. Gastro-
poda: Prosobranchia.

Microcardium Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 878; as new
genus [subgenus of Nemocardium Meek 1876, teste Keen 7m Moore
1969: N589]. Type-species by subsequent designation (Keen 1937:
155, with emended spelling peramabile): Cardium (Fulvia) perama-
bilis Dall [1881: 132]. Cardiidae. Bivalvia.

Microdiscula Thiele 1912b.
Deutsche Siidpolar-Expedition 1901-1903, 13(2): 199; as new
genus. Type-species by monotypy: Microdiscula vanhéffeni [vanhoef-
feni] Thiele 1912. [Vitrinellidae, teste Thiele 1925d: 79; Skeneopsi-
dae, teste Thiele 1929: 176]. Gastropoda: Prosobranchia.

Microhappia Thiele 1927c.
Abhandlungen herausgegeben von der Senckenbergischen Natur-
forschenden Gesellschaft, 40(3): 320; as new genus [section of
Muradiscops H. B. Baker 1925, teste Thiele 1931c: 598; subgenus
of Miradiscops, teste Zilch 1960: 545]. Type-species by monotypy:
Microhappia brasiliensis Thiele 1927. Systrophiidae. Gastropoda:
Pulmonata.

Micropleurotoma Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 362; as new
subgenus of Turris O. F. Miller 1766 [sic] [genus, teste Wenz
1943: 1412]. Type-species by monotypy: Turris (Micropleurotoma)
sptrotropoides (Thiele) [Pleurotoma spirotropoides Thiele 1925].
Conidae [Turridae, teste Wenz 1943: 1412]. Gastropoda: Proso-
branchia.

Mirellia Thiele 1933c.
Sitzungsberichte der Gesellschaft Naturforschender Freunde zu
Berlin, 1933(4-7): 283; as new section of Ptychotrema L. Pfeiffer

38 NEMOURIA No. 39

1853 [subgenus of Ptychotrema, teste Zilch 1960: 577]. Type-species
by monotypy: Ennea prodigiosa E. Smith [E. A. Smith 1903: 316,
pl. 4 fig. 11]. Streptaxidae. Gastropoda: Pulmonata.

Mé6rchiella [Moerchiella] Thiele 1924a.

Mitteilungen aus dem Zoologischen Museum in Berlin, 11(1): 71;
nomen novum pro Morchia A. Adams 1860 (non Albers 1850)
[non Mérchiella G. Nevill 1884; Morchiella Thiele replaced by
Morchinella ‘Vhiele 1931 (q.v.)]; also introduced as new name by
Thiele 1925b: 59 [25], and cited by Thiele 1929: 175 as of that
date. See also Tomlin 1936: 136. Trochidae [Adeorbidae, teste
Thiele 1929: 175]. Gastropoda: Prosobranchia.

Mé6rchinella [Moerchinella] Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 763; nomen
novum pro Morchiella Thiele 1924 (g.v.), non G. Nevill 1884.
Adeorbidae [Tornidae, teste Wenz 1939: 645]. Gastropoda: Proso-
branchia.

Monoeciostraea Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 814; emenda-
tion pro Monoeciostrea Orton 1928, as synonym of Ostrea Linnaeus
1758, s.s.; Stenzel (1971: N1138) listed this as a nomen nullum in
the synonymy of Ostrea. Ostreidae. Bivalvia.

Mopaliella Thiele 1909d.

Zoologica, 22(56/1): 16; as new genus [= Cyanoplax Pilsbry 1892,
teste A. G. Smith 1960: 158; subgenus of Lepidochitona Gray 1821,
teste Kaas & Belle 1980: [ix]; = Lepidochitona s.s., teste Belle 1983:
96-97]. Type-species by monotypy: Chiton bipunctatus Sowerby [G.
B. Sowerby I 1832: 104]. [Callochitonidae, teste Thiele 19091:
116; Lepidochitonidae, teste Thiele 1925d: 36; Ischnochitonidae,
teste A. G. Smith 1960: 158, Belle 1983: 96]. Polyplacophora.

Mopaliopsis Thiele in Troschel & Thiele 1893c.

Das Gebiss der Schnecken, 2(8): 393; as new genus [= Cyanoplax
Pilsbry 1892, teste Thiele 1909d: 4, 7, 16; = Lepidochitona Gray
1821, s.s., teste Belle 1983: 96]. Type-species by monotypy:
Mopalhiopsis cingillata Reeve [Chiton cingillatus Reeve 1847a: pl. 23
fig. 160]; Thiele (1909d: 4, 7) subsequently re-identified the
material previously called Mopaliopsis cingillata as Cyanoplax hartwe-
git (Carpenter 1856), resulting in a misidentified type species in
the sense of ICZN 1985, Art. 70]. Chitonidae [Ischnochitonidae,
teste A. G. Smith 1960: 158, Belle, 1983: 96]. Polyplacophora.

September 30, 1991 Boss & Bieler 39

Nantivitrea Thiele 1927a.
Zoologische Jahrbiicher, 53(1-3): 123, 126; as new section of
Paludinella Pfeiffer 1841 [subgenus of Paludinella, teste Wenz 1939:
633]. Type-species by original designation: Paludinella helicoides
Gundlach [Gundlach in Poey 1865: 70]. Assimineidae. Gastropoda:
Prosobranchia.

Nanula Thiele 1924a.
Mitteilungen aus dem Zoologischen Museum in Berlin, 11(1): 54,
69; as new genus. Type-species by monotypy: Gibbula tasmanica
Petterd [1879: 103]. Trochidae. Gastropoda: Prosobranchia.

Naytiopsis Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 324; as new
subgenus of Nassa Lamarck 1799 [genus, teste Wenz 1943: 1232].
Type-species by monotypy: Nassa (Naytiopsis) granum Lamarck
[Buccinum grana Lamarck 1822: 274]. Nassidae [Nassariidae, teste
Wenz 1943: 1232]. Gastropoda: Prosobranchia.

Neactaeonina Thiele 1912b.
Deutsche Stdpolar-Expedition 1901-1903, 13(2): 219; as new
genus or subgen. of Actaeonina Orbigny 1850. Type-species by
monotypy: Actaeonina cingulata Strebel [1908: 8, pl. 2, fig. 17a-c].
[Actaeonidae, teste Thiele 1926a: 105]. Gastropoda: Opisthobran-
chia.

Nesodiscus Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 571; as new
genus. Type-species by monotypy: Nesodiscus fabrefactus (Pease)
[Helix fabrefacta Pease 1865: 669]. Endodontidae. Gastropoda:
Pulmonata.

Noetiella Thiele & Jaeckel 1931b.

Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
*“Valdivia’’, 21: 176 (18); as new section of Arca [section of Arca
(Noetia) (Gray) H. & A. Adams 1857, teste Thiele 1934: 793;
synonym of Noetia Gray 1857, teste Newell in Moore 1969: N261].
Type-species by monotypy: Arca (Noetiella) congoensis Thiele &
Jaeckel, 1931. Subsequently attributed by various authors to Jaeckel
& Thiele, Thiele & Jaeckel or to Thiele alone (e.g., Thiele 1934:
793). Arcidae [Noetiidae, teste Newell in Moore 1969: N260].
Bivalvia.

Notochiton Thiele 1906d.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .

40 NEMOURIA No. 39

‘Valdivia’, 9(2): 332; as new genus [subgenus of Nuttallochiton
Plate, 1899, teste Thiele 1929: 9; = Nuttallochiton, teste A. G.
Smith 1960: 160; Belle, 1983: 88; Kaas & Belle: 116]. Type-species
by monotypy: Notochiton mirandus Thiele, 1906. [Lepidochitonidae,
teste Thiele, 1925c: 36; Callistoplacidae, teste A. G. Smith 1960:
160; Ischnochitonidae, teste Kaas & Belle 1987: 116]. Polyplaco-
phora.

Notodiaphana Thiele 1917a.
Nachrichtsblatt der Deutschen Malakozologischen Gesellschaft,
49(1): 23; nomen novum pro Diaphanella Thiele 1912 (g.v.), non
Clessin 1880, nec Hesse 1916. [Notodiaphanidae, teste Thiele
1931c: 383]. Gastropoda: Opisthobranchia.

Notodiscus Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 575; as new
genus. Type-species by monotypy: Notodiscus hooker1 (Reeve) [Helix
hookeri Reeve 1854: pl. 208, no. 1474]. Endodontidae. Gastropoda:
Pulmonata.

Notoficula Thiele 1917a.
Nachrichtsblatt der Deutschen Malakozologischen Gesellschaft,
49(1): 24; nomen novum pro Ficulina Thiele 1912, non Gray 1867
[section of Chlanidota Martens 1878, teste Thiele 1929: 315;
subgenus of Chlanidota, teste Wenz 1943: 1201]. [Buccinidae, teste
Thiele 1925b: 179]. Gastropoda: Prosobranchia.

Notomenia Thiele 1897b.
Zoologischer Anzeiger, 20(542): 398; as new genus. Type-species
by original designation: Notomenia clavigera Thiele 1897. [Lepido-
meniidae, teste Thiele 1913e: ix, 22; family position uncertain, teste
Boss 1982: 949]. Aplacophora.

Novovaginula Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 491; as new
subgenus of Pseudoveronicella Germain 1908. Type-species by
monotypy: Pseudoveronicella (Novovaginula) carinata (Thiele) [Veron-
icella (Vaginula) carinata Thiele 1927]. Vaginulidae. Gastropoda:
Pulmonata.

Ochrodermatina Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 552; as new
section of Ochroderma Ancey 1885 [subgenus of Ochroderma, teste
Zilch 1959: 350]. Type-species by monotypy: Ochroderma (Ochroder-
matina) carolinum (Martens) [Stenogyra carolina Martens, 1880:

September 30, 1991 Boss & Bieler 41

147]. Subulinidae. Gastropoda: Pulmonata.

Odostomiopsis Thiele in Martens in Martens & Thiele 1904a.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
‘Valdivia’, 7A: 68 (also, Thiele 72 Martens & Thiele 1904b: 156
[10]); as new genus [= Toledonia Dall 1902, teste Thiele 1931c:
382]. Type-species by indication (ICZN, 1985, Article 68c):
Odostomiopsis typica Thiele in Martens in Martens & Thiele 1904.
[Diaphanidae, teste Thiele 1931c: 382]. Gastropoda: Opisthobran-
chia.

Olivellopsis Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 333; as new
section of Belloliva Peile 1922 [subgenus of Belloliva, teste Wenz
1943: 1272]. Type-species by monotypy: Bellohva (Olivellopsis)
simplex (Pease) [Olivella (Callianax) simplex Pease 1868: 281, pl.
23 fig. 24]. Olividae. Gastropoda: Prosobranchia.

Onithoplax Thiele 1909d.
Zoologica, 22(56/1): 3; as new subgenus of Tonicia Gray 1847
[section of Acanthopleura (Onithochiton) Gray 1847, teste Thiele
19091: 117; section of Jonicia (Lucilina) Dall 1882, teste Thiele
1929: 22; = Lucilina, teste A. G. Smith 1960: 166]. Type-species
by subsequent designation (Belle 1983: 133, who erroneously
credited the designation to Thiele 1929): Chiton mallardi Deshayes
1863 [1863: 38, pl. 5 fig. 14]. [Chitonidae, teste Thiele 19091:
117]. Polyplacophora.

Oopyramis Thiele 1930.
Die Fauna Stidwest-Australiens, 5(8): 580; as new genus, or new
subgenus of Tiberia Monterosato 1875. Type-species by monotypy:
Oopyramis biplicata Thiele 1930. [Pyramidellidae, teste Wenz 1940:
876]. Gastropoda: Prosobranchia.

Opaliopsis Thiele 1928d.
Zeitschrift fiir wissenschaftliche Zoologie, 132: 92; as new subgroup
of Scala (Klein 1753) Bruguiére 1792 [= Epitonium Réding 1798]
[section of Scala (Cirsotrema) Mérch 1852, teste Thiele 1929: 222;
subgenus of Cirsotrema, teste Wenz 1940: 798]. Type-species by
monotypy: Scala elata Thiele 1925. Scalidae [= Epitoniidae].
Gastropoda: Prosobranchia.

Oriella Thiele 1927a.
Zoologische Jahrbticher, 53(1-3): 124, 127; as new subgenus or new
section of Omphalotropis Pfeiffer 1851 [section, teste Thiele 1929:

42 NEMOURIA No. 39

172]. Type-species by original designation: Omphalotropis submari-
tima Quadras & Médllendorff [1894: 33]. Erroneously spelled
““Oviella” by Wenz 1939: 636. Assimineidae. Gastropoda: Proso-
branchia.

Ovassiminea Thiele 1927a.
Zoologische Jahrbticher, 53(1-3): 122, 125; as new section of
Assiminea Fleming 1828 [subgenus of Assiminea, teste Wenz 1939:
632]. Type-species by original designation: Assiminea dohrniana
Nevill [G. Nevill 1880: 162; fig. in Yen 1939: pl. 4 fig. 37].
Assimineidae. Gastropoda: Prosobranchia.

Pachydrobiella Thiele 1928c.
Zoologische Jahrbticher, 55(5-6): 367, 379; as new genus. Type-
species by monotypy: Pachydrobia brevis Bavay [1895: 87, pl. 5 fig.
2]. Hydrobiidae. Gastropoda: Prosobranchia.

Pagodinella Thiele 1917a.
Nachrichtsblatt der Deutschen Malakozologischen Gesellschaft,
49(1): 24; nomen novum pro Pagodula Hesse 1916, non Montero-
sato 1884 [Pagodinella Thiele = Pagodina Stabile 1864, non Beneden
1853; and = Pagodulina Clessin 1876, teste Thiele 1931c: 510].
[Vertiginidae, teste Thiele 1926a: 139; Orculidae, teste Zilch 1959:
156]. Gastropoda: Pulmonata.

Paracerastus Thiele 1933c.
Sitzungsberichte der Gesellschaft Naturforschender Freunde zu
Berlin, 1933(4-7): 308; as new subgenus of Cerastus Albers 1860
[subgenus of Mabilliella Ancey 1886, teste Zilch 1959: 195].
Type-species by monotypy: Ena drymaeoides Thiele [Ena (Rachisel-
lus) drymaeoides Thiele 1911]. Enidae. Gastropoda: Pulmonata.

Parachiton Thiele 1909d.
Zoologica, 22(56/1): 14; as new subgenus of Lepidopleurus (Leach
MS) Risso 1826 [subgenus of Leptochiton Gray 1847, teste Belle
1983: 54]. Type-species by monotypy: Lepidopleurus (Parachiton)
acuminatus Thiele 1909. [Lepidopleuridae, teste Thiele 1909i:
116; Leptochitonidae, teste Belle 1983: 54; Kaas & Belle 1990:
14]. Polyplacophora.

Paranoetia Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 793; as new
section of subgenus Arca (Noetia) (Gray) H. & A. Adams 1857 [=
Sheldonella Maury 1917, teste Newell in Moore 1969: N262].
Type-species by monotypy: Arca (Paranoetia) lateralis Reeve [Arca

September 30, 1991 Boss & Bieler 43

lateralis Reeve 1844: pl. 17 fig. 115]. Arcidae [Noetiidae, teste
Newell 72 Moore 1969: N262]. Bivalvia.

Parasitala Thiele 1931c.
Sitzungsberichte der Gesellschaft Naturforschender Freunde zu
Berlin, 1930: 397, 398; as new genus. Type-species by original
designation: Parasitala osumiensis Thiele 1931. [Ariophantidae, teste
Thiele 1931c: 635; Euconulidae, teste Zilch 1959: 281]. Gastro-
poda: Pulmonata.

Parateuthis Thiele 1920b.
Deutsche Stidpolar-Expedition 1901-1903, 16(8)(4): 463; as new
genus [possible senior synonym of Alluroteuthis N. Odhner 1923,
teste Thiele 1934: 970; genus incertae sedis, teste Nesis 1987:
260]. Type-species by monotypy: Parateuthis tunicata Thiele 1920.
[PAlluroteuthidae, teste Thiele 1934: 970]. Cephalopoda.

Paratilia Thiele 1924b.
Archiv fiir Molluskenkunde, 56(5): 204, 210; as new section of
Pyrene (Mitrella) Risso 1826 [subgenus of Mzitrella, teste Wenz 1941:
1139]. Type-species by monotypy: Columbella alizona Melvill [Co-
lumbella (Mitrella) alizonae Melvill & Standen 1901: 402, pl. 21 fig.
5]. Columbellidae. Gastropoda: Prosobranchia.

Parhedyle Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 443; as new
genus. Type-species by monotypy: Parhedyle tyrtowi (Kowalevsky)
[Hedyle tyrtowu Kowalevsky 1900: 200-202]. Hedylidae [Microhedy-
lidae, teste Pruvot-Fol 1954: 167]. Gastropoda: Opisthobranchia.

Parvoliva Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 335; as new
section of Oliva Bruguiére 1789 [subgenus of Oliva, teste Wenz
1943: 1281]. Type-species by monotypy: Oliva (Parvoliva) dubia
Schepman [Oliva dubia Schepman 1903: 68]. Olividae. Gastropoda:
Prosobranchia.

Paryphantopsis Thiele 1928a.
Zoologische Jahrbiicher, 55(1-3): 125; as new section of Flammulina
Martens 1873 [subgenus, teste Thiele 1931c: 576; genus, teste Zilch
1959: 223]. Type-species by original designation: Flammulina
(Paryphantopsis) lamelligera Thiele 1928. Endodontidae. Gastropoda:
Pulmonata.

Patellidea Thiele in Troschel & Thiele 1891c.

4.4, NEMOURIA No. 39

Das Gebiss der Schnecken, 2(7): 315; as new genus [subgroup of
Patella (Scutellastra) H. & A. Adams 1854, teste Thiele 1929: 40;
subgenus of Patella, teste Wenz 1938: 215]. Type-species by
monotypy: fatella granularis Linnaeus [1758: 782]. Patellidae.
Gastropoda: Prosobranchia.

Patellona Thiele in Troschel & Thiele 1891c.
Das Gebiss der Schnecken, 2(7): 317; as new genus [subgroup of
Patella (Cymbula) H. & A. Adams 1854, teste Thiele 1929: 41].
‘Type-species by subsequent designation (Kobelt 1894: 26): Patella
granatina [Linnaeus 1758: 782]. Patellidae. Gastropoda: Prosobran-
chia.

Patellopsis Thiele in Troschel & Thiele 1891c.
Das Gebiss der Schnecken, 2(7): 324; as new genus, non Patellopsis
Nobre 1886. Type-species: no nominal taxa included; description
based on the radula of an unnamed South African species.
Homonymy recognized by Thiele 1931c: 766; never renamed.
Patellidae. Gastropoda: Prosobranchia.

Patinastra Thiele im Troschel & Thiele 189Ic.
Das Gebiss der Schnecken, 2(7): 325; as new genus [section of
Helcion (Patina) Leach 1852, teste Thiele 1929: 41; subgenus of
Helcion Montfort 1810, teste Wenz 1938: 214]. Type-species by
monotypy: Patella pruinosa Krauss 1848 [p. 56, pl. 3 figs. 9a, b].
Patellidae. Gastropoda: Prosobranchia.

Paucidentella ‘Thiele 1933c.
Sitzungsberichte der Gesellschaft Naturforschender Freunde zu
Berlin, 1933(4-7): 285; as new section of Gulella L. Pfeiffer 1856
[subgenus of Gulella, teste Zilch 1960: 569]. Type-species by
original designation: Gulella conica (Martens) [Ennea conica Martens
1876: 264, pl. 4 figs. 6, 7]. Streptaxidae. Gastropoda: Pulmonata.

Pediculariella Thiele 1925d.
Handbuch der Zoologie, 5(1): 88; as new group [subgenus of
Pedicularia Swainson 1840, teste Wenz 1941: 1001]. Type-species
by monotypy: californica (Newcomb) [Pedicularia californica New-
comb 1864: 121]. Cypraeidae [Amphiperatidae, teste Wenz 1941:
1001; Pediculariidae, teste Vaught 1989: 33]. Gastropoda: Proso-
branchia.

*“Pherusiana’’ [1929].
Handbuch der systematischen Weichtierkunde, 1(1): 226; error pro
Pherusina (Norman MS) Chaster 1898. Aclididae. Gastropoda:

September 30, 1991 Boss & Bieler 45

Prosobranchia.

Phyllomenia Thiele 1913d.
Deutsche Siidpolar-Expedition 1901-1903, 14(1): 38, 45; as new
genus. Type-species by monotypy: Phyllomenia austrina Thiele 1913.
Lepidomeniidae [Phyllomeniidae, teste Boss 1982: 950]. Aplaco-
phora.

Pistris Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 917; nomen
novum pro Pristis Lamy 1918, non Linck 1790 (Latham 1794), nec
(Lepeletier MS) Brullé 1846 [subgenus of Tellana Linnaeus 1758,
teste Keen in Moore 1969: N619]. Tellinidae. Bivalvia.

Plagiolopsis Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 834; new
genus [introduced by Thiele as “nom nov.”’ pro Plagiola sensu
Stimpson; = Plagiola Rafinesque 1819, teste Haas in Moore 1969:
N455; = Ellipsaria Rafinesque 1820, teste Johnson 1978: 248].
Type-species by monotypy: P. securis (Lea) [Unio securis Lea 1829:
437, pl. 11 fig. 17]. Unionidae. Bivalvia.

Pleurobranchella Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition . . .
*‘Valdivia’’, 17(2): 283 [249]; as new genus or new subgenus of
Euselenops Pilsbry 1896 [section of Pleurobranchaea (Euselenops), teste
Thiele 1931c: 419]. Type-species by monotypy: Euselenops (Pleuro-
branchella) nicobarica Thiele 1925. Pleurobranchidae. Gastropoda:
Opisthobranchia.

Plicarcularia Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 324; as new
section of Nassa (Arcularia) Link 1807 [subgenus of Arcularia, teste
Wenz 1943: 1232]. Type-species by monotypy: Nassa (Plicarcularia)
thersites (Bruguiére) [Nassa thersites Lamarck 1816: pl. 394, liste,
p. 1]. Nassidae [Nassariidae, teste Wenz 1943: 1232]. Gastropoda:
Prosobranchia.

Poeciloplax Thiele in Troschel & Thiele 1893e.
Das Gebiss der Schnecken, 2(8): 365; as new genus [section of
Chiton, teste Thiele 1909d: 2; = Chiton (Amaurochiton) Thiele 1893,
teste A. G. Smith 1960: 166; = Chiton Linnaeus 1758, s.s., teste
Belle 1983: 123]. Type-species by monotypy: Poeciloplax glauca
Gray [1828: p. 5], non Quoy & Gaimard 1835 [= viridus Quoy &
Gaimard 1835, non Spengler 1797, = quoyi Deshayes 1836, = var.

46 NEMOURIA No. 39

limosus (Suter MS) Nierstrasz 1905; teste Kaas & Belle 1980: 54}.
Chitonidae. Polyplacophora.

Primovula Thiele 1925d.
Handbuch der Zoologie, 5(1): 88; as new group [genus, teste
Wenz 1941: 1009, 1010]. Type-species by monotypy: becker
(Sowerby) [Amphiperas beckeri G. B. Sowerby III 1900: 5, pl. 1 fig.
13]. Cypraeidae [Amphiperatidae, teste Wenz 1941: 1009, 1010;
Ovulidae, teste Vaught 1989: 33]. Gastropoda: Prosobranchia.

Probuccinum Thiele 1912b.

Deutsche Siidpolar-Expedition 1901-1903, 13(2): 207, 211; as new
genus or new subgenus of Neobuccinum E. Smith 1879 [section of
Glypteuthria Strebel 1905, teste Thiele 1929: 318; subgenus of
Glypteuthria, teste Wenz 1943: 1207]. Type-species by subsequent
designation (Wenz 1943: 1207): Glypteuthria (Probuccinum) tenera
E. A. Smith [Neobuccinum tenerum E. A. Smith 1907: 2, pl. 1, figs.
2, 2a). Buccinidae. Gastropoda: Prosobranchia.

Probythinella Thiele 1928c.
Zoologische Jahrbiicher, 55(5-6): 370, 378; as new subgenus of
Cincinnatia Pilsbry 1891 [later (footnote p. 370 and p. 378) as
section of Hoyia F. C. Baker 1925]; subgenus of Hoyia, teste Wenz
1939: 569]. Type-species by monotypy: Cincinnatia emarginata
(Kuster) [Paludina emarginata “‘Say’”’ Kiister 1852: 50, pl. 10 figs.
3, 4]. Hydrobiidae. Gastropoda: Prosobranchia.

Procalpurnus Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 272; as new
section of Calpurnus Montfort 1810 [subgenus of Calpurnus, teste
Wenz 1941: 1010]. Type-species by monotypy: Calpurnus (Pro-
calpurnus) lacteus (Lamarck) [Ovula lactea Lamarck 1810: 111].
Cypraeidae [Ovulidae, teste Vaught 1989: 33]. Gastropoda: Proso-
branchia.

Prochaetoderma Thiele 1902b.
Zeitschrift fiir wissenschaftliche Zoologie, 72(2-3): 275; as new
genus. Type-species by monotypy: Chaetoderma raduliferum Ko-
walevsky 1901 [Chaetoderma radulifera Kowalevsky 1901a: 264, pl.
10 figs. 1-20]. [Chaetodermatidae, teste Thiele 1913e: ix, 10;
Caudofoveata, Prochaetodermatidae, teste Boss 1982: 948]. Aplaco-
phora.

Prochiton Thiele 1890a.

September 30, 1991 Boss & Bieler 47

Zeitschrift fiir wissenschaftliche Zoologie, 49(3): 428; as new genus
[listed by Belle 1983: 155 as a hypothetical taxon of Plate 1901].
Type-species by monotypy: Chiton rubicundus O. G. Costa 1829
[pl. 1 fig. 3] [= Chiton (Rhyssoplax) corallinus Risso 1826, teste
Kaas & Belle 1980: 31, 112]; Thiele (1902b: 284) subsequently
re-identified the material previously called Chiton rubicundus as
Callochiton doriae (Capellini 1859) [= Callochiton septemvalvis (Mon-
tagu 1803) subsp. euplaeae (O. G. Costa 1829), teste Kaas & Belle
1980: 40], resulting in a misidentified type species in the sense of
ICZN 1985, Art. 70].

Proclava Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 212; as new
subgenus of Cerithium Bruguiére 1789 [subgenus of Clava Martyn
1784, teste Wenz 1940: 761, 1943: 1501]. Type-species by
monotypy: Cerithium (Proclava) pfefferi Dunker [Vertagus pfefferi
Dunker 1877: 75]. Cerithtidae. Gastropoda: Prosobranchia.

Prodoridunculus Thiele 1912b.
Deutsche Stidpolar-Expedition 1901-1903, 13(2): 218, 222; as new
genus. Type-species by monotypy: Prodoridunculus gaussianus Thiele
1912. [Polyceridae, teste Thiele 1931c: 427; Onchidoridae, teste
Powell 1960: 165; Lamellidoridae, teste Odhner im Franc 1968:
861]. Gastropoda: Opisthobranchia.

Profundinassa Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 323; as new
subgenus of Nassa Lamarck 1799 [genus, teste Wenz 1943: 1231].
‘Type-species by monotypy: Nassa (Profundinassa) babylonica Watson
[Nassa (Aciculina) babylonica Watson 1882: 366; 1886: 185, pl. 11
fig. 8]. Nassidae [Nassariidae, teste Wenz 1943: 1231]. Gastropoda:
Prosobranchia.

Prohappia Thiele 1927c.
Abhandlungen herausgegeben von der Senckenbergischen Natur-
forschenden Gesellschaft, 40(3): 313; as new subgenus of Happia
Bourguignat 1889. Type-species by monotypy: Happia besckei
(Dunker) [Helix bescket Dunker in Pfeiffer 1847: 81]. Systrophiidae.
Gastropoda: Pulmonata.

Prolacuna Thiele 1913a.
Zentralblatt fiir Zoologie, 2(3): 86; nomen novum pro Sublacuna
Thiele 1912 (g.v.), non Pilsbry 1895, nec Cossmann 1899
[subgenus of Frovina Thiele 1912, teste Wenz 1941: 1028].

48 NEMOURIA No. 39

[Naticidae, teste Thiele 1929: 260]. Gastropoda: Prosobranchia.

Proneptunea Thiele 1912b.
Deutsche Stidpolar-Expedition 1901-1903, 13(2): 246; as new
genus [subgenus of Thalassoplanes Dall 1908, teste Thiele 1929:
317]. Type-species by monotypy: Proneptunea amabilis Thiele 1912.
Buccinidae. Gastropoda: Prosobranchia.

Proneritula Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 324; as new
section of Nassa (Cyclope) Risso 1826 [subgenus of Cyclope, teste
Wenz 1943: 1234]. Type-species by monotypy: Nassa (Proneritula)
westerlundi (Brusina) [Cyclope westerlundi Brusina 1900: 89]. Nassi-
dae [Nassariidae, teste Vaught, 1989: 49]. Gastropoda: Prosobran-
chia.

Proplanaxis Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 203; as new
subgenus of Planaxis Lamarck 1822. Type-species by monotypy:
Planaxis (Proplanaxis) planicostatus Sowerby [Planaxis planicostatus
G. B. Sowerby I 1825, Appendix p. xiii]. Planaxidae. Gastropoda:
Prosobranchia.

Proreinia Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 531; as new
section of Reimia Kobelt 1876 [subgenus of Reinia, teste Zilch 1959:
391]. Type-species by monotypy: Reimia (Proreinia) eastlakeana
(Mollendorff) [Clausilia eastlakeana MGllendorff 1882: 187; 1883:
262, pl. 8 figs. 7a-c]. Clausiliidae. Gastropoda: Pulmonata.

Proseila Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 218; as new
section of Seda A. Adams 1861 [subgenus of Sezla, teste Wenz
1940: 781]. Type-species by monotypy: Seila (Proseila) capitata
Thiele [Sela capitata Thiele 1925]. Cerithiopsidae. Gastropoda:
Prosobranchia.

Proselenites Thiele 1927c.

Abhandlungen herausgegeben von der Senckenbergischen Natur-
forschenden Gesellschaft, 40(3): 313; as new subgenus of Haplo-
trema Ancey 1881 [= Geomene Pilsbry 1927, teste Zilch 1960:
549]. Type-species by subsequent designation (Baker 1928: 124;
see also Baker 1941: 132, and Pilsbry 1946: 208): Helix concava
Say 1821 [p. 159]. Systrophiidae [Haplotrematidae, teste Zilch
1960: 549]. Gastropoda: Pulmonata.

September 30, 1991 Boss & Bieler 49

Prosipho Thiele 1912b.
Deutsche Siidpolar-Expedition 1901-1903, 13(2): 206; as new
group [genus, teste Thiele 1929: 319]. Type-species by subsequent
designation (Wenz 1943: 1210): Prosipho gaussianus Thiele 1912.
Buccinidae. Gastropoda: Prosobranchia.

Prothalotia Thiele 1930.
Die Fauna Siidwest-Australiens, 5(8): 566; as new section of
Cantharidus Montfort 1810 [subgenus of Thalotia Gray 1847, teste
Wenz 1938: 305]. Type-species by original designation: Cantharidus
(Prothalotia) flindersi (P. Fischer) [Trochus flinderst P. Fischer 1878:
65]. [Trochidae, teste Thiele 1931c: 735]. Gastropoda: Prosobran-
chia.

Protomella Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 182; as new
section of Protoma Baird 1870 [subgenus of Protoma, teste Wenz
1939: 659]. Type-species by monotypy: Protoma (Protomella) pulchra
Sowerby [Protoma pulchra G. B. Sowerby III 1905: 279]. Turritelli-
dae. Gastropoda: Prosobranchia.

Pruvotia Thiele 1894b.
Zeitschrift fiir wissenschaftliche Zoologie, 58(2): 272; as new genus.
‘Type-species by monotypy: Proneomenia sopita Pruvot 1891 [p.
721, pl. 30 fig. 50, pl. 31 fig. 84]. [Proneomeniidae, teste Thiele
1913e: x, 52; Rhopalomeniidae, teste Boss 1982: 952]. Aplaco-
phora.

Pseudactaeon Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
‘Valdivia’, 17(2): 257 [223]; as new group [genus, teste Thiele
1931c: 378; genus Pseudacteon (sic), teste Odhner im Franc 1968:
835]. Type-species by monotypy: Actaeon albus (Sowerby 1873)
[Tornatella albus G. B. Sowerby II 1873: 720, pl. 59 fig. 6].
Actaeonidae. Gastropoda: Opisthobranchia.

Pseudanachis Thiele 1924b.
Archiv ftir Molluskenkunde, 56(5): 208; as new genus [also
described as new by Thiele 1925b: 172 [138]]. Type-species by
monotypy: Columbella duclosiana Sowerby [G. B. Sowerby II 1844:
113, pl. 36 figs. 15, 16; non Columbella duclosiana Orbigny in Sagra
1842: 136, pl. 21 figs. 31-33]. ?Columbellidae. Gastropoda:
Prosobranchia.

Pseudassiminea Thiele 1927a.

50 NEMOURIA No. 39

Zoologische Jahrbticher, 53: 123, 127; as new genus. Type-species
by original designation: Omphalotropis waigiouensis Sykes [1903:
67]. Assimineidae. Gastropoda: Prosobranchia.

Pseudisidora Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 476; as new
subgenus of Lymnaea Lamarck 1799 [genus, teste Zilch 1959: 93].
Type-species by monotypy: Lymnaea (Pseudisidora) rubella Lea
[Lymnea rubella Lea 1844: 12; non Clessin 1873]. Lymnaeidae.
Gastropoda: Pulmonata.

Pseudocleopatra Thiele 1928c.
Zoologische Jahrbticher, 55(5-6): 394, 400; as new genus. Type-
species by monotypy: Pseudocleopatra togoensis ‘Thiele 1928. Melaniidae
[Thiaridae, teste Wenz 1939: 703]. Gastropoda: Prosobranchia.

Pseudocyclotus Thiele 1894a.
Nachrichtsblatt der Deutschen Malakozologischen Gesellschaft,
26(1-2): 23. Type-species by original designation: Cyclostoma no-
vaehiberniae Quoy & Gaimard [Cyclostoma novae hiberniae [sic]
Quoy & Gaimard 1832: 182, pl. 12 figs. 15-19]. Cyclostomidae
[Assimineidae, teste Thiele 1929: 173]. Gastropoda: Prosobranchia.

Pseudogonaxis Thiele 1932a.
Archiv fiir Molluskenkunde, 64(1): 11; as new subgenus of Tayloria
Bourguignat 1881. Type-species by original designation: nsendweenis
[sic] (Putzeys) [Streptaxis nseudweensis Putzeys 1899: lv, fig. 3].
[Streptaxidae, teste Zilch 1960: 558]. Gastropoda: Pulmonata.

Pseudostomatella Thiele 1924a.
Mitteilungen aus dem Zoologischen Museum in Berlin, 11(1): 59,
70; as new section of Stomatia Helbling 1779 [subgenus of Stomatia,
teste Wenz 1938: 322]. Type-species by original designation:
Pseudostomatella papyracea (Chemnitz) [Stomatella papyracea ‘‘Chem-
nitz’”’” A. Adams 1850b: 31]. Trochidae [Stomatiidae, teste Wenz
1938: 322]. Gastropoda: Prosobranchia.

Pseudotopoma Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 128; as new
section of Cyclotopsis Blanford 1864 [subgenus of Cyclotopsis, teste
Wenz 1939: 532]. Type-species by monotypy: Otopoma hinduorum
Blanford [1864: 464]. Pomatiasidae. Gastropoda: Prosobranchia.

Pseudotritonia Thiele 1912b.
Deutsche Siidpolar-Expedition 1901-1903, 13(2): 218, 224; as new

September 30, 1991 Boss & Bieler 51

genus. Type-species by monotypy: Pseudotritonia quadrangularis
Thiele 1912. [Notaeolidiidae, teste Thiele 1926a: 113; Charcotiidae,
teste Odhner im Franc 1968: 880]. Gastropoda: Opisthobranchia.

Psychroteuthis Thiele 1920b.
Deutsche Stidpolar-Expedition 1901-1903, 16(8)(4): 440; as new
genus. Type-species by monotypy: Psychroteuthis glacialis ‘Thiele
1920. Psychroteuthidae. Cephalopoda.

Ptychobela Thiele 1925b.

Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
‘Valdivia’’, 17(2): 215 [181]; as new subgenus of Bela Gray 1847
or as new genus. [= Brachytoma Swainson 1840, teste Thiele 1929:
362]. Type-species by original designation: Pleurotoma crenularis
Lamarck [Clavatula crenularis Lamarck 1816: pl. 440, liste p. 9].
Pleurotomidae [Conidae, teste Thiele 1929: 362; Turridae, teste
Wenz 1941: 1412]. Gastropoda: Prosobranchia.

Ptychocardia Thiele 1912b.
Deutsche Siidpolar-Expedition 1901-1903, 13(2): 232; as new
genus. Type-species by monotypy: Ptychocardia vanhéffen [vanhoef-
feni] Thiele 1912. [Cyamiidae, teste Thiele 1934: 857; Vesi-
comyidae, teste Powell 1960: 177; Cyamiidae, teste Chavan in
Moore 1969: N539]. Bivalvia.

Ptychosyrinx Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
*WValdivia”’, 17(2): 210 [176]; as new group [synonym of Subulata
Martens 1901, teste Wenz 1943: 1402]. Type-species by monotypy:
Drillia bisinuata (Martens) [Pleurotoma (Subulata) bisinuata Martens
1901: 17]. Pleurotomidae [Conidae, teste Thiele 1929: 359;
Turridae, teste Wenz 1943: 1402]. Gastropoda: Prosobranchia.

*“Punigapia’’ [1934].
Handbuch der systematischen Weichtierkunde, 2(3): 911; error pro
Punipagia Iredale 1930; as genus. Semelidae. Bivalvia.

Pustulosa Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 827; nomen
novum pro Bullata Frierson 1927, non Jousseaume 1875; as
subgroup of Quadrula Rafinesque 1820 [synonym of Quadrula
(Amphinaias) Crosse & Fischer 1894, teste Haas in Moore 1969:
N437]. Unionidae. Bivalvia.

Pyrunculus “Thiele.”

52 NEMOURIA No. 39

Indicated by Thiele (1925b: 381 [347]) as being new, but was
introduced by Pilsbry (1895: 229) as new name for Sao H. & A.
Adams 1854, non Billberg 1820. Retusidae. Gastropoda: Opis-
thobranchia.

Radsiella Thiele in Troschel & Thiele 1893e.

Das Gebiss der Schnecken, 2(8): 368; as new genus, non Radsiella
Pilsbry in Tryon & Pilsbry 1892 [Radstella Thiele = Ischnochiton
Gray 1847, teste Thiele 1909d: 2, 6; = Ischnochiton (Haploplax)
Pilsbry 1894, teste A. G. Smith 1960: I56]. Type-species by
subsequent designation (Kobelt 1894: 26): Chiton punctatissimus [sic]
[Chiton punctulatissimus G. B. Sowerby I 1832: 58 [= Chiton pusio
Sowerby 1832, teste Belle 1983: 66]]. Chitonidae [Ischnochi-
tonidae, teste A. G. Smith: 156; Kaas & Belle 1990: 78].
Polyplacophora.

Rhachidina Thiele 1921b.
Archiv fiir Molluskenkunde, 53(3): 150; as new group [genus
Rhachidina Thiele 1911 (sic), teste Zilch 1959: 194]. Type-species
by original designation: R. tumefacta (Reeve) [Bulimus tumefacta
Reeve 1848: Bulimus, no. 374]. Date given as “1911” by Thiele
193lc: 524. [Enidae, teste Thiele 1926a: 139]. Gastropoda:
Pulmonata.

Rhinocochlis Thiele 193 1c.
Handbuch der systematischen Weichtierkunde, 1(2): 633; as new
genus. Type-species by monotypy: Rhinocochlis nasuta (Metcalfe)
[Helix nasuta Metcalfe 1851: 70]. Ariophantidae. Gastropoda:
Pulmonata.

Rhodoplax Thiele in Troschel & Thiele 1893e.

Das Gebiss der Schnecken, 2(8): 384; as new genus [section of
Ischnochiton (Stenoplax) (Carpenter MS) Dall 1878, teste Thiele
1909i: 112, 116; = Ischnochiton Gray 1847, teste A. G. Smith 1960:
155; Belle 1983: 66; Kaas & Belle 1990: 78]. Type-species by
subsequent designation (Kobelt 1894: 26): Chiton squamulosus [C.
B. Adams 1845: 8; = Chiton striolatus Gray 1828, teste Thiele
1909d: 3, Belle 1983: 66]. Chitonidae [Ischnochitonidae, teste
Thiele 19091: 116; Kaas & Belle 1990: 78]. Polyplacophora.

Rhombosepia Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 954; emenda-
tion pro Rhombosepion Rochebrune 1884; as section of Sepia
Linnaeus 1758. Sepiidae. Cephalopoda.

September 30, 1991 Boss & Bieler 53

Rhopalopleura Thiele in Troschel & Thiele 1893e.

Das Gebiss der Schnecken, 2(8): 373; as new genus [= Acan-
thopleura (Mesotomura) Pilsbry 1893, teste Thiele 1909d: 3, 6; =
Acanthopleura Guilding 1830, teste A. G. Smith 1960: 167, Belle
1983: 127]. Type-species by monotypy: Rhopalopleura aculeata
Linnaeus [Chiton aculeatus Linnaeus 1758: 667; = Chiton echinatus
Barnes 1824, teste Belle 1983: 127; for synonyms of echinatus, see
Kaas & Belle 1980: 41]. Chitonidae. Polyplacophora.

Rhyssoplax Thiele in Troschel & Thiele 1893.

Das Gebiss der Schnecken, 2(8): 368; as new genus. Type-species
by designation under the plenary powers (ICZN 1971, Opinion
951): Chiton affinis Issel 1869 [:234] [= Chiton janeirensis sensu
Thiele, non Gray 1828]. Calloplax Thiele 1909 (g.v.) was estab-
lished on the real Chiton janeirensis Gray 1828. Chitonidae.
Polyplacophora.

Roseniella Thiele 193I1c.

Handbuch der systematischen Weichtierkunde, 1(2): 536; nomen
novum pro Thalestris Lindholm 1913, non Claus 1863, nec Gistl
1868, = Rosenia P. Hesse 1916 non Waagen & Wentzel 1886, nec
Schepman 1914, nec Kieffer 1921, nec Schawerda 1922 [subgenus
of Acrotoma O. Boettger 1881, teste Zilch 1960: 441]. Clausiliidae.
Gastropoda: Pulmonata.

Rostellariella Thiele 1929.

Handbuch der systematischen Weichtierkunde, 1(1): 253; as new
section of Rostellaria Lamarck 1799 [subgenus of Tibia Réding
1798, teste Wenz 1940: 934]. Type-species by monotypy: Rostellaria
(Rostellariella) delicatula Nevill [Rostellaria delicatula G. Nevill 1881b:
262]. Strombidae. Gastropoda: Prosobranchia.

Rupacilla Thiele 1927a.

Zoologische Jahrbiicher, 53(1-3): 123, 126; as new section of
Paludinella Pfeiffer 1841 [subgenus of Paludinella, teste Wenz 1939:
633]. Type-species by original designation: ‘‘Omphalotropis”’ filocincta
Quadras & MGllendorff [Omphalotropis filocincta Quadras & Mdllen-
dorff 1896: 14]. Assimineidae. Gastropoda: Prosobranchia.

Sandalomenia Thiele 1913d.

Deutsche Stidpolar-Expedition 1901-1903, 14(6)(1): 41, 38; as new
genus. Type-species by subsequent designation (Salvini-Plawen
1978: 48): Sandalomenia papilliigera Thiele 1913. Lepidomeniidae
[Sandalomeniidae, teste Boss 1982: 949]. Aplacophora.

54 NEMOURIA No. 39

Scansicochlea Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 656; emenda-
tion pro Scansicohlea Pilsbry 1931; as subgenus of Bulimulus Leach
1815. Bulimulidae. Gastropoda: Pulmonata.

Sculptassiminea Thiele 1927a.
Zoologische Jahrbticher, 53(1-3): 126; as new section of Assiminea
(Leach) Fleming 1828 [subgenus of Assiminea, teste Wenz 1939:
632]. Type-species by original designation: Asstminea microsculpta
Nevill [G. Nevill 1880: 165, 1881b: 158, pl. 7 fig. 5]. Assimineidae.
Gastropoda: Prosobranchia.

Segmentella Thiele 1925b.

Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
‘Valdivia’, 17(2): 110 [76]; as new subgenus of Vermetus Daudin
1800. Type-species by subsequent designation (Wenz 1939: 676):
Stephopoma (Segmentella) agulhasensis (Thiele) [Vermetus (Segmen-
tella) agulhasensis ‘Thiele 1925]. Vermetidae [“‘should probably be
classed as annelids,’’ teste Keen, 1961: 184]. Gastropoda: Proso-
branchia.

Semiretusa Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
‘Valdivia’, 17(2): 268 [234]; as new group [subgenus of Retusa
T. Brown 1827, teste Thiele 1931c: 390]. Type-species by
subsequent designation (Zilch 1959: 46): Retusa (Semiretusa) born-
eensis (A. Adams) [Bulla (Utriculus) borneensis A. Adams 1850a:
572, pl. 120 fig. 23]. Retusidae. Gastropoda: Opisthobranchia.

Silicula “Jeffreys”? Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 790; as genus;
see comments above on Beanella. See Introduction, p. 5. Ledidae.
Bivalvia.

Sinomytilus Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 801; as new
section of Mytilus (Chloromya) Mérch 1853. Type-species by
monotypy: Mytilus (Sinomytilus) crosseanus (Morlet) [Dreissena crosse-
ana L. Morlet 1884: 402, pl. 13 figs. 5-5c]. Mytilidae [Dreissenidae,
teste Brandt 1974: 307]. Bivalvia.

Sitalina Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 635; as new |
genus. Type-species by monotypy: Sitalina circumcincta (Reinhardt) —
[Conulus circumcinctus Reinhardt 1883: 85]. Ariophantidae [Euconu- _

September 30, 1991 Boss & Bieler 55

lidae, teste Zilch 1959: 281]. Gastropoda: Pulmonata.

Sitalinopsis Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 612; as new
subgenus of Kaliella Blanford 1863 [genus, teste Zilch 1959: 296].
Type-species by monotypy: Kaliella (Sitalinopsis) conulus (Blanford)
[Nanina (Kaliella?) conula W.T. Blanford 1865: 73]. Ariophantidae
[Helicarionidae, teste Zilch 1959: 296]. Gastropoda: Pulmonata.

*<Slerochiton’’ [19093].
Zoologica, 22(56/2): 95; error pro Sclerochiton (Carpenter MS)
Dall 1882, non Kraatz 1859. [Chitonidae, teste A. G. Smith 1960:
166]. Polyplacophora.

Spathidosepia Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 954; emenda-
tion pro Spathidosepion Rochebrune 1884; as subgroup of Sepia
Linnaeus 1758. Sepiidae. Cephalopoda.

Sphaerocylichna Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
“Valdivia”, 17(2): 276 [242]; as new subgroup of Cylichna Lovén
1846 [section of Cylichna s.s., teste Thiele 1931c: 391]. Type-
species by subsequent designation (Zilch 1959: 25): Cylichna
(Sphaerocylichna) atyoides Thiele [Cylichna atyoides Thiele 1925].
Scaphandridae. Gastropoda: Opisthobranchia.

Spiraxilla Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 773; nomen
novum pro “Spiraxis Pilsbry 1903” [as listed by Thiele 1931c:
675], non Spiraxis C. B. Adams 1850; this was found by Thiele
(1935: 1154) to have been an unnecessary replacement: “‘Spiraxis’’
was an error pro Gyraxis Pilsbry 1903. See also Tomlin 1936:
136. Urocoptidae. Gastropoda: Pulmonata.

Stereoplax Thiele in Troschel & Thiele 1893e.

Das Gebiss der Schnecken, 2(8): 383; as new genus [= Ischnochiton
Gray 1847, teste A. G. Smith 1960: 155; = Ischnochiton (Ischnoplax)
(Carpenter MS) Dall 1879, teste Belle 1983: 71]. Type-species by
monotypy: Stereoplax multicosiata [sic] C. B. Adams [Chiton multi-
costatus C. B. Adams 1845: 8 = Chiton pectinatus G. B. Sowerby
I 1840, teste Belle 1983: 71]. Chitonidae [Ischnochitonidae, teste
A. G. Smith 1960: I55]. Polyplacophora.

Strioconus Thiele 1929.

56 NEMOURIA No. 39

Handbuch der systematischen Weichtierkunde, 1(1): 374; as new
section of Conus Linnaeus 1758 [= Dendroconus Swainson 1840,
teste Thiele 1931c: 742]. Type-species by monotypy: Conus (Strio-
conus) striatus Linnaeus [Conus stratus Linnaeus 1758: 716].
Conidae. Gastropoda: Prosobranchia.

Striokaliella Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 613; as new
section of Kaliella Blanford 1863 [subgenus of Kaliella, teste Zilch
1959: 296]. Type-species by original designation: “‘Sitala’’ crenicincta
Godwin-Austen [Sitala crenicincta Godwin-Austin 1883: 75, pl. 13
fig. 2]. Ariophantidae. Gastropoda: Pulmonata.

Striosubulina Thiele 1933c.
Sitzungsberichte der Gesellschaft Naturforschender Freunde zu
Berlin, 1933(4-7): 301; as new section of Homorus (Neoglossula)
Pilsbry 1909 [subgenus of Homorus, teste Zilch 1959: 343].
Type-species by monotypy: striatella Rang [Helix striatella Rang
1831: 38, pl. 3 fig. 7], non Anthony 1840]. Subulinidae. Gastro-
poda: Pulmonata.

Striotellina Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 917; as new
section of Quadrans Bertin 1878 [objective junior synonym of
Serratina Pallary 1922, teste Keen in Moore 1969: N620]. Type-
species by monotypy: Quadrans (Stnotellina) serratus (Renieri)
[Tellina serrata Brocchi 1814: 510]. Tellinidae. Bivalvia.

Stylopyramis Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 236; as new
genus. Type-species by monotypy: Stylopyramis cerithioides Thiele
1929. Pyramidellidae. Gastropoda: Prosobranchia.

Sublacuna Thiele 1912b.
Deutsche Stidpolar-Expedition 1901-1903, 13(2): 195; as new
genus [non Sublacuna Pilsbry 1895; Sublacuna Thiele replaced by
Prolacuna Thiele 1913, (q.v.)]. Type-species by monotypy: Subla-
cuna indecora Thiele 1912. [Naticidae, teste Thiele 1929: 260].
Gastropoda: Prosobranchia.

Subninella Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 68; as new
section of Zurbo Linnaeus 1758 [subgenus of Turbo, teste Wenz
1938: 351]. Type-species by monotypy: Turbo (Subninella) undulatus
Martyn [Turbo undulatus Gmelin 1791: 3597]. Turbinidae. Gastro-

September 30, 1991 Boss & Bieler 57

poda: Prosobranchia.

Supplanaxis ‘Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 203; as new
subgenus and section of Planaxis Lamarck 1822. Type-species by
monotypy: Planaxis (Supplanaxis) nucleus (Lamarck) [Buccinum
nucleus Bruguiére 1789: 254]. Planaxidae. Gastropoda: Prosobran-
chia.

Suterilla Thiele 1927a.
Zoologische Jahrbticher, 53(1-3): 124, 127; as new section of
Omphalotropis Pfeiffer 1851 [subgenus of Omphalotropis, teste Wenz
1939: 636]. Type-species by monotypy: “‘Cirsonella’’ neozelanica
Murdoch [Cirsonella neozelanica Murdoch 1899: 320, pl. 16 figs.
2-6]. Assimineidae. Gastropoda: Prosobranchia.

Sypharochiton Thiele in Troschel & Thiele 1893e.

Das Gebiss der Schnecken, 2(9): 365; as new genus [section of
Chiton, teste Thiele 1909d: 2; subgenus of Chiton, teste A. G. Smith
1960: 166; = Chiton Linnaeus 1758, s.s., teste Belle 1983: 123].
Type-species by monotypy: Sypharochiton pellisserpentis [sic] Quoy
& Gaimard [Chiton pelliserpentis Quoy & Gaimard 1835: 381, pl.
74 figs. 17-22]. For synonyms of pelliserpentis, see Kaas & Belle
1980: 97. Chitonidae. Polyplacophora.

Tellinangulus Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 919; as new
subgenus of Angulus Megerle von Miuhlfeld 1811 [subgenus of
Telina Linnaeus 1758, teste Keen im Moore 1969: N620]. Type-
species by monotypy: Angulus (Tellinangulus) aethiopicus (Jackel &
Thiele [sic]) [Yellina (Tellinella) aethiopica Thiele & Jaeckel 1931).
Tellinidae. Bivalvia.

Thaumastochiton Thiele 1909d.

Zoologica, 22(56/1): 34; as new subgenus of Craspedochiton Shuttle-
worth 1853 [= Craspedochiton, teste A. G. Smith 1960: 169; =
Notoplax (Spongiochiton), teste Belle 1983: 145]. Type-species by
monotypy: Craspedochiton (Thaumastochiton) mébiusi [moebiusi] Thiele
1909 [= Onithochiton isipingoensis Sykes 1901, teste Kaas & Belle
1980: 86]. [Cryptoplacidae, teste Thiele, 1909i: 116; Acanthochi-
tonidae, teste A. G. Smith 1960: 169]. Polyplacophora.

**Toniciella’’ [1893e].
Das Gebiss der Schnecken, 2(8): 389; error pro Jonicella Carpenter
1873. Chitonidae [Ischnochitonidae, teste A. G. Smith 1960: 158].

58 NEMOURIA No. 39

Polyplacophora.

Tonicina Thiele 1906e.

Expédition Antactique Francaise . . . Sciences naturelles: docu-
ments scientifiques, p. [i]; as new genus [subgenus of Nuttalina
(Carpenter MS) Dall 1871, teste Belle 1983: 101]. Type-species
by monotypy: Chiton zscham Pfeffer 1886 [Pfeffer 1x Martens &
Pfeffer 1886: 105, pl. 3 fig. 2]. Also described as new by Thiele
1906f: 549. [Ischnochitonidae, teste Thiele 1908c: 20]. Polyplaco-
phora.

Toniciopsis Thiele in Troschel & Thiele 1893e.
Das Gebiss der Schnecken, 2(8): 371; as new genus [= Tonicia
(Lucilina) Dall 1882; teste Thiele 1909d: 3, 6]. Type-species by
subsequent designation (Kobelt 1894: 26): Chiton pictus Reeve 1847
[1847a: pl. 15 fig. 29], non Blainville 1825, nec Bean in Thorpe
1844, [= Chiton lamellosus Quoy & Gaimard 1835, teste Belle 1983:
132]. Chitonidae. Polyplacophora.

Triboplax Thiele in Troschel & Thiele 1893e.

Das Gebiss der Schnecken, 2(8): 366; as new genus [section of
Chiton, teste Thiele 1909d: 2; = Sypharochiton Thiele 1893, teste
Thiele 1929: 20; = Chiton Linnaeus 1758, s.s., teste Belle 1983:
123]. Type-species by monotypy: TJriboplax scabricula Sowerby
[Chiton scabriculus G.B. Sowerby I 1832]; Thiele (1909d: 2)
subsequently re-identified the material previously called Triboplax
scabricula as Chiton pelliserpentis Quoy & Gaimard 1835, resulting
in a misidentified type species in the sense of ICZN 1985, Art.
70]. Chitonidae. Polyplacophora.

Trochaclis Thiele 1912b.
Deutsche Stidpolar-Expedition 1901-1903, 13(2): 192; as new
genus. Type-species by monotypy: Jrochaclis antarctica Thiele 1912.
?Aclidae, teste Thiele 1925d: 85; >?Trochaclididae, teste Thiele
1929: 179]. Gastropoda: Prosobranchia.

Turbacmella Thiele 1927a.
Zoologische Jahrbticher, 53(1-3): 123, 126; as new genus. Type-
species by original designation: ‘‘Adelomorpha’’ doherty: Fulton
[Adelomorpha dohertyi Fulton 1899: 219, pl. 11 fig. 19]. As-
simineidae. Gastropoda: Prosobranchia.

Typhlosyrinx Thiele 1925b.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
“Valdivia”, 17(2): 218 [184]; as new subgroup of Pleurotomella

September 30, 1991 Boss & Bieler 59

Verrill 1873 [genus, teste Thiele 1929: 365]. Type-species by
monotypy: Pleurotomella (Typhlosyrinx) vepallida Martens [Pleurotoma
(Leucosyrinx) vepallida Martens 1903: 204, 1904: pl. 2 fig. 6).
Pleurotomidae [Conidae, teste Thiele 1929: 365; Turridae, teste
Wenz 1943: 1425]. Gastropoda: Prosobranchia.

Umbotrochus (Monterosato MS) Thiele 1924a.
Mitteilungen aus dem Zoologischen Museum in Berlin, 11(1): 58,
69; as genus, based on “Umbotrochus Monterosato in coll.’? [non
Umbotrochus Perner 1904; Umbotrochus Thiele replaced by Callum-
bonella Thiele 1924 (g.v.)]. Type-species by monotypy: Gzbbula
gorgonarum P. Fischer [1883: 393]. Trochidae. Gastropoda: Proso-
branchia.

Utriculastra Thiele 1925b.

Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition. . .
“Valdivia”, 17(2): 269 [235]; as new subgenus of Retusa T. Brown
1827 [synonym of Acteocina Gray 1847, teste Mikkelsen & Mikkelsen
1984: 168]. Type-species by original designation: Retusa canaliculata
(Say) [Volvaria canaliculata Say 1826: 211]. Retusidae [not a
member of Retusidae, teste Mikkelsen & Mikkelsen 1984: 164-165].
Gastropoda: Opisthobranchia.

Venustaconcha Thiele 1934.
Handbuch der systematischen Weichtierkunde, 2(3): 837; nomen
novum pro Venusta Frierson 1927, non O. Boettger 1877, nec
Barrande 1881; as subgroup of Lampsilis Rafinesque 1820 [subgenus
of Lampsilis, teste Haas in Moore 1969: N450]. Unionidae. Bivalvia.

WalkRerilla Thiele 1928c.
Zoologische Jahrbticher, 55(5-6): 372, 379; as new section of
Birgella (Somatogyrus) Gill 1863 [subgenus of Somatogyrus, teste
Wenz 1939: 576]. Type-species by monotypy: Somatogyrus coosaensis
Walker [Somatogyrus coosaensis Walker 1915: 40]. Hydrobiidae.
Gastropoda: Prosobranchia.

Watsonella Thiele 1912b.
Deutsche Siidpolar-Expedition 1901-1903, 13(2): 237; as new
group [non Watsonella W. Grabau 1900; Watsonella Thiele replaced
by Boogina Thiele 1913, (q.v.)]. Type-species by monotypy: Rissoa
(Setia) sinapi Watson [1886: 610, pl. 46 fig. 13]. [Rissoidae, teste
Thiele 1929: 167; Cingulopsidae, teste Ponder 1985: 100]. Gastro-
poda: Prosobranchia.

*‘Zeacumanthus’’ [1929].

60 NEMOURIA No. 39

Handbuch der systematischen Weichtierkunde, 1(1): 208; error pro
Zeacumantus Finlay 1927. Potamididae. Gastropoda: Prosobranchia.

Zebrinops Thiele 1931c.
Handbuch der systematischen Weichtierkunde, 1(2): 525; as new
genus. Type-species by monotypy: Zebrinops revoil1 (Bourguignat)
[Limicolaria revoili Bourguignat im Revoil 1882: 42, pl. 2 figs. 24,
25 (26)]. Enidae. Gastropoda: Pulmonata.

Zemiropsis Thiele 1929.
Handbuch der systematischen Weichtierkunde, 1(1): 332; as new
genus [= Babylonia Schliiter 1838, teste Barnard 1959: 60, 147].
Type-species by monotypy: Zemiropsis papillaris (Sowerby) [Eburna
papillaris G. B. Sowerby I 1825: Appendix, p. xxii]. ? Olividae.
Gastropoda: Prosobranchia.

September 30, 1991 Boss & Bieler 61

LITERATURE CITED
(Authorities for type-species, subsequent designations and placement)

Adams, A. 1850a. Monograph of the family Bullidae. [In:] Sowerby, George
Brettingham, Junior [II]. Thesaurus Conchyliorum or figures and descrip-
tions of Recent shells. Volume II. Part XI. Containing monographs of the
genera Navicella and Bulla, with a supplementary plate of Cyclostoma.
London, Sowerby, pp. 553-608, pls. 119-125.

— 1850b. An arrangement of Stomatellidae, including the characters of a new
genus, and of several new species. Proceedings of the Zoological Society
of London, 1850: 19-40.

— 1853. Descriptions of new shells, from the Cumingian Collection; with a
note on the genus Nematura. Proceedings of the Zoological Society of
London, CCXXXIII, Part XIX, 1851: 224-226.

— 1855. Descriptions of thirty-nine new species of shells, from the collection
of Hugh Cuming, Esq. Proceedings of the Zoological Society of London,
Part XXII, 1854: 130-138, pl. 28.

Adams, A. & L. Reeve. 1850. [In:] Adams, A. (ed.). The Zoology of the Voyage
of H.M.S. Samarang; under the command of Captain Sir Edward Belcher,
C. B., F.R.A.S., F.G.S., during the years 1843-1846. Mollusca. Part III,
pp. 45-87, pls. 18-24.

Adams, C. B. 1845. Specierum novarum conchyliorum, in Jamaica repertorum,
synopsis. Pars 1. Proceedings of the Boston Society of Natural History, 2:
1-10.

Baker, H. B. 1928. Thiele’s Brazilian land snails. The Nautilus, 41: 124-129.

— 1941. Some Haplotrematidae. The Nautilus, 54: 130-136, text-figs. 1-12.

Barnard, K. H. 1959. Contributions to the knowledge of South African marine
Mollusca. Part II. Gastropoda: Prosobranchiata: Rachiglossa. Annals of the
South African Museum, 45(1): 1-237, 51 figs.

Bavay, A. 1895. Coquilles nouvelles, provenant des récoltes de M. L. Levay,
dans les rapides du Haut-Mékong, pendant la campagne du Massie,
1893-1894-1895. Journal de Conchyliologie, 43: 82-94, pls. 5, 6.

Belle, R. A. van. 1983. The systematic classification of the chitons (Mollusca:
Polyplacophora). Informations de la Société Belge de Malacologie, (11)1-2:
1-178, 13 pls.

Bergh, L. S. R. 1891. Die cladohepatischen Nudibranchien. Zoologische
Jahrbticher, Abteilung fiir Systematik, 1890, 5: 1-75.

Berry, S. S. 1911. Preliminary notices of some new Pacific cephalopods.
Proceedings of the United States National Museum, 40(1838): 589-592.

Bieler, R. & K. J. Boss. 1989. Johannes Thiele and his contributions to Zoology.
Part 1. Biography and bibliography. Nemouria, 34: 1-30.

62 NEMOURIA No. 39

Blainville, H. M. D. de. 1832. Disposition méthodique des espéces récentes et
fossiles des genres pourpre, ricinule, licorne et concholépas de M. Lamarck,
et description des espéces nouvelles ou peu connues, faisant partie de la
collection du Muséum d’Histoire Naturelle de Paris. Nouvelles Annales
du Muséum d’Histoire Naturelle, Paris, 1(2): 189-263, pls. 9-12.

Blanford, W. T. 1864. On the classification of the Cyclostomacea of eastern
Asia. Annals and Magazine of Natural History, (3)13: 441-465.

— 1865. Contributions to Indian Malacology. Descriptions of some land-shells
from Arakan, Pegu, and Ava, with notes on distribution of described
species. Journal of the Asiatic Society of Bengal, 34(2): 66-105.

— 1870. Contributions to Indian Malacology. No. XI. Descriptions of new
species of Paludomus, Cremnoconchus, Cyclostoma, and of Helicidae from
various parts of India. Journal of the Asiatic Society of Bengal, 39(2): 9-25,
pl. 3.

Boettger, O. 1887. Aufzahlung der zur Gattung Assiminea Fleming gehérigen
Arten. Jahrbuch der Deutschen Malakozoologischen Gesellschaft, 14:
147-234, pl. 6, figs. 6-11.

Boss, K. J. 1971. Critical estimate of the number of Recent Mollusca. Occasional
Papers on Mollusks, Department of Mollusks, Museum of Comparative
Zoology, Harvard University, Cambridge, Massachusetts, 3(40): 81-135.

— 1982. Mollusca [and] Classification of Mollusca. [In:] Parker, S. P. Synopsis
and Classification of Living Organisms. McGraw-Hill, New York, 1:
945-1166; 2: 1092-1096.

— 1989. Preface. Occasional Papers on Mollusks, Department of Mollusks,
Museum of Comparative Zoology, Harvard University, Cambridge, Massa-
chusetts, 4: xi-xvi.

Boss, K. J., J. Rosewater & F. A. Ruhoff. 1968. The zoological taxa of William
Healey Dall. United States National Museum, Bulletin 287: 427 pp.

Boss, K. J. & L. Urbanski. (in press). Oblomovism revisited. Journal of Molluscan
Behaviour.

Bourguignat, J. R. 1882. Mollusques terrestres et fluviatiles. [In:] G. Reévoil.
Faune et Flore des Pays comalis (Afrique orientale). Challamel, Ainé,
Paris, 108 pp., 4 pls.

Brandt, R. A. M. 1974. The non-marine aquatic Mollusca of Thailand. Archiv
fiir Molluskenkunde, 105: 1-423, 30 pls.

Brocchi, G. B. 1814. Conchiologia fossile Subapennina con osservazioni geolo-
giche sugli Apennini e sul suolo adiacente. Volume 1: 1-240, Volume 2:
241-712, pls. 1-16; Reale, Milano.

Broderip, W. J. & G. B. Sowerby I. 1829. Observations on new or interesting
Mollusca contained, for the most part, in the Museum of the Zoological
Society. The Zoological Journal, 4: 359-379, pls. 9.

Bruguiére, J.-G. 1789. Encyclopédie Méthodique. Histoire Naturelle des Vers,

September 30, 1991 Boss & Bieler 63

1(1): 1-344; Panckoucke, Paris.

— 1792. Encyclopédie Méthodique. Histoire Naturelle des Vers, 1(2): 345-
757; Panckoucke, Paris.

Brusina, S. 1900. Zur Molluskenfauna des Mittelmeeres. Nachrichtsblatt der
Deutschen Malakozoologischen Gesellschaft, 32: 86-89.

Carpenter, P. P. 1856. Descriptions of (supposed) new species and varieties of
shells, from the Californian and West Mexican coasts, principally in the
collection of Hugh Cuming, Esq. Proceedings of the Zoological Society of
London, Part 23, 1855: 228-235.

Chavan, A. 1969. [In:] Moore, R. C. (ed.). Treatise on Invertebrate Paleontology.
The Geological Society of America, Inc. and the University of Kansas,
Part N, Mollusca 6, Bivalvia, Volume 2 (of 3): N537-N539.

Cossmann, A. E. M. 1921. Essais de Paléoconchologie comparée. Paris [privately
published]. Douziéme livraison: 348 pp., 6 pls. + leaflets A-D.

Costa, O. G. 1829. Catalogo sistematico e ragionato dei Testacei delle sue Sicilie.
Tipografia della Minerva, Naples, 1-8 + i-cxxxii pp., 2 pls.

Crosse, H. 1862. Description d’une espéce nouvelle appartenant au genre Pisania.
Journal de Conchyliologie, 10: 251-252, pl. X, fig. 5 [sic, 7].

— 1869. Description d’espéces inédites provenant de la Nouvelle-Calédonie.
Journal de Conchyliologie, 17: 24-28, pls. 1-2

Dall, W. H. 1879. Report on the limpets and chitons of the Alaskan and Arctic
regions, with descriptions of genera and species believed to be new.
Scientific Results of the Exploration of Alaska, Article 4, p. 63-126.

— 1881. Reports on the results of dredging, under the supervision of Alexander
Agassiz, in the Gulf of Mexico, and in the Caribbean Sea, 1877-79, by the
United States Coast Survey Steamer ‘“‘Blake,’? Lieutenant-Commander C.
D. Sigsbee, U.S.N., and Commander J. R. Bartlett, U.S.N., commanding.
XV. Bulletin of the Museum of Comparative Zoology, 9(2): 33-144.

—— 1908. Reports on the dredging operations off the west coast of Central
America to the Galapagos, to the west coast of Mexico, and in the Gulf
of California, in charge of Alexander Agassiz, carried on by the U.S. Fish
Commission Steamer “‘Albatross,”? during 1891. Lieut.-Commander Z. L.
Tanner, U.S.N., commanding. XXXVII. Reports on the scientific results
of the expedition to the eastern tropical Pacific, in charge of Alexander
Agassiz, by the U.S. Fish Commission Steamer ‘‘Albatross,”’ from October,
1904, to March, 1905. Lieut.-Commander L. M. Garrett, U.S.N.,
commanding. XIV. The Mollusca and Brachiopoda. Bulletin of the
Museum of Comparative Zoology, 43(6): 205-487, pls. 1-22.

Deshayes, G. P. 1855. Descriptions of new shells from the collection of Hugh

Cuming, Esq. Proceedings of the Zoological Society of London, Part XXII,
1854: 317-371.

— 1863. Catalogue des Mollusques de I’Ile de la Réunion (Bourbon). Pp.

64 NEMOURIA No. 39

1-144, pls. 28-41 [1-13]; Dentu, Paris.

Dunker, W. 1847. [In:] Pfeiffer, L. Diagnosen neuer Heliceen. Zeitschrift fiir
Malakozoologie, 4: 81-84.

— 1877. Mollusca nonnulla nova maris Japonici. Malakozoologische Blatter,
24: 67-75.

Eichwald, E. 1838. Fauna Caspii maris primitiae. Bulletin de la Société Impériale
des Naturalistes de Moscou, 11(2): 125-174.

Fischer, P. 1878. Diagnoses trochorum novorum. Journal de Conchyliologie, 26:
62-67.

— 1883. Diagnoses d’espéces nouvelles de mollusques recueillis dans le cours
de l’expédition scientifique du Talisman (1883). Journal de Conchyliologie,
31: 391-394.

Folin, A. G. L. de, Marquess [and L. Périer]. 1867-1879. Les Fonds de la
Mer. Etude internationale sur les particularités nouvelles des régions
sous-marines, etc. Tom. I-III. Paris. [Fischer, P., L. de Folin & L. Périer.
1873. Les Fonds de la Mer. Vol. 2 (cover gives date as 1872); pp. 161-208
published in 1873].

Forbes, E. & S. Hanley. 1852-1853. A history of British Mollusca, and their
shells. Volume IV. Pulmonifera and Cephalopoda. Pp. i-vi, 1-301, pls.
I-CXXXIII; John van Voorst, London.

Frembly, J. 1827. A description of several new species of chitons, found on the
coast of Chili, in 1825; with a few remarks on the method of taking and
preserving them. The Zoological Journal, 3: 193-205.

Fulton, H. 1899. A list of the species of land Mollusca collected by Mr. W.
Doherty in the Malay Archipeiago; with descriptions of some supposed new
species and varieties. Proceedings of the Malacological Society of London,
3: 212-219, pl. 11.

Gmelin, J. F. 1791. Caroli a Linné, Systema Naturae per Regna Tria Naturae,
Secundum Classes, Ordines, Genera, Species, cum characteribus, differen-
tiis, synonymis, locis. Editio Decima Tertia, Aucta, Reformata. Tom. 1(6),
Vermes, pp. 3021-3910. Georg Emanuel Beer, Lipsiae [for a collation of
this work see Kabat & Petit (1988)].

Godwin-Austen, H. H. 1883. Land and Fresh-water Mollusca of India. Part III.
Pp. 67-94, pls. 13-21; Taylor and Francis, London.

Gould, A. A. 1846. [Shells collected by the United States Exploring Expedition
under the command of Charles Wilkes]. Proceedings of the Boston Society
of Natural History, 2: 141-145.

— 1852. Mollusca and shells. Vol. 12 [In:] United States Exploring Expedition
during the years 1839-1842 under the command of Charles Wilkes U.S.N.
Boston, December 1852, xv + 510 pp. [for a complete collation of this
work, see Haskell (1942)].

Gray, J. E. 1828-30. Spicilegia Zoologica; or original figures and short systematic

September 30, 1991 Boss & Bieler 65

descriptions of new and unfigured Animals. Richard Taylor, London, Parts
I and II, pp. 12, 11 pls.

Gredler, P. V. 1887. Zur Conchylien-Fauna von China. [X. Stiick. Malakozoolo-
gische Blatter, Neue Folge, 9: 1-20.

Gundlach, J. [In:] Poey, F. 1865. Descripcion de tres moluscos terrestres.
Repertorio Fisico-Natural de la isla de Cuba, 1: 69-70.

Haas, F. 1969. [In:] Moore, Raymond C. (ed.) Treatise on Invertebrate
Paleontology. The Geological Society of America, Inc. and The University
of Kansas, Part N, Mollusca 6, Bivalvia, Volume 1 (of 3): N411-N489.

Haskell, D. C. 1942. The United States Exploring Expedition, 1838-1842, and
its publication, 1844-1874. New York Public Library, 188 pp.

Heath, H. 1911. Reports on the scientific results of the expedition to the tropical
Pacific, in charge of Alexander Agassiz, by the U.S. Fish Commission
Steamer ‘‘Albatross’”? from August, 1899, to June, 1900, Commander
Jefferson F. Moser, U.S.N., Commanding. XIV. The Solenogastres.
Memoirs of the Museum of Comparative Zoology, 45(1): 1-179, pls. 1-36.

Hertlein, L. 1969. [In:] Moore, R. C. (ed.). Treatise on Invertebrate Paleontol-
ogy. The Geological Society of America, Inc. and the University of Kansas,
Part N, Mollusca 6, Bivalvia, Volume 1 (of 3): N348-N373.

Hesse, P. 1916. Kritische Fragmente. Nachrichtsblatt der Deutschen Malakozo-
ologischen Gesellschaft, 48(3): 122-124.

Hwass, C. H. [In:] Bruguiére, J.-G. 1792. Cone. [In:] Encyclopédie Méthodique.
Histoire Naturelle des Vers. Tome Premier, Part 2, pp. 586-757; Paris,
Panckoucke.

[ICZN] International Commission on Zoological Nomenclature. 1961. Interna-
tional Code of Zoological Nomenclature. xvi (+1) + 176 pp.; International
Trust for Zoological Nomenclature, London.

[ICZN] International Commission on Zoological Nomenclature. 1971. Opinion
951. Rhyssoplax Thiele, 1893 (Amphineura): designation of a type-species
under the Plenary Powers. Bulletin of Zoological Nomenclature, 28: 18-19.

[ICZN] International Commission on Zoological Nomenclature. 1980. Opinion
1154. Drupella Thiele, 1925 (Mollusca, Gastropoda): designation of a type
species by the use of Plenary Powers. Bulletin of Zoological Nomenclature,
37: 85-88.

[ICZN] International Commission on Zoological Nomenclature. 1985. Interna-
tional Code of Zoological Nomenclature. Third Edition. xx + 338 pp.;
International Trust for Zoological Nomenclature/British Museum (Natural
History), London; University of California Press, Berkeley and Los Angeles.

Issel, A. 1869. Malacologia del Mar Rosso richerche zoologische e paleiontolo-
giche. Biblioteca Malacologica, Pisa, 387 pp., 5 pls.

Johnson, R. I. 1978. Systematics and zoogeography of Plagiola (= Dysnomia =
Epioblasma), an almost extinct genus of freshwater mussels (Bivalvia:

66 NEMOURIA No. 39

Unionidae) from middle North America. Bulletin of the Museum of
Comparative Zoology, 148(6): 239-320, 15 pls.

Kaas, P. & R. A. van Belle. 1980. Catalogue of living chitons (Mollusca:
Polyplacophora). W. Backhuys, Rotterdam, 10 unnumbered pp. + 144 pp.

Kaas, P. & R. A. van Belle. 1985a. Monograph of living chitons (Mollusca:
Polyplacophora). Volume 1. Order Neoloricata: Lepidopleurina. E. J.
Brill/W. Backhuys, Leiden, 240 pp.

Kaas, P. & R. A. van Belle. 1985b. Catalogue of living chitons (Mollusca:
Polyplacophora). Volume 2. Suborder Ischnochitonina. Ischnochitonidae:
Schizoplacinae, Callochitoninae & Lepidochitoninae. E. J. Brill/W. Backhuys,
Leiden, 198 pp.

Kaas, P. & R. A. van Belle. 1987. Catalogue of living chitons (Mollusca:
Polyplacophora). Volume 3. Suborder Ischnochitonina. Ischnochitonidae:
Chaetopleurinae, & Ischnochitoninae (pars). Additions to Vols. 1 & 2. E.
J. Brill/W. Backhuys, Leiden, 302 pp.

Kaas, P. & R. A. van Belle. 1990. Catalogue of living chitons (Mollusca:
Polyplacophora). Volume 4. Suborder Ischnochitonina. Ischnochitonidae:
Ischnochitoninae (continued). Additions to vols. 1, 2 and 3. E. J. Brill,
Leiden, 298 pp.

Kabat, A. R. & R. E. Petit. 1988. The two printings of J. F. Gmelin’s Systema
Naturae, 13th edition (1788-1796). Nautilus, 102(4): 164-166.

Keen, A. M. 1937. Nomenclatural units of the pelecypod family Cardiidae.
Bulletin du Musée royal d’Histoire naturelle de Belgique, 13(7): 1-22.

— 1961. A proposed reclassification of the gastropod family Vermetidae.
Bulletin of the British Museum Neha History), Zoology Series, 7(3):
181-213, pls. 54, 55.

— 1969. [In:] Moore, Raymond C. (ed.). Treatise on Invertebrate Paleontol-
ogy. The Geological Society of America, Inc. and the University of Kansas,
Part N, Mollusca 6, Bivalvia, Volume 1 (of 3): N230-N231, N269-N270,
Volume 2 (of 3): N583-N594, N613-N643, N852-N857.

[Kobelt, W.}] 1894. Literaturbericht. Troschel, Dr. H., Das Gebiss der Schnecken
zur Begriindung einer naturlichen Classification untersucht. Fortgesetzt von
Dr. J. Thiele . . . Nachrichtsblatt der Deutschen Malakozoologischen
Gesellschaft, 26(1-2): 25-27.

Kowalevsky, A. 1900. Comptes rendu des mes études zoologiques a Sebastopol
pendant 1’été de 1899 (Communication préliminaire) [Otchet’’ 0 moikh”’
zoologicheskikh”’ v’’ Sevastopoli litom”’ 1899 goda]. Bulletin de 1‘Académie
Impériale des Sciences de St.-Pétersbourg [Izvestiya Imperatorskoi Akade-
mii Nauk, S. Peterburg], (5)12(2): 193-204.

—— 190la. Sur le genre Chaetoderma. Archives de Zoologie Expérimentale et
Générale, 9: 261-283, pls. 10-12.

—— 1901b. Les Hedylides. Etude anatomique. Mémoires de l’Académie Impériale

September 30, 1991 Boss & Bieler 67

des Sciences de St. Pétersbourg, (8)12(6): 1-32, pls. 1-5.

Krause, A. 1885. Ein Beitrag zur Kenntniss der Mollusken-Fauna des Ber-
ingsmeeres. Brachiopoda und Lamellibranchiata. Archiv ftir Naturges-
chichte, 51: part 1, 14-40, pl. 3; part 2, 256-302, pls. 16-18.

Krauss, F. 1848. Die Sidafrikanischen Mollusken. Von Ebner & Seubert,
Stuttgart, [i-iv] + 1-140 pp, 6 pls.

Kiister, H. C. 1852. Die Gattungen Paludina, Hydrocaena und Valvata. [In:]
Systematisches Conchylien-Cabinet von Martini und Chemnitz. 2nd Edi-
tion. Bauer und Raspe, Niirnberg, 1(21): 96 pp., 14 pls.

Lamarck, J. B. P. A. d. M. d. 1798. Tableau Encyclopédie et Méthodique des
trois régnes de la nature. Vers, coquilles, mollusques et polypiers. Agasse,
Paris, 2(3): pls. 287-390.

— 1810. Suite de la détermination des espéces de mollusques testacés.
Continuation du genre porcelaine, et genre ovule. Annales du Muséum
National d’Histoire Naturelle, Paris, 16: 89-114.

— 1816. Tableux Encyclopédique et méthodique des trois régnes de la nature.
Vers, coquilles, mollusques et polypiers. 3(4): 391-488; Agasse, Paris.

—— 1822. Histoire naturelle des animaux san vertebres, . . . 7: 1-711. Chez
auteur, au Jardin de Roi, Paris.

— 1827. Tableau Encyclopédie et Méthodique des trois régnes de la nature.
Vers, coquilles, mollusques et polypiers. Agasse, Paris, 1: i-viii, 1-180.

Le Guillou, M. 1845 (May). Description de six Hélices nouvelle. Revue
Zoologique par la Société Cuvierienne, 8: 187-188.

Lea, I. 1829. No. XIII. Description of a new genus of the family of naiades,
including eight species, four of which are new; also the description of eleven
new species of the genus Unio from the rivers of the United States: with
observations on some of the characters of the naiades. Transactions of the
American Philosophical Society, new series, 3(4): 403-457.

—— 1844. Continuation of Mr. Lea’s paper on fresh water and land shells.
Transactions of the American Philosophical Society, new series, 9 (1846):
1-31.

Leche, W. 1878. Ofversigt 6fver de af Svenska Expeditionerna till Novaja Semlja
och Jenissej} 1875 och 1876 insamlade hafs-mollusker. Kongliga Svenska
Vetenskaps-Akademiens Handlingar, 16(2): 1-86, pls. 1-2.

Linnaeus, C. [afterwards Carl von Linné]. 1758. Caroli Linnaei . . . Systema
Naturae per Regna tria Naturae, secundum classes, ordines, genera, species,
cum characteribus, differentiis, synonymis, locis ... Editio decima
reformata. Impensis Direct. Laurentii Salvii, Holmiae, Tom. |. Animalia.
[iv] + 823 pp.

—— 1767. Caroli a Linné ... Systema Naturae per Regna tria Naturae,
secundum classes, ordines, genera, species, cum characteribus, differentiis,
synonymis, locis . . . Editio duodecima, reformata. Impensis Direct.

68 NEMOURIA No. 39

Laurentii Salvii: Holmiae, Tom. 1. Regnum Animale. Pars II. Classis V-VI,
Nomina Generica. Nomina specierum propria: Nomina trivialia. Nomina
trivialia Papilionum and Phalaenarum. Synonyma. Termini Artis. Appendix
Synonymorum. Addenda. Errata, pp. 533-1327.

Martens, E. von. 1870. Conchylien aus dem obern Nilgebiet. Malakozoologische
Blatter, 17: 32-36.

— 1876. Die von Prof. Dr. R. Buchholz in Westafrika gesammelten Land-und
Stisswasser-Mollusken. Monatsberichte der KGniglichen Preussischen Akade-
mie der Wissenschaften zu Berlin, 1876: 253-274, 5 pls.

— 1878. Vebersicht der von Hrn. J. M. Hildebrandt in Ost-Afrika gesammelten
Land- und Siisswasser-Conchylien. Monatsberichte der KGniglichen Preus-
sischen Akademie der Wissenschaften zu Berlin, 1878: 288-299, 2 pls.

— 1880. Herr v. Martens zeigte einige Landschnecken vor, welche Dr. O.
Finsch auf den Karolinen gesammelt hat. Sitzungsberichte der Gesellschaft
Naturforschender Freunde zu Berlin, 1880: 143-147.

— 1885. Vorlaufige Mittheilung tiber die Molluskenfauna von Stid-Georgien.
Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin,
1885: 89-94.

— 1897. Stiss- und Brackwasser Mollusken des Indischen Archipels. [In:]
Zoologische Ergebnisse einer Reise in Niederlandisch Ost-Indien, 4: 1-331,
12 pls.

— 1901. Einige neue Meer-Conchylien von der deutschen Tiefsee-Expedition.
Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin,
1901: 14-26.

— 1903. Herr von Martens legte einige neue Arten von Meer-Conchylien aus
den Sammlungen der deutschen Tiefsee-Expedition under der Leitung von
Prof. Carl Chun 1890-99 vor. Sitzungsberichte der Gesellschaft Natur-
forschender Freunde zu Berlin, 1902: 237-244.

—- 1904. Systematisch-geographischer Theil. [In:] Die beschalten Gastropoden
der deutschen Tiefsee-Expedition, 1898-1899. Wissenschaftliche Ergebnisse
der deutschen Tiefsee-Expedition auf dem Dampfer ‘‘Valdivia’”’ 1898-1899,
7(A): 1-146, pls. 1-5 [title page states 1903").

Martens, E. von & G. Pfeffer. 1886. Die Mollusken von Stid Georgien nach der
Ausbeute der Deutschen Station, 1882-1883. Jahrbuch der Hamburgischen
Wissenschaftlichen Anstalten, 3: 63-135, 4 pls.

Martens, E. von & F. Wiegmann. 1898. Land- und Stisswasser-Mollusken der
Seychellen. Mitteilungen aus der Zoologischen Sammlung des Museums
fur Naturkunde in Berlin, 1(1): 1-94, pls. 1-4. [Conchyliologischer Theil
by von Martens; Zootomischer Theil by Wiegmann].

Melvill, J. C. & R. Standen. 1901. The Mollusca of the Persian Gulf, Gulf of

Oman and Arabian Sea, as evidenced mainly through the collections of
Mr. F. W. Townsend, 1893-1900; with descriptions of new species.

September 30, 1991 Boss & Bieler 69

Proceedings of the Zoological Society of London, 1901: 327-460. pls. 21-24.

Menke, K. T. 1843. Molluscorum Novae Hollandiae Specimen, quod ad
celebrandum diem jubilaeum vicesimum sextum mensis februarii anni
MDCCCXLIII quo per viginti quinque annos professoris muneri in illustri
gymnasio academico hamburgensi parefuit vir doctissimus, clarissimus
Joannes Georgius Christianus Lehmann, med. et philos. doctor, &c., scripsit
Carolus Theodorus Menke. Hannoverae, in libraria Aulica Hahniana, pp.
46.

Metcalfe, W. 1851. An enumeration of species of Recent shells, received by W.
J. Hamilton, Esq., from Borneo, in November 1850, with descriptions of
new species. Proceedings of the Zoological Society of London, 1851: 70-74.

Mikkelsen, P. S. & P. M. Mikkelsen. 1984. Comparison of Acteocina canaliculata
(Say, 1826), A. cande: (d’Orbigny, 1841), and A. atrata spec. nov.
(Gastropoda: Cephalaspidea). The Veliger, 27(2): 164-192, 13 text-figs.

Mollendorff, O. F. von. 1882. Diagnoses specierum novarum Chinae meridion-
alis. Jahrbuch der Deutschen Malakozoologischen Gesellschaft, 9: 179-188.

— 1883. Materialien zur Fauna von China. Jahrbuch der Deutschen Malakozo-
ologischen Gesellschaft, 10: 228-269, pl. 8.

Morlet, L. 1884. Description d’espéces nouvelles de coquilles, recueillies par
M. Pavie, au Cambodge. Journal de Conchyliologie, 32: 386-403, pls.
11-13.

Murdoch, R. 1899. Description of Sigaretus? drewi, n.sp. (fossil), and Cirsonella?
neozelandica, n. sp. from New Zealand; with notes on some New Zealand
land Mollusca. Proceedings of the Malacological Society of London, 3:
320-325, pl. 16.

Nesis, K. N. 1987. Cephalopods of the world. Translated from Russian by B.
S. Levitov & edited by Lourdes A. Burgess. T.F.H. Publications, Neptune
City, New Jersey, 351 pp., 88 figs.

Nevill, G. 1880. New species of brackish-water mollusks. Journal of the Asiatic
Society of Bengal, 49(2): 159-166.

—— 188la. Note on little known Mollusca of the Indo-Malayan Fauna. Journal
of the Asiatic Society of Bengal, 50(2): 125-167, pls. 5-7.

—— 1881lb. Description of a new species of Rostellaria from the Bay of Bengal.
Journal of the Asiatic Society of Bengal, 50(2): 262.

Newcomb, W. 1864. Description of a new species of Pedicularia. Proceedings of
the California Academy of Sciences, 3: 121-122.

Newell, N. D. 1969. [In:] Moore, R. C. (ed.). Treatise on Invertebrate
Paleontology. The Geological Society of America, Inc. and the University
of Kansas, Part N, Mollusca 6, Bivalvia, Volume 1 (of 3): N261-N267.

Nierstrasz, H. F. 1903. Neue Solenogastren. Zoologische Jahrbiicher, Abteilung
fiir Anatomie, 18: 359-386, 2 pls.

70 NEMOURIA No. 39

Odhner, N. H. 1968. Opisthobranches. [In:] Franc, A. Gastéropodes. [In:] P.-P.
Grassé (ed.). Traité de Zoologie, 5: 608-893. Masson, Paris.

Orbigny, A. d’. 1842. Mollusques. [In:] Sagra, R. Histoire Physique, Politique
et Naturelle de l’Ile de Cuba. Paris, vol. 2, 264 pp., 28 pls.

Pease, W. H. 1865. Descriptions of new species of land shells from the islands
of the Central Pacific. Proceedings of the Zoological Society of London,
1864: 668-676.

— 1868. Descriptions of sixty-five new species of marine gasteropodae,
inhabiting Polynesia. American Journal of Conchology, 3: 271-297, pls.
23-24.

Petit de la Saussaye, S. 1843. Descriptions of new species of shells belonging to
the genus Auncula, collected by H. Cuming, Esq. Proceedings of the
Zoological Society of London, Part X, 1842: 201-202.

Petterd, W. F. 1879. New species of Tasmanian marine shells. Journal of
Conchology, 2: 102-105.

Pfeiffer, L. 1839. Bericht tiber die Ergebnisse meiner Reise nach Cuba im Winter
1838-1839. Archiv ftir Naturgeschichte, 5(1): 346-358.

Philippi, R. A. 1836. Enumeratio molluscorum Siciliae cum viventium tum in
tellure Tertiaria fossilium quae in itinere su observavit. 1: i-xiv + 1-268
pp., pls. 1-12. Schroppii, Berolini [Berlin].

— 1844. Enumeratio molluscorum Siciliae cum viventium tum in tellure

Tertiaria fossilium quae in itinere su observavit. 2: i-iv + 1-304 pp., pls.
13-28. Anton, Halis Saxonum [Halle].

— 1846. Diagnosen neuer Conchylien-Arten. Zeitschrift fir Malakozoologie,
3: 47-55.

Pilsbry, H. A. 1895. Polyplacophora. Acanthochitidae, Cryptoplacidae and
Appendix. Tectibranchia. Manual of Conchology (1)15(60): 181-436, pls.
43-50, 59-61.

—— 1932. Libera or Garrettina (Endodontidae). The Nautilus, 46(2): 70-71.

— 1946. Land Mollusca of North America (North of Mexico). Monograph
of the Academy of Natural Sciences of Philadelphia, Number 3, 2(1): 1-vi,
1-520, figs. 1-281.

Ponder, W. F. 1982. The anatomy and relationships of Hydrococcus braziert CT.
Woods). Journal of Molluscan Studies, 48: 64-79, 21 text-figs.

—— 1985. A review of the genera of the Rissoidae (Mollusca: Mesogastropoda:
Rissoacea). Records of the Australian Museum (1984), Supplement 4:
1-221, 153 figs.

Ponder, W. F. and A. Warén. 1988. Appendix. Classification of the Caenogastro-
poda and Heterostropha-—A list of the family-group names and higher taxa.
Malacological Review, Supplement 4: 288-326.

Potiez, V.-L.-V. & A.-L.-G. Michaud. 1838. Galerie des Mollusques ou catalogue

September 30, 1991 Boss & Bieler Toh

méthodique, descriptif et raisonné des mollusques et coquilles du Muséum
de Douai. Bailliére, Paris, pp. [i]-xxxvi, 1-560, [1-4]; Atlas, pp. 1-79, 1-70.

Powell, A. W. B. 1960. Antarctic and Subantarctic Mollusca. Records of the
Auckland Institute and Museum, 5(3 and 4): 117-193.

Pruvot, G. 1891. Sur organisation de quelques Néoméniens des cétes de France.
Archives de Zoologie Expérimentale et Générale, 9: 699-805.

Pruvot-Fol, A. 1954. Mollusques Opisthobranches. Faune de France, 58.
Lechevalier, Paris, 460 pp.

Putzeys, S. 1899. Diagnoses de quelques coquilles et d’un sous-genre nouveau
provenant de |’etat indépendant du Congo. Annales de la Société Royale
Malacologique de Belgique, 34: L-LX, figs. 1-16.

Quadras, J. F. & O. F. de Moellendorff. 1894. Diagnoses specierum novarum a
J. F. Quadras in insulis Mariannis collectarum scripserunt. Nachrichtsblatt
der Deutschen Malakozoologischen Gesellschaft, 26: 33-42.

Quadras, J. F. & O. F. de Moellendorff. 1896. Diagnoses specierum novarum
ex insulis Philippinis auctoribus. Nachrichtsblatt der Deutschen Malakozo-
ologischen Gesellschaft, 28: 1-15.

Quoy, J.J. C. & J. P. Gaimard. 1832-1835. Voyage de Découvertes de |’Astrolabe.
Zoologie. J. Tastu, Paris, 2(1): 1-320 (1832), (2): 321-686 (1833); 3(1):
1-366 (1834), (2): 367-954 (1835); Atlas, Mollusques, plates 1-93 (1833).

Rang, S. 1831. Descriptions des coquilles terrestres recueillies pendant un voyage

a la cdte occidentale d’Afrique, et au Brésil. Annales des Sciences
Naturelles, Paris, 24: 5-63, pls. 1-3.

Reeve, L. A. 1844 June). Conchologica Iconica: or, illustrations of the shells of
molluscous animals. Vol. II, Containing monographs of the genera. . .
Area... ., species 115,-plate 17, fig. 115.

— 1847a (May). Conchologica Iconica: or, illustrations of the shells of
molluscous animals. Vol. IV, Monograph of the genus Chiton, plates 15
and 23.

— 1847b. Descriptions of new species of shells collected in the Eastern
Archipelago by Capt. Sir Edward Belcher and Mr. Adams during the
voyage of H.M.S. Samarang. Proceedings of the Zoological Society of
London, 1847: 24-26.

—— 1848 (December). Conchologica Iconica: or, illustrations of the shells of
molluscous animals. Vol. V, Containing monographs of the genera Bulimus
» s plate 256.

—— 1854 (December). Conchologica Iconica: or, illustrations of the shells of
molluscous animals. Vol. VII, Containing a monograph of the genus Helix,
plate 208.

Reinhardt, O. 1883. Herr O. Reinhardt legte einige ihm von Herrn Dr. W.
Kobelt in Schwanheim a. M. zur Begutachtung wtbersandte, von Herrn
Hungerford gesammelte japanische Hyalinen vor. Sitzungsberichte der

72 NEMOURIA No. 39

Gesellschaft Naturforschender Freunde zu Berlin, 1883: 82-86.

Risbec, J. 1928. Contribution a |’étude des nudibranches Néo-calédoniens. Faune
des Colonies Francaise. Société d’Editions, Paris, 2(1): 1-328, pls. 1-12,
text-figs. 1-98.

Salisbury, A. E. 1934. On the nomenclature of Tellinidae, with descriptions of
new species and some remarks on distribution. Proceedings of the
Malacological Society of London, 21: 74-91, pls. 9-14.

Salvini-Plawen, L. von. 1978. Antarktische und subantarktische Solenogastres.
Eine Monographie: 1898-1974. Zoologica, 44(2/128/2): 157-315, pls. I-II.

Sars, G. O. 1878. Bidrag til Kundskaben om Norges arktiske Fauna. I. Mollusca
Regionis Arcticae Norvegiae. Oversigt over de i Norges arktiske region
forekommende bldddyr. Universitets program for forste halvaar 1878.
Brogger, Christiania, 466 pp.

Say, T. 1821. Descriptions of univalve shells of the United States. Journal of the
Academy of Natural Sciences of Philadelphia, 2: 149-179.

— 1826. Descriptions of marine shells recently discovered on the coast of the
United States. Journal of the Academy of Natural Sciences of Philadelphia,
5: 207-221.

Schepman, M. M. 1903. Descriptions of three new species of Oliva from the
Siboga Expedition. Tijdschrift der Nederlandsche Dierkundige Vereeniging,
Leiden, (2)8: 67-69

— 1911. The Prosobranchia of the Siboga Expedition. Part IV. The Rachiglossa.
Siboga-Expeditie, 49d: 243-363, 7 pls.

— 1911 [1913]. The Prosobranchia of the Siboga Expedition. Part V.
Toxoglossa. Siboga-Expeditie, 49e: 365-452, 6 pls., 1 text-fig.

Sherborn, C. D. & B. B. Woodward. 1906. On the dates of publication of the
natural history portions of the “Encyclopédie Méthodique.” Annals and
Magazine of Natural History, (7)17(97): 577-582.

Smith, A. G. 1960. Amphineura. [In:] Moore, R. C. (ed.). Treatise on
Invertebrate Paleontology. The Geological Society of America, Inc. and
the University of Kansas, Part I, Mollusca 1: 141-176, figs. 31-45.

Smith, E. A. 1893. Descriptions of new species of Helix, Rissoina and Actaeon.
The Conchologist, 2(5): 97-100, text-figs.

— 1902. Mollusca. Report on the collections of natural history made in the
antarctic regions during the voyage of the “Southern Cross.’’ London, pp.
201-213, pls. 24, 25.

—— 1903. Descriptions of new species of land shells from Central Africa. Journal
of Conchology, 10(10): 315-319, pl. 4.

—— 1904a. On a collection of marine shells from Port Alfred, Cape Colony.
The Journal of Malacology, 11(2): 21-43, pls. 2-3.

—— 1904b. Natural history notes from H. M. Indian Marine Survey Steamer

September 30, 1991 Boss & Bieler 73

‘Investigator,’ Commander T. H. Heming, R.N. Series III, No. 1. On
Mollusca from the Bay of Bengal and the Arabian Sea. Annals and
Magazine of Natural History, (7)14: 1-14.

— 1907. Mollusca II. Gastropoda. [In:] National Antarctic (‘““Discovery’’)
Expedition, 1901-1904. Natural History, Vol. II, Zoology. (Vertebrata;
Mollusca; Crustacea). British Museum (Natural History), London, pp.
1-12, pls. 1-2.

Sowerby, G. B. [I]. 1825. A catalogue of the shells contained in the collection
of the late Earl of Tankerville, arranged according to the Lamarckian
Conchological System; together with an appendix containing descriptions
of many new species. Stirling, London, pp. i-vii, 1-92, i-xxxiv, 9
unnumbered plates.

— 1832. Characters of new species of Mollusca and Conchifera, collected by
Mr. Cuming. Proceedings of the committee of science and correspondence
of the Zoological Society of London, Part II, 1832: 25-33, 50-61, 104-108.

— 1838. A descriptive catalogue of the species of Leach’s genus Margarita.
The Malacological and Conchological Magazine, London, Part 1, pp.
23-27.

Sowerby, G. B., Junior [II]. 1842. Thesaurus Conchyliorum or figures and
descriptions of Recent shells. Part II. Containing monographs of the genera
Pteroceras, Pecten, and Hinnites. Sowerby, London, pp. 41-82, pls. 11-20.

—— 1844. Thesaurus Conchyliorum or figures and descriptions of Recent shells.
Part IV. Containing monographs of the genera Scalaria, and Columbella.
Sowerby, London, pp. 83-146, pls. 32-50.

— 1855. Thesaurus Conchyliorum or figures and descriptions of Recent shells.
Part XVI. Containing a monograph of Cerithium . . . Sowerby, London,
pp. 847-899, pls. 176-186.

—— 1873. Descriptions of twelve new species of shells. Proceedings of the
Zoological Society of London, 1873: 718-722, pl. 59.

Sowerby, G. B. III. 1900. On some marine shells from Pondoland and the
Kowie, with descriptions of seventeen new species. Proceedings of the
Malacological Society of London, 4: 1-7, pl. 1.

—— 1905. Descriptions of seven new species of marine Mollusca from the
collection of the late Admiral Keppel. Proceedings of the Malacological
Society of London, 6: 279-282, 6 text-figs.

Stenzel, H. B. 1971. [In:] Moore, R. C. (ed.). Treatise on Invertebrate
Paleontology. The Geological Society of America, Inc. and the University
of Kansas, Part N, Mollusca 6, Bivalvia, Volume 3 (of 3): N953-N1124.

Strebel, H. 1905. Beitrage zur Kenntnis der Molluskenfauna der Magalhaen-
Provinz. No. III. Zoologische Jahrbticher, Abteilung fiir Systematik, 22:
575-666, 4 pls.

— 1908. Die Gastropoden (mit Ausnahme der nackten Opisthobranchier).

74 NEMOURIA No. 39

Wissenschaftliche Ergebnisse der Schwedischen Stidpolar-Expedition 1901-
1903, 6(1): 1-112, 6 pls.

Sykes, E. R. 1903. Descriptions of new land shells from the Austro-Malayan
Region. Journal of Malacology, 10: 64-67.

Tate, R. 1893. On some new species of Australian marine Gastropoda.
Transactions of the Royal Society of South Australia, 17: 189-197, pl. 5.

Tenison-Woods, J. E. 1876. Descriptions of new Tasmanian shells. Papers and
Proceedings of the Royal Society of Tasmania for 1875: 134-162.

Thiele, J. [see Bieler & Boss 1989].

Tomlin, J. R. le B. 1936. Book notes. Proceedings of the Malacological Society
of London, 22(3): 135-137.

Valenciennes, A. [In:] Lamarck, J. B. P. A. d. M. d. 1827. Tableau Encyclopédie
et Méthodique des trois regnes de la nature. Vers, coquilles, mollusques
et polypiers. Agasse, Paris, 1: i-viii, 1-180 [p. 154 is caption to Plate 287
with figure 3 listed as Téllina plicata, Val.].

Vaught, K. C. 1989. A Classification of the Living Mollusca. American
Malacologists, Inc., Melbourne, Florida, xii + 195 pp.

Vélain, C. 1877. Passage de Vénus sur le Soleil (9 Décembre 1874). Expédition
francaise aux Iles Saint-Paul et Amsterdam. Zoologie. Observation générales
sur la faune des deux iles suivies d’une description des mollusques. Archives
de Zoologie Expérimentale et Générale, 6: 1-144, pls. 1-5.

Vokes, H. E. 1980. Genera of the Bivalvia: a systematic and bibliographic
catalogue (revised and updated). Paleontological Research Institution,
Ithaca, New York, xxvii + 307 pp.

Walker, B. 1915. Apical characters in Somatogyrus, with descriptions of three new
species. The Nautilus, 29: 37-41; 49-53, 4 text-figs.

Warén, A. 1980. Marine Mollusca described by John Gwyn Jeffreys, with the
location of the type material. Conchological Society of Great Britain and
Ireland, Special Publications No. 1, 60 pp., 8 pls.

Watson, R. B. 1882. Mollusca of H.M.S. ‘Challenger’ Expedition. Part XIII.
Journal of the Linnean Society of London, 16: 358-372.

— 1886. Report on the Scaphopoda and Gasteropoda collected by H.M.S.
‘Challenger’ during the years 1873-1876. Report on the Scientific Results
of the Challenger Expedition, Zoology, 15(42): 1-756, pls. 4-50.

Wenz, W. 1938-1944. Gastropoda. Teil 1: Allgemeiner Teil und Prosobranchia.
[In:] Schindewolf, O.H. (ed). Handbuch der Paldozoologie, 6(1): 1-xii,
1-1639. Borntraeger, Berlin [pp. 1-240, March 1938; 241-480, October
1938; 481-720, July 1939; 721-960, August 1940; 961-1200, October
1941; 1201-1506, October 1943; 1507-1639, i-xii, November 1944].

Yen, T.-C. 1939. Die chinesischen Land- und Sitisswasser-Gastropoden des
Natur-Museum Senckenberg. Abhandlungen der Senckenbergischen Natur-

September 30, 1991 Boss & Bieler ihe

forschenden Gesellschaft, 444: 1-233, 16 pls.

Zilch, A. 1959-1960. Gastropoda von Wilhelm Wenz. Teil 2. Euthyneura. [In:]
Schindewolf, O.H. (ed). Handbuch der Paléozoologie, 6(2): i-xii, 1-835,
2515 figs. Borntraeger, Berlin [pp. 1-200, 17 July 1959; 201-400, 25
November 1959; 401-600, 30 March 1960; 601-835, i-xii, 15 August
1960].

76 NEMOURIA No. 39
ACKNOWLEDGMENTS

We thank Prof. Dr. Rudolf Kilias (Museum fiir Naturkunde, Humboldt
Universitat, Berlin) for providing Thiele’s photograph as well as the
sample of his handwriting, and Ms. Kathie Way (The Natural History
Museum, London) for a copy of Thiele’s 1923 letter to that museum.
Dr. Alan Kabat (National Museum of Natural History, Washington)
directed our attention to omissions in our previous paper; Ms. Paula
M. Mikkelsen kindly supplied information on Thiele’s opisthobranch
taxa, and Mr. Richard I. Johnson commented expertly on an earlier draft
of the manuscript.

The manuscript was reviewed by Dr. Alan R. Kabat (National Museum
of Natural History, Washington, DC), Ms. Paula M. Mikkelsen (Harbor
Branch Oceanographic Museum, Ft. Pierce, FL) and Mr. Richard E.
Petit (North Myrtle Beach, SC).

September 30, 1991 Boss & Bieler fe)

ADDITIONS AND CORRECTIONS TO PART 1
(Nemouria, 34, 1989):

Page 3, fourth paragraph: for 1960, read 1860.
Page 5, last paragraph: for 1911f, read 1911b.

Add asterisks to the following entries in section A. Publications on
Mollusca: 1925d (p. 21), 1928d (p.22).

Previously omitted malacological publications by J. Thiele:

Carl Chun (1852-1914) is best known among malacologists for his
fundamental monograph on the ‘Valdivia’ cephalopods. He died before
completing this work and Thiele wrote several sections toward the end.
Thiele should be credited with the authorship of those new taxa (as
“Thiele in Chun 1915’’):

* 1915d. Polypus (pp. 485-487, English edition pp. 378-380), Boli-
taenidae (pp. 490-494, English edition pp. 383-386), Amphitre-
tidae (pp. 531-533, English edition pp. 415-416), Vampyro-
teuthidae (pp. 534-536, English edition pp. 419-420), Cirro-
teuthidae (pp. 536-538, English edition pp. 421-422). [In:]
Chun, C. Die Cephalopoden. II. Teil: Myopsida, Octopoda.
Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition
auf dem Dampfer ‘‘Valdivia” 1898-1899, 18(2); Jena (Gustav
Fischer). [English language translation: Carl Chun (C. F. E.
Roper & I. H. Roper, eds.), The Cephalopoda. Israel Program
for Scientific Translations, Keter Publishing House Jerusalem
Ltd., 1975]

Another posthumous publication contains some of Thiele’s contributions
to the Ktikenthal-Krumbach Handbuch der Zoologie. As this work was
issued in signatures, his last portion on cephalopods (pp. 257-258) was
not published until 1956; it was apparently issued with S. Jaeckel’s
addendum to the Mollusca (pp. 259-275):

1956. Mollusca = Weichtiere. [In:] Kiikenthal, W. & Krumbach, T.
(eds.): Handbuch der Zoologie. Eine Naturgeschichte der Stamme
des Tierreiches, 5(4): 257-258, text-figs. 341-343; Berlin and
Leipzig (Walter de Gruyter), 1956 [posthumously continued from
1925d, 1926a and 1926b].

Founding Editor Managing Editor
JOHN E. DU PONT RUDIGER BIELER

Consulting Editors

WILLIAM K. EMERSON, American Museum of Natural History, New
York, NY.

M. G. HARASEWYCH, National Museum of Natural History, Smithsonian
Institution, Washington, DC.

KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh,
PA.

RICHARD W. THORINGTON, JR., National Museum of Natural History,
Smithsonian Institution, Washington, DC.

Published by the

DELAWARE MUSEUM OF NATURAL HISTORY
4840 Kennett Pike, Wilmington,
Delaware 19807, U.S.A.

BT!
N4YAG
NH
NEMOURIA

Occasional Papers of the Delaware Museum of Natural History

NUMBER 40 DECEMBER 5, 1995

A SMALL COLLECTION OF BIRDS
FROM THE PHILIPPINES
WITH NOTES ON BODY MAS

Johannes Erritzoe’

ABSTRACT. The importance of avian body mass is well understood. In
this paper, masses of 199 individuals from 84 species of Philippine birds
are reported, along with sex, month, and the bird’s condition; interesting
occurrences and plumage notes are presented for 12 other specimens.

INTRODUCTION

Many have already outlined the value of avian body mass data as one
of the most important avian measurements (e.g., Clench & Leberman,
1978; Thomas, 1982; Dunning, 1984; Smith, et al., 1986). Several
papers have supplied weight data for Philippine birds. For example,
Amadon & Jewett (1946) gave the mass of 31 species; Ripley & Rabor
(1958), 483; Ripley & Rabor (1962), 7; and Parkes (1988), 1. But I
know of no papers where the sex, month, and the bird’s condition have
been reported.

The mass of a bird is biologically more meaningful when related to
its fat index (Clench & Leberman, 1978). In this paper, masses of 199
individuals from 84 species are listed; 12 others for which the mass is not
given but which have other interesting information are discussed at the
end of the paper along with the more unusual cases.

"House of Bird Research, Taps Old Rectory, DK-6070 Christiansfeld, Denmark.

2 NEMOURIA No. 40

MATERIALS AND METHODS

The birds were all collected on Luzon and Palawan between 1965
and 1980, most of them by Hartvig Jensen, and are now housed in
Denmark. The collection is available for scientific research.

Masses were obtained using Pesola spring balances of 50, 100 and
300 g. Only birds which were sexed by dissection and by gonad mea-
surement are included in this paper, with one exception (Porzana pusilla).
Age determination was made by studying skull ossification, the size of
gonads, the form of the oviduct, and the presence or absence of bursa
Fabricii, all methods described by Erritzoe (1985). When the color of
bare parts described on the bird-skin labels differed from the literature
(du Pont, 1971; Ripley, 1977; Hayman, et al., 1986), it is so stated in
the species account. Any interesting information about stomach contents
is also given. Taxonomy follows Dickinson, et al. (1991), with one excep-
tion (Pluvialis fulva). Birds were identified only to species level.

Abbreviations used are: ad, adult; bF, bursa Fabricii; P, purchased
in the bird pet-market in Manila; imm, immature; 1, lean (breastbone
keen, lacking 1/3 or more of breast-muscle); I-n, a little lean (breastbone
a little sharp); n, normal (no or only very little fat visible); n-f, a little
fat; f, fat (fat fills furcula area which is level); vf, very fat (furcular area
bulges with fat).

Sites (all Luzon except Iwahig) mentioned in the text are: Alicia (16°
80’ N, 121° 75’ E); Apalit (14° 57’ N, 120° 46’ E); Aparri (18° 22’ N,
121° 39’ BE); Bataan Province (14° 30’ N, 120° 25’ E); Bulacan
Province (15° 00’ N, 124° 05’ E); Cabanatuan (15° 29’ N; 1207/58 15);
Cagayan Province (18° 00’ N, 121° 50’ EE); Casq (14° 40’ N, 121° 30’
E); Chico River (17° 37’ N, 121° :30’ E); Clayeria (18° 37’ NvJ217.05-
E); Conner (17° 50’ N, 121° 10’ E); Gonzaga (18° 15’ N, 122° 20’ BE);
Isabela Province (17° 00’ N, 122° 00’ E); Iwahig, Palawan (09° 45’ N,
118° 40’ E); Jones (16° 50’ N, 121° 70’ E); Laguna de Bai (14° 23’ N,
121° 15’ E); Laguna Kaliraya (14° 10’ N, 121° 35’ E); Los Bafios (14°
13’ N, 121° 11’ E); Maddela (16° 21’ N, 121° 41’ E); Magot River (16°
25’ N, 121° 17’ E); Malamag Province (17° 17’ N, 121° 48° E)3) Me:
Maquiling (14° 08’ N, 121° 12’ E); Munting Buhangin (14° 02’ N, 120°
30’ E); Nassiping (17° 58’ N, 121° 37’ E); Palaui Island (18° 35’ N,
122° 08’ E); Pata (17° 38’ N, 121° 30’ E); Pefia Blanca (17° 40’ N, 121°
45’ E); Punta (12° 50’ N, 121° 37’ E); San Mariano (17° 00’ N, 122°
00’ E); Santa Ana (18° 30’ N, 122° 20’ E); Santa Fe (16° 10’ N, 120°
57' E); Santa Teresita Beach (18° 30’ N, 122° 00’ E); Tabang (17° 50’
N, 121° 307.E):

December 5, 1995 Johannes Erritzoe 3

SYSTEMATIC LIST
FAMILY RALLIDAE

Gallirallus philippensis (Linne, 1766)
2 ad, 15 October, Apalit. 172 g (n-f); largest eggs in ovary 7.9, 6.8,
4.4, and 3.9 mm.

Gallirallus striatus (Linné, 1766)
6 ad, 13 February, Laguna de Bai. 98 g (n); iris brown, bill red-
dish-brown, legs and feet slate-grey.
6 ad, 01 March, Laguna de Bai. 128 g (n). Stomach contents: fresh-
water snails.

Gallirallus torquatus (Linné, 1766)
6 ad, 25 February, Laguna de Bai. 276 g (f).

Rallina eurizonoides (Lafresnaye, 1845)
? ad, 20 April, Pefa Blanca. 79 g (n).

Porzana pusilla (Pallas, 1776)
6, 06 February, Laguna de Bai. Found decayed. Det. Kenneth C.
Parkes.
2, 21 October, Santa Fe. 39 g (f); iris orange-red, bill and legs
greenish. Habitat: tall grass. Stomach contents: rice.

Porzana fusca (Linné, 1766)
6 imm, 03 December, Santa Fe. 51 g, bF 9.0 x 3.7 mm. Stomach
contents: rice.
¢ imm, 06 December, Santa Fe. 51 g, bF 7.5 x 3.5 mm.
? imm, 03 December, Santa Fe. 40 g (n), bF approx. 4.4 x 1.9 mm;
iris and eye-ring orange-red, bill dark green, legs dirty red.

Porzana tabuensis (Gmelin, 1789)
6 imm, 04 December, Santa Fe. 33 g (f), bF 7.3 x 3.9 mm; iris
orange, legs red-brown.

Porzana cinerea (Vieillot, 1819)
3, 21 May, Alicia. 65 g.
2 ad, 21 May, Alicia. 72 g.

4 NEMOURIA No. 40

Amaurornis phoenicurus (Pennant, 1769)
2 ad, 08 February, Laguna de Bai. 120 g, bF 11.6 x 5.4 mm.
2 imm, 13 February, Laguna de Bai. 141 g.

Gallicrex cinerea (Gmelin, 1789)
3d ad, 20 May, Alicia. 443 g (n); testes: left 19.4 x 13.8, right 21.0
x 11.5 mm; iris brown.
2 ad, 20 May, Alicia. 292 g; largest egg 4.7 mm.

FAMILY ROSTRATULIDAE

Rostratula benghalensis (Linné, 1758)
6 ad, 12 February, Tabang. 145 g; iris brown.
6 ad, 13 February, Laguna de Bai. Iris dark brown.
2 ad, 26 March, Cabanatuan. 163 g; largest egg 8.0 mm.
2 ad, 24 April, Chico River.
? ad, 20 May, Alicia. 178 g (n); small feathers in molt; largest eggs:
12.8, 7.7, and 4.3 mm.

FAMILY CHARADRIIDAE

Pluvialis fulva (Gmelin, 1789)
6 ad, 24 April, Chico River. 136 g (f).
2? imm, 08 March, Cagayan. Legs blue-grey. Stomach contents:
small snails.
2 imm, 31 March, Aparri. Stomach contents: small snails.

Charadrius dubius Scopoli, 1786
3d ad, 26 May, Chico River. 33 g (n).
? ad, 10 March, Magot River. 34 g (n); largest eggs: 2.6, 2.5, 2.2,
and 1.7 mm.

Charadrius alexandrinus Linné, 1758
6 imm, 31 March, Aparri. 43 g (I).
2 imm, 03 March, Aparri. 39 g (I).
2 imm, 22 March, Aparri. 38 g (1).

Charadrius peronu “Temminck” Schlegel, 1865
3d ad, 22 February, Casq. Was alone.

December 5, 1995 Johannes Erritzoe 5

Charadrius mongolus Pallas, 1776
6 ad, 25 May, Aparri. 53 g (n-f). Stomach contents: small insects,
maggots, and snails.
2 imm, 03 May, Aparri. Legs blue-grey.

Charadrius leschenaultu Lesson, 1826
103 3, 31 March - 22 April, Aparri. 60-81 g (mean 68.8 g).
4 22, 03-21 April, Aparri. 70-81 g (mean 75.8 g).

FAMILY SCOLOPACIDAE

Tringa stagnatilis (Bechstein, 1803)
6 ad, 11 May, Chico River. 76 g; testes: left 7.5 x 4.2 mm, right 7.5
x 4.6 mm.

Tringa nebularia (Gunnerus, 1767)
6 ad, 09 April, Aparri. 154 g (n).

Tringa glareola Linné, 1758
d ad, 23 March, Isabela. 58 g (1).
2 imm, 12 March, Chico River. 77 g (vf); in molt. Stomach con-
tents: small snails.
2 22 imm, 12 March, Chico River. 77 and 65 g.
? imm, 31 March, Aparri. 58 g.

Actitis hypoleucos (Linné, 1758)
2 ad, 23 April, Chico River. 49 g (vf).

Arenaria interpres (Linné, 1758)

3 imm, 20 September, Aparri. 92 g (vf). Stomach contents: mag-
gots.

Gallinago megala Swinhoe, 1861
dS imm, 07 October, Pata. 134 g (f), bF 7.5 x 5.0 mm.

Calidris alba (Pallas, 1764)
3 ad, 09 April, Aparri. 58 g (f). Stomach contents: small snails.
3 ad, 21 May, Santa Teresita Beach. 55 g (f).
2 ad, 07 May, Aparri. 54 g (n).

Calidris ruficollis (Pallas, 1776)
3 66, 02-10 April, Aparri. 25, 26, and 29 g.

6 NEMOURIA No. 40

3d, 20 May, Chico River. 28 g.

6, 02 September, Aparri. 31 g.

3d, 05 September, Aparri. 30 g.

3d, 04 October, Aparri. 27 g.

6 22, 21-24 April, Chico River. 24-34 g (mean 29.3 g).
2 22,16 and 20 May, Chico River. 37 and 34 g.

2, 04 September, Aparri. 31 g.

Calidris subminuta (Middendorff, 1853)
2 66, 25 and 26 March, Chico River. 26 g and 27 g (n).
3d, 02 April, Chico River. 28 g (vf).
6, 21 April, Chico River. 30 g (f).
36, 24 April, Chico River. 25 g (f).
3d, April, Chico River. 30 g (vf).
2, 26 March, Chico River. 28 g (n).
?, March, Chico River. 28 g (n).
2, 02 April, Aparri. 30 g (f).
2, April, Chico River. 30 g.

Calidris acuminatus (Horsfield, 1821)
2 36 ad, 21 and 24 April, Chico River. 72 g and 73 g (f).
6 ad, 05 May, Chico River. 69 g (vf) (fat = 6 g).
2 imm, 20 April, Chico River. 62 g.
? imm, 06 May, Chico River. 55 g.
2 imm, 20 May, Chico River. 66 g (f).

Calidnis ferruginea (Pontoppidan, 1763)
3d ad, 04 May, Chico River. 56 g.
6 ad, 10 May, Chico River. 55 g.

Calhdris alpina (Linne, 1758)
2 ad, 01 March, Aparri. 61 g (n-f).

FAMILY STERNIDAE

Sterna albifrons Pallas, 1764
2 ad, 14 April, Aparri. 56 g (f).

FAMILY PSITTACIDAE

Prioniturus discurus (Vieillot, 1822)
6, 26 October, P. 86 g (1).

December 5, 1995 Johannes Erritzoe |

Tanygnathus lucionensis (Linné, 1766)
d,June, P. 168 g (1-n).
6 ad, June, P. 148 g (1).
6, 30 December, P. 168 g (I).

Tanygnathus sumatranus (Raffles, 1822)
6,06 June, P. 173 g (1).

Loriculus philippensis (P. L. S. Muller, 1776)
6 imm, 06 June, P. 31 g.
? imm, 07 December, P. 35 g (n).

FAMILY CUCULIDAE

Phaentcophaeus superciliosus Dumont, 1823

6 ad, 20 April, Cagayan. 100 g (n). Stomach contents: large inde-
terminable insects and locusts.

3 ad, 15 August, Bulacan. 112 g (n).

Phaenicophaeus curvirostris (Shaw, 1810)
6 ad, 16 May, Iwahig. 144 g (n).

Phaenicophaeus cumingi Fraser, 1839
2 ad, 15 March, P. 139 g.
2 ad, 15 May, P. 104 g (I).

Centropus bengalensis (Gmelin, 1788)

6,12 April, Pata. In molt; iris yellow (!). Stomach contents: large
indeterminable insects and locusts.

Centropus unirufus (Cabanis & Heine, 1863)

2 ad, 19 June, Santa Fe. 162 g (n); legs slate grey. Stomach con-
tents: passerine bird (!) and insects.

FAMILY APODIDAE
Collocalia esculenta (Linné, 1758)

3 363 imm, 28 August, Clayeria.
2, 17 April, Malamag.

8 NEMOURIA No. 40

FAMILY ALCEDINIDAE

Alcedo atthis (Linné, 1758)
3 ad, 14 April, Santa Fe. 29 g (f).
3d, 09 October, Chico River. 24 g (n); breast feathers in molt.
2, 29 March, Chico River. 27 g (f).

Halcyon coromanda (Latham, 1790)
6, 21 April, Palaui Island. 108 g. Stomach contents: snails.
2, 18 October, Santa Fe. 87 g (f); primaries on left wing very worn.

Halcyon smyrnensis (Linne, 1758)
6 ad, May, Bulacan. 68 g.

Halcyon chloris (Scopoli, 1786)

6 ad, 16 April, Laguna de Bai. 55 g (n). Stomach contents: cater-
pillar and other insects.
2 ad, 18 April, Santa Fe. 57 g.

FAMILY MEROPIDAE

Merops viridis Linné, 1758
2 ad, 04 April, Punta. 40 g.

Merops philippinus Linné, 1766
6 ad, 29 March, Chico River. 36 g (n).

FAMILY PITTIDAE

Pitta erythrogaster Temminck, 1823
6 ad, 16 June, Santa Fe. 49.7 g (n).
? ad, 23 November, Santa Fe. 47 g (n).
? imm, 04 December, Santa Fe. 47 g (n), bF 5.0 x 3.1 mm.
2 imm, 09 December, Santa Fe. 46 g (n).

Pitta sordida (P. L. S. Muller, 1776)
6 ad, 25 April, Santa Fe. 59 g (f).
3d ad, 16 May, Iwahig. 64 g.
2 22 ad, 17 and 18 April, Santa Fe. 45.4 g (I-n) and 48 g (n).

December 5, 1995 Johannes Erritzoe 9

FAMILY ALAUDIDAE

Murafra javanica Horsfield, 1821
3 ad, 24 May, Pata. 21 g (n).
2 36,13 and 19 October, Santa Fe. 23 g (n) and 19 g. Stomach
contents: rice.
3 $d, 06 and 11 December, Santa Fe. 20, 21, and 21 g. Stomach
contents: rice, insects and plant matter.

Alauda gulgula Franklin, 1831
6 ad, 25 March, Chico River. 26 g. Stomach contents: insects.

FAMILY HIRUNDINIDAE

HMirundo daurica Linneé, 1771
6 ad, 13 May, Santa Ana. 31 g (f).

FAMILY PYCNONOTIDAE

Pycnonotus urostictus (Salvadori, 1870)
6 ad, 20 April, Pefia Blanca. 26 g (n); eye-ring lemon-yellow.

Pycnonotus goiavier (Scopoli, 1786)
2 ad, 31 October, Pefia Blanca. 31 g (n).

FAMILY ORIOLIDAE
Oriolus chinensis Linné, 1766
° ad, 12 February, Bulacan. 106 g (vf). Stomach contents: larvae,
fruit, berries.
FAMILY PARIDAE
Parus elegans Lesson, 1831
3S ad, 18 April, Laguna de Bai. 15 g (n).
3 ad, 11 May, Santa Fe. 14 g (n).
FAMILY SITTIDAE

Sitta frontalis Swainson, 1820
3 ad, 11 May, Santa Fe. 17 g (n).

10 NEMOURIA No. 40

FAMILY RHABDORNITHIDAE

Rhabdornis mystacalis (Temminck, 1825)
6 ad, 19 April, Laguna de Bai. 28 g (n).

FAMILY TURDIDAE

Copsychus luzoniensis (Kittlitz, 1832)
6 ad, 04 March, Mt. Maquiling. 25 g.

Copsychus niger (Sharpe, 1877)
6 ad, 16 May, Iwahig. 30 g (n).

Saxicola caprata (Linné, 1766)
6 ad, 01 February, Luzon. 15 g.
3d ad, 04 May, Maddela. 14 g (n).
6 ad, 26 May, Pata. 14 g.
6 imm, 06 September, Pata. Stomach contents: berries.
2 ad, 07 March, North Luzon. 16 g.

Monticola sohtarius (Linné, 1758)
6, 24 January, Santa Fe. 41 g (1); right side of breast in molt.
2 ad, October, Chico River. 52 g (n).

Turdus chrysolaus Temminck, 1831
6, February, Santa Fe. 71.5 g.

FAMILY SYLVITDAE

Phylloscopus borealis (Blasius, 1858)
? imm, 18 October, Santa Fe. 8 g (I).

Acrocephalus stentoreus (Ehrenberg, 1833)
3, 29 March, Laguna de Bai. 23 g.
2 22,13 and 20 March, Laguna de Bai. 21 g (n) and 19.8 g (n).

Acrocephalus orientalis (Temminck & Schlegel, 1847)
6, 20 March, Laguna de Bai. 24 g (n).
6, 03 April, Laguna de Bai. 27 g.
6, 19 October, Santa Fe. 25 g (n).
6, 06 November, Santa Fe. 26 g (f).
2 22 imm, 13 and 19 October, Santa Fe. 21 g (n) and 26 g.

December 5, 1995 Johannes Erritzoe 11

Locustella ochotensis (Middendorff, 1853)
6 imm, 06 November, Santa Fe. 17 g (n).

Locustella lanceolata (Temminck, 1840)
2 366, 03 and 07 December, Santa Fe. 12 g (f) and 10 g (f).
2 22,08 and 18 October, Santa Fe. 11 g and 10 g (f).

Megalurus palustris Horsfield, 1820
6 ad, 24 February, Jones. 47 g. Stomach contents: plant matter and
insects.
3d ad, 03 April, Magot River. 58 g.
2, 03 April, Laguna de Bai. 35 g (n).
? imm, 02 October, Laguna de Bai. 35 g (n), bF 7.7 x 3.4 mm.

Cisticola juncidis (Rafinesque, 1810)
6 ad, 25 May, Gonzaga. 10 g (n).

FAMILY MUSCICAPIDAE

Cyornis rufigastra (Raffles, 1822)
6,17 February, Nassiping. 17 g.
6, 29 December, Munting. 20 g.
2, 28 December, Munting. 17 g; one albino feather on back.

Rhipidura cyaniceps (Cassin, 1855)
6, 03 January, Laguna de Bai. 16 g (n).
3d, 01 December, Los Bafios. 18 g (n).
2, 18 April, Laguna Kaliraya. 14 g.
2 ad, 19 May, Santa Fe. 13 g (n); legs slate-grey.

Rhipidura javanica (Sparrman, 1788)
3d, 27 December, Bataan. 18 g (n).
2, 26 December, Bataan. 18 g (n).

Aypothymis azurea (Boddaert, 1783)
3 ad, 01 April, Pata. Mouth and tongue yellow.
2 ad, 21 April, Palaui Island. 12 g (n).

FAMILY MOTACILLIDAE

Motacilla flava Linné, 1758
3, 03 February, Magot River. 15 g.
3, 18 May, Santa Fe. 31 g (vf).

12 NEMOURIA

Anthus novaeseelandiae (Gmelin, 1789)
d ad, 06 March, Pata. 28 g.
d ad, 24 May, Pata. 25 g.
6 ad, 25 May, San Mariano. 22 g.

2 3664, 26 May, Chico River. 24 g and 27 g (n).

6 ad, 10 September, Isabela. 27 g.
2,08 March, North Luzon. 27 g.

2 ad, 25 May, Aparri. 26 g (n).

2 imm, 07 September, Pata. 27.5 g.

FAMILY ARTAMIDAE

Artamus leucorhynchus (Linné, 1771)
6 ad, 24 April, Conner. 42 g (n).

FAMILY LANIIDAE
Lantus schach Linné, 1758
6 imm, 02 October, Luzon. 37 g.
2 ad, 26 May, Pata. 36 g (n).
FAMILY STURNIDAE

Sarcops calvus (Linné, 1766)
2 ad, 18 February, P. 82 g (I).

Acridotheres cristatellus (Linné, 1766)
3 ad, 06 May, Santa Fe. 122 g (n-f).

FAMILY NECTARINITIDAE

Nectarinia jugularis (Linné, 1766)
6 ad, 06 March, Laguna de Bai. 9 g.
3d ad, 01 June, Santa Fe. 9 g (n).
2 ad, 07 March, Laguna Kaliraya. 8 g (n).

FAMILY DICAEIDAE
Dicaeum australe (Hermann, 1783)

6 ad, 19 April, Laguna Kaliraya. 9 g (n).
2 ad, 12 March, Laguna Kaliraya. 9 g (n).

No. 40

December 5, 1995 Johannes Erritzoe 13

FAMILY ZOSTEROPIDAE

Zosterops montanus Bonaparte, 1851
66, 02 and 03 January, Laguna de Bai. 11 g (f) and 12 g.

FAMILY PLOCEIDAE

Passer montanus (Linné, 1758)
6 ad, 10 January, Cagayan. 19.5 g (J).

FAMILY ESTRILDIDAE

Lonchura punctulata (Linné, 1758)
? ad, 03 April, Laguna de Bai. 13 g.

DISCUSSION

Not surprisingly, the Rallidae with their hidden habits offered the
most interesting records. Among 16 skins and one skeleton, seven species
are uncommon to rare (Dickinson, et al., 1991):

Uncommon resident: Rallina eurizonoides (n = 1); Porzana fusca (n = 3).

Uncommon, probably winter visitor: Porzana pusilla (n = 2).

Rare and extremely secretive resident: Porzana tabuensis (n = 1).

In all cases, except the Baillon’s Crake (Porzana pusilla) found dead and
decomposed, the local people had caught the birds in snares for con-
sumption.

The large Gallicrex cinerea male was still in its non-breeding plumage
without any trace of molt despite the large gonads and breeding reported
to start only one month later (Ripley, 1977).

The Greater Painted-snipe, Rostratula benghalensis, is an uncommon,
local resident (Dickinson, et al., 1991). The five specimens collected one-
at-a-time on Luzon suggest that the population on this island is over-
looked and more common than previously believed.

The Marsh Sandpiper, Jringa stagnatilis, is a passage migrant or win-
ter visitor (Dickinson, et al., 1991). The late date, 11 May, and the low
body mass (76 g; range 55.0 - 120.0 g, Dunning, 1993) indicate that it
may have been a summer visitor. The record of a Common Greenshank,
Tringa nebularia, is also noteworthy because it is reported as an uncom-
mon passage migrant and winter visitor (Dickinson, et al., 1991).

The Sanderling, Calidris alba, is classified as rare and only seen on
Luzon between November and May (Dickinson, et al., 1991). All three
Sanderlings were from April-May with small- (April) to medium-sized

14 NEMOURIA No. 40

gonads. The record from 21 May might possibly have been a non-breed-
ing summer visitor because many non-breeders remain south all year
(Hayman, et al., 1986). The most remarkable shorebird is the Dunlin,
Cahdris alpina, found dead on the sand beach at Aparri, 01 March 1988,
the first record of this species for the Philippines (Erritzoe, 1994).

The single record of a Little Tern, Sterna albifrons, from Aparri is also
interesting, because this bird is an uncommon winter visitor and a rare
resident only breeding on Mindoro (Dickinson, et al., 1991).

The female Ruddy Kingfisher, Halcyon coromanda, collected on 18
October was abnormal in having all primaries new except three in the
right wing, primaries 7-8 with normal wear, and primary 10 about 2 mm
shorter. Compared with this, the primaries on the left wing were all worn
and shortened, some about one-quarter of the normal length, and the
outer webs on most primaries had notches. It looked most similar to the
feather-eating commonly seen, for instance, in captive parrots, but I have
never seen this reported in kingfishers. The bird was otherwise in quite
normal condition.

I compared a White-breasted Kingfisher, Halcyon smyrnensis, from
Selangor, Malaysia, and another skin from Bangkok, Thailand [both of
the subspecies fusca (Boddaert, 1783)], with the specimen from the
Philippines [subspecies gularis (Kuhl, 1820)], and found a surprising
detail. On an area of the lower breast, where the nominate and H. s. fusca
are white and the Philippine bird is typically red-brown, a faint tinge of
white was visible. On the same area, the bases of the feathers were black.
On the other hand, on the upper breast and lower throat, the bases of the
red-brown feathers were white, but here the feathers had no white tinge.
This color pattern has not been mentioned in other works (Forshaw &
Cooper, 1985; Fry, et al., 1992).

The two males of Zosterops montanus were determined to belong to

this species in spite of the lowland locality because of black lores which
in Z. meyent Bonaparte, 1850, are paler (Mees, 1957: 170).

ACKNOWLEDGMENTS

This paper is dedicated to my dear brother, Thorkild Erritzoe, who
for many years was the Danish Consul General for the Philippines. I
acknowledge Dr. Kenneth C. Parkes and Prof. Anders Pape Moller for
making valuable comments on an earlier draft of this manuscript.

December 5, 1995 Johannes Erritzoe 15

LITERATURE CITED
Amadon, D. & S.T. Jewett. 1946. Notes on Philippine birds. Auk, 63: 541-559.

Clench, M. H. & R. C. Leberman. 1978. Weights of 151 species of Pennsylvania
birds analyzed by month, age, and sex. Bulletin of the Carnegie Museum of
Natural History, 5, 87 pp.

Dickinson, E. C., R. Kennedy, & K. C. Parkes. 1991. The Birds of the
Philippines: an annotated check-list. British Ornithologists’ Union Check-
list 12, 507 pp.

Dunning, J. B., Jr. 1984. Body weights of 686 species of North American birds.
Western Bird Banding Association, Monograph 1, 38 pp.

— 1993. Avian Body Masses. CRC Press, Boca Raton, 371 pp.

du Pont, J. E. 1971. Philippine Birds. Delaware Museum of Natural History,
Greenville, 480 pp.

Erritzoe, J. 1985. Geschlechts- and Altersbestimmung bei Végeln. Der
Praparator, 31: 81-93.

— 1994. First record of the Dunlin from the Philippines. Bulletin of the British
Ornithologists’ Club, 114: 128-129.

Forshaw, J. M. & W. TIT. Cooper. 1985. Kingfishers and Related Birds, vol. 1.
Lansdowne, Melbourne, 266 pp.

Fry, C. H., K. Fry, & A. Harris. 1992. Kingfishers, Bee-Eaters & Rollers.
Princeton University Press, Princeton, New Jersey, 324 pp.

Hayman, P., J. Marchant, & T. Prater. 1986. Shorebirds, an identification guide to
the waders of the world. Houghton Mifflin Company, Boston, 412 pp.

Mees, G. F. 1957. A systematic review of the Indo-Australian Zosteropidae. Part
1. Zoologische Verhandelingen (Leiden), 35: 1-204.

Parkes, K. C. 1988. Three new subspecies of Philippine birds. Nemouria, 30: 1-
8.

Ripley, S. D. 1977. Rails of the World. David R. Godine, Boston, 406 pp.

— &D.S. Rabor. 1958. Notes on a collection of birds from Mindoro Island,
Philippines. Peabody Museum of Natural History, Yale University, Bulletin,
13: 1-83.

— & —— 1962. New birds from Palawan and Culion Islands, Philippines.
Postilla, 73: 1-16.

Smith, J. N. M., P. Arcese, & D. Schluter. 1986. Song sparrows grow and shrink
with age. Auk, 103: 210-212.

Thomas, B. T. 1982. Weights of some Venezuelan birds. Bulletin of the British
Ornithologists’ Club, 102: 48-52.

Founding Editor Managing Editor
JOHN E. DU PONT PAULA M. MIKKELSEN

Consulting Editors

WILLIAM K. EMERSON, American Museum of Natural History, New
York, NY.

M. G. HARASEWYCH, National Museum of Natural History, Smithsonian
Institution, Washington, DC.

KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh,
PA.

RICHARD W. THORINGTON, JR., National Museum of Natural History,
Smithsonian Institution, Washington, DC.

Published by the

DELAWARE MUSEUM OF NATURAL HISTORY
4840 Kennett Pike, Wilmington,
Delaware 19807, U.S.A.

67
36
NA NEMOURIA

Occasional Papers of the Delaware Museum of Natural History
NUMBER 41 NOVEMBER 30, 1997

ANNOTATED CATALOG OF TYPE SPECIMENS IN
THE MALACOLOGICAL COLLECTION OF THE
DELAWARE MUSEUM OF NATURAL HISTORY

PART II. ADDITIONS AND CORRECTIONS
TO PART I (PROSOBRANCHS, HETEROSTROPHA,
AND OPISTHOBRANCHIA), PLUS BIVALVIA,
SCAPHOPODA, AND POLYPLACOPHORA

Paula M. Mikkelsen! and Alison Bradford’

ABSTRACT. An annotated catalog of name-bearing type specimens in the
mollusk collection at Delaware Museum of Natural History is continued. This
contribution is Part II of the series, following Part I (marine prosobranchs and
opisthobranchs) by Bieler & Bradford (1991, Nemouria, No. 36, 1-48). This
part lists all remaining type specimens, excluding Pulmonata. A total of 143
type lots of 122 species-group taxa 1s recognized, and taxonomic and author
indices to all taxa in Parts I and II are provided.

INTRODUCTION

In accordance with Recommendation 72G(4) of the International Code
of Zoological Nomenclature (ICZN, 1985), this publication continues the
annotated catalog of name-bearing type specimens [holotypes, paratypes,
lectotypes, and syntypes (= cotypes)] in the mollusk collection of the Delaware
Museum of Natural History (DMNH). Part I (Bieler & Bradford, 1991)
included 189 lots of marine “Prosobranchia” (now recognized as an artificial
taxon; Bieler, 1992) and Opisthobranchia. This part includes corrections and
additions to Part I plus entries for 143 lots of non-pulmonate mollusks not
previously listed. Following completion of ¢ project to isolate all name-bearing

' Department of Invertebrates, American Museum of Natural History, Central Park
West at 79th Street, New York, New York 10024-5192.

* Department of Malacology, Delaware Museum of Natural History, P. O. Box 3937,
Wilmington, Delaware 19807-0937.

2 NEMOURIA No. 41

types from the main collection storage cabinets into a segregated type
collection, it became obvious that, contrary to an earlier statement (Bieler &
Bradford, 1991: 1, 4), most of the DMNH types were not marine
prosobranchs and opisthobranchs, but rather pulmonates. At present,
presumed types of Pulmonata comprise 788 lots, or 70% of the total type
collection; these have not yet been verified and await future study.

Since the previous catalog, the malacological collection at DMNH has
surpassed 208,000 cataloged lots, maintaining its status within the top ten
largest mollusk collections in the U. S. A. Moreover, DMNH undoubtedly
ranks among the most completely computerized of major molluscan collections
(now surpassing 72%), and still claims a remarkable capacity for collection
growth in terms of available collection storage space. Contributing toward this
catalog since 1991 were significant private collections containing type material
donated by Dr. Lawrence F. Kienle (Medford, New Jersey; Cypraeidae),
Helene Eker (Phoenix, Arizona; Mitridae and Costellariidae), and Eugenia I.
Wright (Phoenix, Arizona; Muricidae). Type specimens of newly named
species were also received from B. J. Piech (DMNH), D. Tippett (National
Museum of Natural History), and J. K. Tucker (Illinois Natural History
Survey, Wood River). In addition, one archival resource of particular note was
acquired as part of the Eker donation. This is a set of approximately 2,000 10
x 16.5 cm alphabetically arranged index cards maintained as a research tool by
Walter O. Cernohorsky, listing most of the described species of Mitridae and
Costellariidae. Type material, original description information, geographical
distribution, and Cernohorsky's own research notes are included on each card.
DMNH also acquired approximately 1,300 color photographic transparencies,
photographed by Cernohorsky, of type specimens of Miutridae and
Costellariidae in the world's museum collections.

Following a short list of additions and corrections to Part I (Bieler &
Bradford, 1991), this list is arranged in the same general format as that of
Bieler & Bradford (1991). Alphabetical order of taxa is used within major
molluscan groups: Polyplacophora, prosobranchs (additions to Part I + non-
marine taxa), Heterostropha, Opisthobranchia (additions to Part I), Bivalvia,
and Scaphopoda. Each entry includes the original combination, original status
(exactly as expressed), full citation, number and kind of types, DMNH catalog
number, locality data, source of DMNH material (with source lot catalog
number if known), specimen dimension(s), present (and original) family
placement, and remarks (justification of type status if not cited specifically by
original description, condition of specimens, etc.). The locality information
given is as cited on the collection label, not modified according to the original
description or other source, unless so noted. Like Part I, this is not a critical
catalog in that no attempt has been made to fully research each taxon with
regard to present systematic status, other type material, etc.; however, the valid

November 30, 1997 Mikkelsen & Bradford s)

name, if different from the orginal through Known synonymy or as used in the
DMNH or ANSP collections, is given following the family status.

As in Part I, we excluded specimens that were previously recognized as
types simply because they were part of the original material of the author if the
original description specifically restricted the type repositories. Much of the
type material in this catalog originated as "split lots" from the Academy of
Natural Sciences of Philadelphia (ANSP); the ANSP collection and its catalog
records were consulted as additional evidence either for or against type status
of the DMNH lots. Many of the ANSP source lots are now divided into "type
+ paratypes"; although these cases are noted as such in the text below, none
of these notations (nor any other statement herein) are intended to be, nor
should they be interpreted as, lectotype designations.

The specimen dimensions were determined to the nearest 0.1 mm either
with calipers or a calibrated eye-piece in a dissecting stereomicroscope.
Dimensions are presented slightly differently for the different shell forms
reflected by the various classes. Polyplacophorans are given as Lx W, where
L = anteroposterior length, and W = lateral width; gastropods, as H x W,
where H = anteroposterior height, and W = lateral width; bivalves, as Lx H,
Where L = anteroposterior length, and H = dorsoventral height; and
scaphopods, as Lx W, where L = length of the long axis, and W = maximum
width or diameter of the tube. Generally, a range is provided if the lot contains
more than four specimens. Discrepancies between published and actual
dimensions are also noted.

The entry format of this paper differs from Part I (Bieler & Bradford,
1991) in only two minor ways: (1) Each original citation includes all pages and
figures, not just those figuring DMNH type material. Any DMNH specimens
that are believed to be figured types are listed as such. (2) Family placement
is expressed differently for ease of readability. Part I listed the original family,
followed by the current family (in brackets) according to the classification of
Ponder & Warén (1988). In this Part, the current family is presented first,
followed by the original placement, if different, in brackets. Current family
assignment was guided by Ponder & Warén (1988, prosobranchs only),
Vaught (1989), and Turgeon, er al. (in press).

This list includes 122 taxa, 143 lots, 880 specimens, and 8 holotypes.
Polyplacophora comprise 3 lots (3 taxa), additional prosobranchs 79 lots (62
taxa), Heterostropha 1 lot (1 taxon), additional Opisthobranchia 1 lot (1
taxon), Bivalvia 58 lots (54 taxa), and Scaphopoda 1 lot (1 taxon). No
cephalopod types presently reside at DMNH. Parts I and II of this catalog (all
species except Pulmonata) comprise a total of 277 taxa, 334 lots, 1,339
specimens, and 46 holotypes. Four taxa are covered by both parts of this
catalog.

The type entries are followed by two indices, taxonomic (new in this Part)

= NEMOURIA No. 41

and author (total = 105), which include names from both published parts of
this catalog. The Literature Cited includes all original citations plus any other
cited references.

Abbreviations used throughout the text are: AIM, Auckland Institute &
Museum, New Zealand; AMNH, American Museum of Natural History, New
York; ANSP, Academy of Natural Sciences of Philadelphia, Pennsylvania;
DMNH, Delaware Museum of Natural History, Wilmington; D[T]MNH,
Dallas Museum of Natural History, Texas; FLMNH, Florida Museum of
Natural History, Gainesville; FMNH, Field Museum of Natural History,
Chicago; MCZ, Museum of Comparative Zoology, Harvard University,
Cambridge, Massachusetts; UF, University of Florida [= FLMNH]; USNM,
National Museum of Natural History [United States National Museum],
Smithsonian Institution, Washington, D. C.; WAM, Western Australian
Museum, Perth, Western Australia.

TYPE CATALOG

ADDITIONS AND CORRECTIONS
TO PART I (BIELER & BRADFORD, 1991)

Page 6, aureola, Pyrene, Howard, 1963. Add: (non Duclos in Chenu, 1846).
Emended (one month after original description) by Howard (1963) to
aureomexicana.

Page 7, axelrodi, Conus, Walls, 1978. Next-to-last sentence should read: "Also
described by Walls (1979a: 183 ff.) with reference to earlier publication
(Walls, 1978)."

Page 8, boholensis, Conus, Petuch, 1979. DMNH 126400 (holotype, 38.4
mm) and 126401 (2 paratypes, 26 and 17 mm) were examined and
measured by Coomans, et al. (1982: 32) and determined to be
synonymous with C. borneensis Adams & Reeve, 1848.

Page 8, boschi, Cymatium, Abbott & Lewis, 1970. Add: DMNH holotype
listed by Henning & Hemmen (1993: 74).

Page 10, clovert, Conus, Walls, 1978. Next-to-last sentence should read "Also
described by Walls (1979a: 338 ff.) with reference to earlier publication
(Walls, 1978)."

Page 14, fayae, Anachis (?Costanachis), Keen, 1971. Read (?Costoanachis).

Page 15, floridanus, Microdochus, Rehder, 1943. Exact number of paratypes
is 78.

Page 16, guesti, Pterynotus, Harasewych & Jensen, 1979. Add: DMNH
holotype listed by Houart (1994).

Page 19, Rerstitchi, Conus, Walls, 1978. Next-to-last sentence should read

November 30, 1997 Mikkelsen & Bradford 5

"Also described by Walls (1979a: 615 ff.) with reference to earlier
publication (Walls, 1978)..."

Page 21, ightbournt, Prerynotus, Harasewych & Jensen, 1979. Add: DMNH
holotype listed by Houart (1994).

Page 26, philippinensis, Axymene, Petuch, 1979. Add: DMNH holotype
listed by Houart (1994, as A. philippensts [sic]), and by Bouchet & Warén
(1986) who synonymized the species with Eostpho dentatus (Schepman,
1911) of Buccinidae.

Page 26, radwint, Prerynotus, Harasewych & Jensen, 1979. DMNH holotype
listed by Houart (1994).

Page 27, sandwichensis, Conus suturalis, Walls, 1978. Read suturatus.

Page 30, vernicosum, Mangilia (Calliotectum), Dail, 1889. Read Dall, 1890.
Add: Placed in Volutidae by subsequent authors, e. g., Weaver & duPont
(1970), Emerson & Sage (1986),-Poppe & Goto (1992).

Page 43, Dall, W. H., 1889. Read 1890.

POLYPLACOPHORA

andersoni, Acanthochitona [as new species]

Watters, 1981: 173-176, pls. 2e-g, 41.

1 figured paratype: DMNH 095383; 35 ft [published as 10.6 m], by
scuba, west of Picquet Rocks, Bimini Island, Bahamas. Coll. G.
Watters; ex G. Watters. Dimensions (L x W, greatest dimensions
of strongly curved, dried animal): 4.8 x 2.8 mm.

Remarks: "On Strombus gigas" and "April 17, 1973" were in the original
description of this type lot, but are not on the specimen label.
Three illustrations of this paratype appear in pls. 2e-g, which
although they look like dissociated plates, were presumably drawn
from this still-intact specimen.

Acanthochitonidae [originally Cryptoplacidae].

lineata, Acanthochitona [as sp. nov.]
Lyons, 1988: 90-92, figs. 42-51.
1 paratype: DMNH 095381; Water Island, Virgin Islands. Coll. Weber,
July 1959; ex G. Watters. Dirnensions (L x W): 8.5 x 4.3 mm
[published as 9.7 mm}.
Acanthochitonidae.

portobelensis, Callistochiton [as new species]
Ferreira, 1976: 46-49, figs. 1-6.
1 paratype: DMNH 102101; 1-5 ft, under side af loose rock, about 5 mi

6 NEMOURIA No. 41

[published as 5 km] west of Portobelo, Panama. Coll. A. J.
Ferreira, 15 February 1975; ex A. J. Ferreira. Dimensions (L x W):
5,5 x 2.0) mim.

Remarks: The original description mentioned paratypes in the author's
collection, the source of this material.

Ischnochitonidae [originally Callistoplacidae].

PROSOBRANCHS
(ADDITIONS TO PART I; BIELER & BRADFORD, 1991)

acanthus, Mirachelus [as new species]

Quinn, 1991b: 168, figs. 4-6.

Holotype (figured): DMNH 187589; dredged, off Castle Roads,
Bermuda. Coll. A. Guest (lot #582); ex J. E. Quinn. Dimensions
(H x W): 3.1 x 2.5 mm [published as 3.45 x 2.95 mm].

3 paratypes: DMNH 187590; dredged, off Castle Roads, Bermuda.
Coll. A. Guest (lot #582) via J. E. Quinn. Dimensions (H x W):
(1) 2.9 x 2.4 mm, (2) 2.8 x 2.4 mm, G) 2.7-x 2.3 mm:

Remarks: Although the caption to the published scanning electron
micrographs (Quinn, 1991b: figs. 4-6) indicated "Paratype,
DMNH 187590," it is the holotype (DMNH 187589) that has
been coated for SEM. The published depth of 100 m is not
indicated on the DMNH labels.

Trochidae [originally Trochidae: Eucyclinae: Chilodontini].

alcalde1, Chondropoma (Chondropoma) [as n. sp.]

Jaume & Sanchez de Fuentes, 1943a: 17-18, pl. 2, fig. 1.

4 paratypes: DMNH 135179; Mogote Las Damas, Central Dolores,
Pedro Betancourt, Matanzas, Cuba. Ex O. Alcalde (Jackson
catalog) via R. W. Jackson (#6628). Dimensions (H x W): (1)
14.8 x 9.1 mm, (2) 14.5 x 8.7 mm, (3) 14.2 x 9.1 mm, (4) 13.1%
8.8 mm.

Remarks: The original description mentioned paratypes in private
collections, 7. e., "en las colecciones de los autores y en la del Sr.
Oscar Alcalde Ledon." The Jackson collection catalog in the
DMNH archives indicates the source of these specimens as
Alcalde.

Annulariidae [none specified originally].

alcaldei, Chondrothynum (Plicathyrium) [as nueva especie]
Jaume & Sanchez de Fuentes, 1943c: 24-25, pl. 4, fig. 1.

November 30, 1997 Mikkelsen & Bradford 7

1 paratype: DMNH 135184; Palmarito de Gonzales, Cienfuegos, Las
Villas, Cuba. Ex O. Alcalde via R. W. Jackson [#6613, as C. (P.)
alcaldet alcaldei]. Dimensions (H x W): 14.6 x 10.2 mm.

Remarks: The original description mentioned paratypes in "las
colecciones de los autores y en la del Sr. O. Alcalde."

Annulariidae [none specified originally].

alcoviensis, Somatogyrus [as new species]

Krieger, 1972: 120-125, figs. 1-4.

12 paratypes (including figured): DMNH 041741; Cedar Shoals, Yellow
River, Newton County, Georgia. Coll. K. A. Krieger, 1968-1971;
ex K. A. Kneger. Dimensions (range): H = 2.2-4.3 mm, W = 2.1-
3.6 mm.

12 paratypes: DMNH 041742; Newton Factory Shoals, Alcovy River,
Newton County, Georgia. Coll. K. A. Krieger, 1970-1971; ex K.
A. Krieger. Dimensions (range): H = 1.5-3.0 mm, W = 1.6-2.9
mm.

Remarks: Krieger's (1972) fig. 1 showed three views of two paratypes
from DMNH 041741; they measure 4.3 x 3.6 mm [the largest
specimen] and 4.1 x 3.6 mm (both larger than the stated
dimensions from fig. 1 of 3.0 x 2.9 mm). Most of the paratypes
have opercula plus dried soft parts inside the shell.

Hydrobiidae.

backae, Scabricola (Scabricola) [as sp. n.]

Cernohorsky, 1973: 133-135, figs. 1-5.

1 paratype: DMNH 197613; Flinders Bay, Augusta, southwestern
Australia. Coll. Mrs. F. Back, 13 September 1972; ex W. O.
Cernohorsky via H. Eker. Dimensions (H x W): 22.0 x 9.1 mm.

1 paratype: DMNH 197616; Flinders Bay, Augusta, southwestern
Australia. Ex Auckland Institute & Museum via H. Eker.
Dimensions (H x W): 23.3 x 9.0 mm. Remarks: The original label
indicates two paratypes, but this lot contains only one, with "E"
written in pencil on inside of outer lip.

Remarks: The Cernohorsky index card in DMNH does not list the Eker
collection, but the original description mentioned two paratypes in
the Eker collection (including one at 23.1 x 9.0 mm = DMNH
197616), and one in Cernohorsky's collection (given as 22.0 x 9.0
mm = DMNH 197613), all from Flinders Bay.

Mitridae [originally Mitridae: Imbricariinae]. This is now a synonym of
Scabnicola sowerbyi melvilli (Sowerby, 1882) (Cernohorsky, 1991).

8 NEMOURIA No. 41

barretti, Odostomia [as new species]

Morrison, 1965: 220, fig. 4.

2 paratypes: DMNH 108366; 4 ft, northeast corner of Heron Bay,
Mississippi. Coll. J. P. E. Morrison; ex USNM 635631 (wd?).
Dimensions (H x W): (1) 2.0 x 1.0 mm (worn shell, apertural lip
broken), (2) 1.7 x 0.9 mm (fresh shell).

Remarks: These also serve as types for the replacement name Hydrobia
booneae Morrison (1973: 28) [new name for Odostomia barretn
Morrison, 1965, non O. bareti L. Morlet, 1885].

Hydrobiidae [originally Pyramidellidae].

bermudense, Calliostoma [as new species]

Quinn, 1992: 86, figs. 23-24.

Holotype (figured): DMNH 187591; dredged, 45 fms, off Castle Roads,
Bermuda. Coll. A. Guest & J. R. H. Lightbourn, 1975-1976; ex J.
E. Quinn. Dimensions (H x W): 11.6 x 8.5 mm [published as 11.9
x 8.9 mm].

9 paratypes: DMNH 187592; dredged, 45 fms, off Castle Roads,
Bermuda. Coll. A. Guest & J. R. H. Lightbourn, 1975-1976; ex J.
E. Quinn. Dimensions (range): H = 5.7-8.8 mm, W = 4.8-7.0
mm.

Trochidae [originally Trochidae: Calliostomatinae].

brito1, Eutudora limbifera [as nueva subespecie]

Jaume & Sanchez de Fuentes, 1943b: 19-20, pl. 3, fig. 1.

6 paratypes: DMNH 135217; El Hatillo, San Miguel de los Banos,
Matanzas, Cuba. Ex O. Alcalde (Jackson catalog) via R. W.
Jackson (#6641). Dimensions (range): H = 10.1-12.7 mm, W =
9.7-9.8 mm.

Remarks: The original description mentioned paratypes in private
collections, 7. e., "en las colecciones de los autores y del Sr. Oscar
Alcalde y Ledon." The Jackson collection catalog in the DMNH
archives indicates the source of these specimens as Alcalde. All
specimens have opercula in place, possibly with dried soft parts.

Annularidae.

caribaea, Coralliophila [as new species]
Abbott, 1958: 66-67, text-fig. 3 (1-2), pl. 1g-h.
3 paratypes: DMNH 024282; Vera Cruz, Mexico. Coll. Heilprin & F.
C. Baker, 1890; ex ANSP 61054 (vd). Dimensions (H x W): (1)
19.4 x 15.1 mm, (2) 1879'x 11-9'mm, G) 18:2 x 123mm:

November 30, 1997 Mikkelsen & Bradford 9

Remarks: All live-collected, with dried soft parts in shells.
Muricidae [originally Magilidae].

carinatus, Lamellitrochus [as new species]

Quinn, 1991a: 84-85, figs. 7-12, 34-35.

Holotype: DMNH 179393; southwest of Egmont Key, Florida, 200-230
fms. Coll. J. Moore, May 1962; ex D. Steger. Dimensions (H x
W): 2.3 x 2.5 mm [published as 2.50 x 2.70 mm]. Remarks: Split
from DMNH 104803 [Lamelhitrochus lamellosus (Verrill & Smith,
1880)].

31 paratypes: DMNH 179392; 80 fms, rubble, 100 mi southwest of
Egmont Key, Florida. Ex D. Steger. Dimensions (range): H = 1.1-
2.5 mm, W = 1.2-2.6 mm. Remarks: Split from DMNH 104814
[L. lamellosus (Verrill & Smith, 1880)].

22 paratypes (including figured): DMNH 186768; southwest of Egmont
Key, Florida. Coll. J. Moore, May 1962; ex D. Steger. Dimensions
(range): H = 1.3-2.5 mm, W = 1.7-2.6 mm. Remarks: Includes
two specimens gold-coated for scanning electron microscopy (figs.
7-9). Split from DMNH 179393; this is now the holotypic lot
(above) of this species, but which according to DMNH cataloging
records initially comprised 16 specimens. The discrepancy in the
number of specimens (15 versus 22; initially miscounted?
additional specimens added from original lot, DMNH 104803?) is
unresolved.

Trochidae [orginally Trochidae: Solariellinae].

casta, Notocypraea [as nov. spec. ]

Schilder & Summers, 1963: 66.

1 paratype (labeled "e"): DMNH 196180; Port MacDonnell, South
Australia. Coll. by Ian Carrison, 1960; ex P. Trenberth via L. F.
Kienle. Dimensions (H x W): 21.0 x 12.8 mm [original label as
WS) poveayp

Remarks: This specimen is pure white. The original description
mentioned paratypes (although un-numbered or -lettered) "in colls.
Schilder, Summers, Trenberth, et. [szc] al."

Cypraeidae [none specified originally]. This is a junior synonym of C.
comptonn Gray, 1847 (Burgess, 1985).

chocolatum, Incidostoma [as n. sp.]
Morrison, 1955: 158-159, figs. 10-12.
5 paratypes: DMNH 135098; Papallagta, Ecuador. Ex R. W. Jackson
(#6382). Dimensions (range): H = 12.2-14.3 mm, W = 17.7-22.6

10 NEMOURIA No. 41

mm.

Remarks: The original description noted "additional paratypes from the
original lot are in the collection of Mr. Jackson." The Jackson
collection catalog indicates "det. Dr. Morrison."

Poteriidae [originally Cyclophoridae: Neocyclotinae]. Curated in the
genus Cyrtotoma in the DMNH collection.

clover1, Cancilla (Ziba) [as sp. n.]

Cernohorsky, 1971: 133-135, figs. 4-8.

1 figured paratype: DMNH 197615; 70 fms, 75 mi southwest of
Kaushiung, Formosa Strait, Taiwan. Coll. 1969; ex W. O.
Cernohorsky via H. Eker. Dimensions (H x W): 17.0 x 5.5 mm.

Remarks: The DMNH specimen is "paratype no. 2" described as "in the
author's coll." and figured in the original description (fig. 6). Its
intact protoconch was also illustrated in a line drawing (fig. 8). The
paratype is listed as in Cernohorsky's collection on Cernohorsky's
index card at DMNH, but was not included in the list of type
material given by Cernohorsky (1991).

Mitridae [originally Mitridae: Imbricariinae].

colombiensts, Aroapyrgus [as n. sp.]

Malek & Little, 1971: 20-25, fig. 1.

Holotype: DMNH 041542; tributaries of Rio Pichende, near Pichende,
Municipio de Cali, Departamento de Valle de Cauca, Colombia.
Coll. E. A. Malek, 1968; ex E. A. Malek. Dimensions (H x W): 3.2
x 1.4 mm.

91 paratypes: DMNH 041543; tributaries of Rio Pichende, near
Pichende, Municipio de Cali, Departamento de Valle de Cauca,
Colombia. Coll. E. A. Malek, 1968; ex E. A. Malek. Dimensions
(range): H = 2.0-3.1 mm, W = 1.0-1.6 mm.

Hydrobiidae.

crosnieri1, Cancilla scrobiculata [as new subspecies]

Cernohorsky, 1970a: 95, 98-99, figs. 1-2.

1 paratype ("No. 3"): DMNH 197604; 150-200 m, 06°05'S, 02°37'E,
coast of Dahomey, West Africa. Coll. A. Crosnier, 24 October
1963; ex H. Eker. Dimensions (H x W): 75.1 x 19.9 mm; height
of aperture = 38.0 mm [published as 76.8 x 21.0 mm, height of
aperture = 38.6 mm].

Remarks: The holotype (DMNH 022392) and paratype (nos. 4-5,
DMNH 028221) lots were described in Part I (Bieler & Bradford,
1991: 11). At the tme that paper was published, this paratype was

November 30, 1997 Mikkelsen & Bradford 1]

"in the author's collection, Auckland Institute & Museum";

Cernohorsky's numbered collection label accompanies the lot.

"Paratype No 3 of crosmiert, coasts of Dahomey" 1s written in ink on

the inner lip of the shell. Cernohorsky (1991) listed the DMNH

holotype, but omitted mention of the two DMNH paratypic lots.
Mitridae.

diminutum, Incidostoma [as n. sp.]

Morrison, 1955: 159, figs. 7-9.

1 paratype: DMNH 146762; Papallagta, Ecuador. Ex R. W. Jackson
(#6383) via M. L. Walton (#8951). Dimensions (H x W): 12.7 x
13.38 mm.

12 paratypes: DMNH 149232; Papallagta, Ecuador. Coll. R. W.
Jackson; ex R. W. Jackson (#6383). Dimensions (range): H = 8.3-
12,7 mm, W = 10.5-20,0 mm.

Remarks: The original description mentioned paratypes "in the Jackson
collection.” ‘The Jackson collection catalog indicates "det. Dr.
Morrison."

Poteriidae [originally Cyclophoridae: Neocyclotinae]. Curated in the
DMNH collection as a junior synonym of Cyrtotoma granulatum
(Pieitfer, 1862).

dondant1, Mitra [as sp. n.]

Cernohorsky, 1985: 56-58, figs. 19-24.

1 paratype: DMNH 197607; dredged at 100 fms, Panglao, Bohol,
Philippines. Coll. January 1985; ex H. Eker. Dimensions (H x W):
Sdladex f2. 57mm:

Remarks: Original label indicates a "51.56 mm giant" of Mitre [sic]
deynsert Cernohorsky for the price of $150.00 from Derry's Shell
Shelter, La Habra, California. Paratypes from the type locality
were given in the original description as in several private
collections, including that of Eker. The original description listed

the maximum recorded shell length as 50.0 mm.
Mitridae.

earle1, Mitra (Nebularia) [as sp. n.]
Cernohorsky, 1977: 121-123, figs. 1-5.
1 paratype: DMNH 197617; 100 fms, Pokai Bay, Oahu, Hawaii. Coll.
R. Salisbury, 11 October 1976; ex Auckland Institute & Museum
via H. Eker. Dimensions (H x W): 13.8 x 4.9 mm.
Remarks: The original description listed paratypes in the Eker collection,
the source of this specimen. Cernohorsky (1977, 1991) twice listed

iZ NEMOURIA No. 41

DMNH among repositories for paratypes, without number
reference; because no specimens of this species were part of the
DMNH collection prior to the Eker donation in September 1995,
it is assumed that DMNH never received the intended paratypic
lot. The specimen contains dried blackened material within the
shell, perhaps dried soft parts.

Mitridae.

edgerly1, Gyrineum perca [as n. var.]

Richards;-1 933: 57, pl. 6, fig. 2.

Holotype (figured): DMNH 195251; Japan. Ex Mrs. E. R. Edgerly via
Mystic Museum. Dimensions (H x W): 62.9 x 45.7 mm [published
as 6.6 x 3.7 cm).

Remarks: Although this specimen does not exactly match the published
dimensions, it very closely approximates the orignal figure,
including several visible breaks and growth flaws.

Ranellidae [none specified originally]. Placed in synonymy with G. perca
(Perry, 1811) by Henning & Hemmen (1993), but without citing
any type material for G. perca or G. edgerlyr.

ekerae, Scabricola (Swainsonia) ocellata [as subsp. n.]

Cernohorsky, 1973: 135-137, figs. 8-12.

1 paratype: DMNH 197605; 22°43'30"S, 113°40'E, southwest of Point
Cloates, Western Australia, Australia. Coll. Ningaloo Expedition,
O7 September 1968; ex WAM via H. Eker. Dimensions (H x W):
23.0 x. Oo) aim:

Remarks: This specimen is listed here as a possible paratype. The
Cernohorsky index card in the DMNH collection does not list the
source (Eker) collection specifically; neither was mention made of
the Eker collection in the list of paratypes in the original
description. However, the original description listed six paratypes
from Point Cloates, five in WAM (confirmed by Wells, 1977) and
one "in author's collection." The paratype listed as in the author's
collection closely approximates the dimensions of this specimen
(33.0 x 10.0 mm); we assume that this is the DMNH specimen.

Mitridae [originally Mitridae: Imbricarinae].

eugeniae, Notadusta [as spec. nov.]
C. NaCate, 19752260, fig. 5.
1 paratype: DMNH 196224; Hopetoun, southern Western Australia,
Australia. Ex L. F. Kienle. Dimensions (H x W): 27.1 x 15.2 mm.
1 paratype ("No. 1"): DMNH 201199; Hopetoun, southern Western

November 30, 1997 Mikkelsen & Bradford 13

Australia, Australia. Coll. 14 March 1974; ex E. Wnght (#C290,
"paratype 1, sub-adult, 27.5 mm, $75.00"). Dimensions (H x W):
216 = Ail im,

Remarks: The measurements of DMNH 201199 exactly match those
cited in the original description ("paratype I").

Cypraeidae [originally Cypraeidae: Erroneinae]. Synonymized with
Cypraea declivis Sowerby, 1870 (Burgess, 1985).

fergusoni, Acmaea [as sp. nov.]

Wheat, 1913: 17-20, pl. 1.

1 syntype: DMNH 023402; Hempstead Bay, Long Island, New York.
Ex Brooklyn Museum via ANSP 185472 (vidz). Dimensions (H x
W): 19.3 x 14.8 mm; depth = 6.5 mm.

Remarks: This specimen is listed here as a possible syntype. The original
description listed "Types from the north shore of Long Island, N.
Y., in the Museum of The Brooklyn Institute of Arts and Sciences,
Accession No. 12656." The Brooklyn Museum collection was
subsequently transferred to ANSP (Robertson, et a/., 1981: 19).
Hempstead Bay was not listed among the author's collecting sites.
The source lot (ANSP 185472) is labelled "types" and now
contains 61 specimens filed as Acmaea testudinalis var. fergusont;
other labels with the lot indicate "Central Museum," catalog no.
12656, and "Hempstead Bay, VI.17.13" QGune 17, 1913?).
Robertson, et al. (1981) listed the source lot as syntypic.

Lottiidae [none specified originally]. Now known as Tectura testudinalis
ferguson (fide Jacobson & Emerson, 1971; Lindberg, 1986).

filareginae, Vexillum regina [as subspec. nov. |]

J. M. Cate, 1961: 80-81, pl. 18, figs. 6a-b, pl. 19, fig. 6, pl. 20, figs. 1-
10, table.

1 figured paratype (#6): DMNH 197602; Zamboanga, Philippines. Ex
H. Eker. Dimensions (H x W): 67.0 x 17.0 mm [published as 67.4
x 17.5 mm]. Remarks: A sticker inside the aperture indicates "124
Para #6." The specimen has a complete thickened outer lip. This
specimen was figured by Cate on pl. 19, fig. 6 and pl. 20, fig. 7.

1 figured paratype (#7): DMNH 197603; Zamboanga, Philippines. Ex
H. Eker. Dimensions (H x W): 65.0 x 15.0 mm [published as 65.2
x 15.4 mm].). Remarks: A sticker inside the aperture indicates
"124 Para #7." The specimen has a thin, chipped outer lip. This
specimen was figured by Cate on pl. 20, fig. 8.

Remarks: The original description listed paratypes #6 and #7 as in the
d'Attilio collection. Although these specimens, from the Eker

14 NEMOURIA No. 41

collection, do not indicate a source, they are clearly paratypes #6
and #7 (via stickers in the aperture and matching the published
figures).

Costellariidae [originally Mitridae]. This subspecies was synonymized
with Vexillum regina (Sowerby, 1828) by Cernohorsky (1970b).

fuentesi, Chondropoma (Chondropoma) [as spec. nov.]

Jaume & Alcalde y Ledon, 1944: 39-40, pl. 6.

3 paratypes: DMNH 124279; finca de Lewis, Arimao, Cienfuegos, Las
Villas, Cuba. Ex O. Alcalde (Jackson catalog) via R. W. Jackson
(#6633). Dimensions (H x W): (1) 14.5 x 8.5 mm, (2) 12.9 x 8.4
Mim, (3) 12.2 x 8.2 mim:

Remarks: The original description mentioned paratypes in private
collections, 1. e., "en las colecciones del Dr. Luis Sanchez de
Fuentes y de los autores." The Jackson collection catalog in the
DMNH archives indicates the source of these specimens as
Alcalde. All specimens have "4153" written on them; significance
of this number is unknown.

Annularidae [none specified originally].

fuentes1, Chondrothyrium crenimargo [as nueva subespecie]

Alcalde y Ledon, 1943: 11-13, pl. 1, fig. 1.

1 paratype: DMNH 135185; Loma #3 del Marafion, Gavilan,
Cienfuegos, Las Villas, Cuba. Ex O. Alcalde via R. W. Jackson
(#6612). Dimensions (H x W): 15.0 x 10.5 mm.

Remarks: The original description mentioned paratypes "en las
collectiones del autor, del Dr. Luis Sanchez de Fuentes y de
Miguel L. Jaume." The DMNH shell has "1449" written on it; the
original [pon-DMNH] labels have "6612" inscribed. The Jackson
collection catalog in the DMNH archives indicates Alcalde as the
source for #6612. The DMNH specimen does not match the
dimensions of any of the 12 specimens cited in the omnginal
description.

Annulariidae [none specified originally].

guamensts, Favartia [as n. sp.]
Emerson & D'Attilio, 1979: 4-5, figs. 11-12.
1 paratype: DMNH 201173; 60 ft, under dead coral, scuba, Orote
Cliffs, Orote Peninsula, Guam Is., Mariana Islands. Coll. 1978
[published as 1977]; ex A. Deynzer via E. Wright (#M991, "6.4
mm, $25.00"). Dimensions (H x W): 6.7 x 4.3 mm.
Remarks: Although A. Deynzer's collection label is not present, Wnght's

November 30, 1997 Mikkelsen & Bradford 15

collection catalog indicates that this specimen was purchased from
Deynzer.

Muricidae [originally Muricidae: Muricopsinae]. This is "probably a
junior synonym of Favartia crouchi (Sowerby, 1894)" according to
Houart (1994: 54).

guesti, Calliostoma [as new species]

Quinn, 1992: 106-107, figs. 92-93.

Holotype: DMNH 096994; 100-120 fms, 2.5 mi off South Shore,
Bermuda. Coll. A. Guest & J. R. H. Lightbourn, 21 September
1973; ex A. Guest & J. R. H. Lightbourn. Dimensions (H x W):
27.4 x 27.1 mm [published as 27.5 x 27.5 mm].

3 paratypes (including figured): DMNH 187588; 100-120 fms, 2.5 mi
off South Shore of Bermuda. Coll. A. Guest & J. R. H. Lightbourn,
21 September 1973; ex A. Guest & J. R. H. Lightbourn.
Dimensions (H x W): (1) 23.1 x 22.7 mm, (2) 22.7 x 21.9 mm
[figured specimen], (3) 16.2 (apex broken) x 17.4 mm. Remarks:
Split from DMNH 096994.

Remarks: The figured specimen (Quinn, 1992: figs. 92-93) is not the
holotype as stated, but rather is the second largest paratype,
photographed when the specimen was still part of lot DMNH
096994. The cited dimensions of the holotype in the figure
caption, although not exactly matching measurements presented
here, are presumably those of the holotype and not the figured

paratype.
Trochidae [originally Trochidae: Calliostomatinae].

hadari, Drupa ricinus [as new subspecies]

Emerson & Cernohorsky, 1973: 23-24, pl. 2, figs. 9-10, pl. 18.

2 paratypes: DMNH 051118; on coral reefs, Eilat, Gulf of Aqaba, Israel.
Coll. D. Peled, April 1971; ex D. Peled. Dimensions (H x W,
including spines): (1) 35.0 x 36.8 mm, (2) 27.1 x 29.2 mm.

Remarks: The original description indicated DMNH 051119 as a
paratypic lot with (at least) one specimen of 28.0 mm length x
29.0 mm width - roughly equal to the smaller specimen in this lot.
The original specimen label (ex Auckland Institute & Museum)
indicates "2 paratypes DMNH 51118/9." DMNH 051119 was
assigned to a lot of another species, so we believe this is the type
lot.

Muricidae [originally Muricidae: Thaidinae].

16 NEM OURIA No. 41

hansenae, Vexillum (Pusia) [as sp. n.]
Cernohorsky, 1973: 138, figs. 13-16.
1 paratype: DMNH 197620; Cape Riche, ca. 90 mi east of Albany,
Western Australia. Coll. M. P. Marrow, 26 December 1971; ex H.
Eker. Dimensions (H x W): 10.3 x 4.2 mm (worn). Remarks: Lot
listed (not as type) on the Cernohorsky index card at DMNH. The
original description listed ten specimens from Cape Riche from the

collection of M. Marrow, including one measured as 10.8 x 4.7 x

5.9 mm, which could be this one.

2 paratypes: DMNH 197624; Augusta, southwestern Australia. Ex
Auckland Institute & Museum via H. Eker. Dimensions (H x W):
(1) 15.6 x 6.0 mm, (2) 16.0 x 6.2 mm. Remarks: Lot listed (not as
type) from Sarge Bay (= type locality) on the Cernohorsky index
card at DMNH. The onginal AIM label indicates three paratypes,
but only two are in this lot.

Remarks: The original description mentioned paratypes in the Eker
collection, the source of these two lots.

Costellariidae [originally Vexillidae].

hilli, Anachis (Glyptanachis) [as n. sp.]

Pilsbry & Lowe, 1932: 73, pl. 5, fig. 6.

2 paratypes: DMNH 024377; Mazatlan, Mexico. Coll. H. N. Lowe; ex
ANSP 154677 (vidi). Dimensions (H x W): (1) 8.2 x 4.0 mm, (2)
7.2 x 3.4 mm.

Remarks: The original description mentioned "paratypes in Lowe
collection." The ANSP source lot (orginally ten specimens, now
eight) was donated by H. N. Lowe and is presently considered
paratypic in that collection.

Columbellidae.

inspinata, Melongena corona [as n. var.]

Richards,.1933:575 pl. 6, fig.-1.

Holotype (figured): DMNH 195252; Gulf of Mexico, Florida. Coll. J.
R. Holmes; ex Mystic Museum. Dimensions (H x W): 93.2 x 62.8
mm [published as 10.1 x 6.2 cm].

Remarks: Although this specimen does not exactly match the published
dimensions, it very closely approximates the orignal figure,
including several visible growth flaws. DMNH records do not
reflect the original location of the material ["collection of Mrs. E.
R. Edgerly, of Trenton, N. J."].

Melongenidae [none specified originally]

November 30, 1997 Mikkelsen & Bradford 17

jacksont, Incidostoma [as n. sp.]

Morrison, 1955: 158, figs. 13-15.

16 paratypes: DMNH 135090; Mera, Oriente, Ecuador. Ex R. W.
Jackson (#6384). Dimensions (range): H = 11.6-16.1 mm, W =
18.2-24.9 mm.

2 paratypes: DMNH 146761; Mera, Oriente, Ecuador. Ex R. W.
Jackson (#6384) via M. L. Walton (#8952). Dimensions (H x W):
(1): 15.4 %25.7 min, (2) 13.0 x 22.9 mim.

Remarks: The original description noted "numerous paratypes
comprising the remainder of this lot are in Mr. Jackson's
collection." The Jackson collection catalog indicates "det. Dr.
Morrison."

Poteriidae [originally Cyclophoridae: Neocyclotinae]. Curated in the
DMNH collection in the genus Cyrtotoma.

gacksonti, Scissurella [as n. sp.]

Melvill, 1904: 160-161, pl. 10, fig. 5.

1 syntype: DMNH 023171; Gulf of Oman, Arabia. Coll. F. W.
‘Townsend, "1935"; ex ANSP 164838 (vid7). Dimensions (H x W):
1.4x 1.5 mm.

Remarks: No repository was designated in the original publication. The
source lot, ANSP 164838, was listed as a syntypic lot by
Robertson, et al. (1981) and by Trew (1987); the DMNH lot was
not acknowledged by Trew (1987). The date 1935 associated with
this lot is that of acquisition/purchase [by ANSP], not the year
collected.

Scissurellidae [none specified originally].

jaumet1, Chrondrothyrium violaceum [as nueva subespecie]

Alcalde y Ledon, 1943: 14-16, pl. 1, fig. 2.

4 paratypes: DMNH 135092; La Travesada, Rio Chiquito, Cienfuegos,
Las Villas, Cuba. Ex R. W. Jackson. Dimensions (range): H =
18.3-21.3 mm, W = 13.5-15.1 mm.

Remarks: The original description mentioned paratypes "en la colleccion
del autor, y en las de M. L. Jaume y Dr. L. Sanchez de Fuentes."
Each DMNH specimen has "1381" written on the shell; two
original [pon-DMNH] labels have "6614" inscribed. The DMNH
specimens do not match the dimensions of any of the 12 specimens
cited in the original description. The Jackson collection catalog in
the DMNH archives indicates Alcalde as the source for no. 6614.

Annulariidae [none specified originally].

18 NEMOURIA No. 41

gaumet, Eutrochatella (Troschelviana) [as spec. nov.]

Aguayo & Jaume ["Torre (in sched)"], 1957: 120-121, pl. 1, fig. 8.

4 paratypes: DMNH 161859; "La Esperanza," San Andres, Consolacion
del Norte, Pinar del Rio, Cuba. Leg. M. L. Jaume (Jackson
catalog), ex R. W. Jackson (#4198). Dimensions (H x W): (1) 7.4
x 5.4 mm (flaring outer lip), (2) 7.1 x 5.1 mm (flaring outer lip),
(3) 6.8 x 4.8 mm, (4) 5.9 x 4.4 mm.

Remarks: The original description mentioned paratypes in the collection
of M. L. Jaume. Specimens received in the Jaume Collection (e. g.,
ANSP 167331), were considered paratypic by Robertson, er al.
(1986). All four specimens are with opercula.

Helicinidae. Curated as a junior synonym’ of Troschelviana
(Troscheluiana) chrysochasma Poey, 1853, in the DMNH collection.

jenniernestae, Distorsio (Distorsio) [as sp. nov.]

Emerson & Piech, 1992: 111-114, figs. 5-8, 16-18, 24.

2 paratypes: DMNH 189600; dredged between Cébaco and Coiba
Islands, Pacific Panama, 240 ft. Coll. Ernest, 1990; ex B. J. Piech.
Dimensions (H x W): 58.5 x 36.3 mm (cleaned), (2) 55.3 x 35.2
mm (with periostracum).

Remarks: Both specimens are with opercula.

Personidae.

lopesi, Mitra (Cancilla) [as sp. n.]

Matthews & Coelho, 1969: 3-6, figs. 1-3.

3 paratypes: DMNH 197614; 16 fms, ex pisces [from digestive tract of
toadfish, Amphichthys cryptocentrus (Cuvier & Valenciennes,
1837)], off Mucuripe, Fortaleza, Ceara, Brazil. Coll. by fishermen,
April 1968; ex H. R. Matthews via W. O. Cernohorsky via H.
Eker. Dimensions (H x W): (1) 22.6 x 8.9 mm (worn, apex
broken, hole in ventral body whorl), (2) 19.4 x 7.5 mm (worn,
apex broken), (3) 17.7 x 7.1 mm (fresh specimen, broken outer lip,
hole in spire).

Remarks: Four specimens listed as ex pisces from this locality (collected
in 1969, not 1968) were cited in the original description from other
depositories [Instituto Oswaldo Cruz (IOC), Rio de Janeiro;
Laboratorio de Ciéncias do Mar da Universidade Federal do Ceara
(LABOMAR), Fortaleza; Museu Nacional (MN), Rio de Janeiro;
and Museu Oceanografico de Rio Grande (MORG), Rio Grande
do Sul]. Although the original description therefore did not
specifically allow for the DMNH specimens, they are listed as
paratypes in the Cernohorsky collection on Cernohorsky's index

November 30, 1997 Mikkelsen & Bradford 19

card in DMNH,, therefore they are retained here as possible type
material.

Mitridae. This is a jumior synonym of Ziba candida (Reeve, 1845)
according to Cernohorsky (1991).

loutsianae, Vioscalba [as new species and type species of new genus]

Morrison, 1965: 217, 219-220, figs. 1-2.

23 paratypes: DMNH 108365; off Frenier Beach, Lake Ponchartrain,
Louisiana. Coll. 05 November 1959, J. P. E. Morrison; ex USNM
635629 (with label indicating DMNH donation, zvid7). Dimensions
(range): H = 2.7-3.4 mm, W = 1.7-2.1 mm.

Remarks: The original description mentioned "more than twenty-five
additional paratype lots" in USNM.

Hydrobiidae. Curated as a junior synonym of Probythinella protera
(Pilsbry, 1953) in the USNM collection.

mactanensis, Murexiella [as n. sp.]

Emerson & D'Attilio, 1979: 7, figs. 7-8.

1 paratype: DMNH 201172; 100+ fms, Panglao, Bohol Island,
Philippines. Coll. 1977-1978 [published as 1977]; ex A. Deynzer
via E. Wright (#M939, "19.5 mm, w/o [with operculum],
$35.00"). Dimensions (H x W): 19.7 x 12.7 mm.

Remarks: Deynzer's collection label is present.

Muricidae [originally Muricidae: Muricopsinae].

marrow, Vexillum (Pusia) [as sp. n.]

Cernohorsky, 1973: 140-142, figs. 19-23.

2 paratypes: DMNH 197626; Margaret River, southwestern Australia.
Ex Auckland Institute & Museum via H. Eker. Dimensions (H x
W): (1) 10.0 x 5.7 mm, (2) 9.4 x 4.7 mm. Remarks: The original
AIM label indicates four paratypes, but only two are found in this
lot. Lot listed (not as type, from south of Cape Naturaliste) on the
Cernohorsky index card at DMNH.

1 paratype: DMNH 197627; Rottnest Island, Western Australia,
Australia. Coll. Kendrick, 1969; ex W. O. Cernohorsky via H.
Eker. Dimensions (H x W): 11.1 x 6.2 mm. Remarks: Listed (not
as type, from Lady Adeline Bay) on the Cernohorsky index card at
DMNH.

1 paratype: DMNH 197628; North Beach, Perth, Western Australia.
Coll. Rossell, 1950; ex W. O. Cernohorsky via H. Eker.
Dimensions (H x W): 11.1 x 6.2 mm (apex and outer lip broken).
Remarks: Listed (not as type) on the Cernohorsky index card at

20 NEMOURIA No. 41

DMNH.

1 paratype: DMNH 197629; South Cottesloe Beach, Western Australia.
Coll. T. Whitehead, November 1966; ex W. O. Cernohorsky via
H. Eker. Dimensions (H x W): 11.7 x 5.7 mm (outer lip chipped).
Remarks: Listed (not as type) on the Cernohorsky index card at
DMNH.

Remarks: The original description mentioned paratypes in the Eker
collection, the source of all four of these lots. Margaret River was
listed as the locality of four specimens "In H. Eker coll.,"
presumably including the two in DMNH 197626. Lady Adeline
Bay - Rottnest Island, North Beach - Perth, and South Cottesloe
Beach - Perth were each listed as including "1 spec. in author's
coll." = DMNH 197627, 197628 and 197629, respectively.

Costellariidae [originally Vexillidae].

martinorum, Vexillum (Costellaria) [as sp. n.]

Cernohorsky, 1986: 50-52, figs. 15-20.

1 paratype: DMNH 197611; 183-207 m, off Coamen Island, west
Bohol reef, Philippines. Coll. R. Martin, 1986; ex H. Eker.
Dimensions (H x W): 19.6 x 6.7 mm.

Remarks: The original description mentioned paratypes in the Martin
collection.

Costellariudae.

mawson1, Parmaphorella [as n. sp.]

Powell, 1958: 180-181, text-figs. 1-3, pl. 3, fig. 9.

1 paratype: DMNH 023360; off Enderby Land, Antarctica. Coll. A. W.
B. Powell; ex ANSP 226808 (BANZARE sta. 41, 193 m, widz).
Dimensions (H x W): 21.0 x 11.3 mm; depth = 5.1 mm.

Remarks: The original description indicated type material only in the
South Australian Museum (Adelaide), so under strict
interpretation, this could not be type material. However, the
ANSP source lot was listed by Robertson, et al. (1981: 22), as
syntypic, and the "catalog shows two specimens received. Only one
specimen is now in the collection." Because the original description
designated a holotype, remaining type specimens must be
considered paratypes, not syntypes.

Fissurellidae. Curated in the DMNH oollection in Tugal
(Parmaphorella).

morenot, Chondrothyrium (Chondrothyrium) violaceum [as n. s. sp.]
Jaume & Sanchez de Fuentes, 1943d: 50-51, pl. 8.

November 30, 1997 Mikkelsen & Bradford Ze

2 paratypes: DMNH 135187; loma #3 del Maranon, Gavilan,
Cienfuegos, Las Villas, Cuba. Ex O. Alcalde (Jackson catalog) via
R. W. Jackson (#6619). Dimensions (H x W): (1) 16.9 x 11.5
mm, (2) 14.2 x 10.0 mm.

Remarks: The original description mentioned paratypes in private
collections, 7. e., "en las colecciones de los autores y en la del Sr.
Oscar Alcalde Ledon." The Jackson collection catalog indicates
Alcalde as the source of these specimens. Both specimens have
"1414" written on them.

Annulariidae [none specified originally].

nanus, Strombus raninus [as new subspecies]

Bales, 1942: 19, pl. 4, figs. a-b.

2 paratypes: DMNH 201259; southern end of Lake Worth, Palm Beach
County, Florida. Ex R. T. Abbott (#1544 on label that states
"rec'd from R. T. Abbott"). Dimensions (H x W): (1) 49.9 x 33.4
mm ("1544" written on thin but fully developed outer lip); (2) 37.8
x 26.7 ("1082" written on thickened outer lip).

Remarks: The original description indicated [holo]type and paratypes in
ANSP, MCZ, and private collections of Bales, McGinty, and
Koto. The origin of this material, other than R. T. Abbott, is
unrecorded. Another paratypic lot, DMNH 001490, was described
in Part I (Bieler & Bradford, 1991: 23-24). These specimens are
listed here as possible paratypes.

Strombidae [none specified originally].

nodospiculum, Vexillum (Costellaria) [as new species]

Cernohorsky, 1970a: 101-102, figs. 6, 7, 11.

1 paratype ("No. 47"): DMNH 197618; U. S. Bureau of Fisheries Sta.
5592, 305 fms, green mud, 43.3°F, off Silungan Island, Borneo,
Indonesia. Ex H. Eker. Dimensions (H x W): 6.0 x 2.1 mm (with
intact, but somewhat encrusted, protoconch).

1 paratype ("No. 13"): DMNH 197622; U. S. Bureau of Fisheries Sta.
5394, 153 fms, green mud, off Pt. Dumurug, Masbate,
Philippines. Ex H. Eker. Dimensions (H x W): 5.4 x 2.2 mm (with
intact protoconch).

Remarks: At the time of publication, both paratypes were "in the author's
collection, Auckland Institute & Museum"; Cernohorsky's
numbered collection labels are with both lots. No paratypes are
listed on the Cernohorsky index card for this species at DMNH.

Costellariidae [originally Mitridae].

Ze NEMOURIA No. 41

orr1, Muricopsis [as sp. n.]

Cernohorsky, 1976: 116-119, figs. 12-20.

2 paratypes (including figured): DMNH 201174; dredged, 30 fms,
south of the Andaman Islands, Indian Ocean. Coll. January 1974;
ex Auckland Institute & Museum/W. O. Cernohorsky via E.
Wrght (#M783). Dimensions (H x W, including spines): (1) 32.3
x 22.1 mm, (2) 28.8 x 22.6 mm (with operculum in place,
figured).

Remarks: These specimens were mentioned in the original description
from the Wright collection. The measured dimensions differ
slightly from those listed for Wnght paratypes 1-3 in Cernohorsky's
text: 32.5 x 20.0, 29.1 x-22.0,;and 29.0 x 20:2, mm. Whese
specimens are apparently paratypes 1 and 3; Cernohorsky's figs.
14-15 are of paratype 3 = DMNH paratype (2). According to
Wright's collection catalog, four specimens were originally
purchased from Vic Wee for $5.00 each as Muricopsis infans (E. A.
Smith, 1884) (as 26, 26, 27 [circled], and 30 mm). The entry also
indicates "2 w/o [with operculum], trawled in 30 fms, one
specimen kept as holotype, others paratypes; 1 paratype sent to
John Orr, 12/21/76."

Muricidae. Now in the genus Attiliosa according to Houart (1994).

petiti, Fenimorea [as new species]

Tippett, 1995: 133, figs. 17; 33.

1 paratype: DMNH 202363; dredged, 30 fms, Gulf of Mexico west of
Crystal River, Florida. Coll. Jim Moore, May 1963; ex D. Tippett.
Dimensions (H x W): 14.7 x 5.1 mm.

Turridae.

rhecta, Meganipha [as new species]

F. G. Thompson, 1978: 43-48, figs. 1-4.

3 paratypes: DMNH 111264; 700 m, Loma del Puerto, Yaroa, Puerto
Plato Province, Dominican Republic, Hispaniola. Coll. F. G.
Thompson & B. E. Johnson (#2659), 12 January 1976; ex
FLMNH (= UF) 22747. Dimensions (H x W): (1) 10.3 x 8.0 mm
(with operculum), (2) 9.9 x 9.1 mm, (3) 2.1 x 2.5 mm Guvenile).

Remarks: The source lot, UF 22747, originally contained 114 paratypes,
according to the original description.

Annulariidae [originally Annulariidae: Annulariinae].

salisbury1, Vexillum (Pusia) [as sp. n.]
Cernohorsky, 1976: 114-116, figs. 6-9, 11.

November 30, 1997 Mikkelsen & Bradford 23

1 paratype: DMNH 155900; Pupukea Beach, Oahu, Hawaii. Coll. R.
Salisbury, April 1976; ex C. Kauffman. Dimensions (H x W): 3.8
x 1.9 mm (protoconch missing, outer lip badly broken).

2 paratypes (#47): DMNH 197619; in beach sand, Pupukea, Hawaiian
Islands. Coll. R. Salisbury, 1975; ex Auckland Institute & Museum
via H. Eker. Dimensions (H x W): (1) 4.3 x 2.0 mm (very pale
pink, outer lip broken), (2) 4.0 x 2.0 mm.

Remarks: Paratypic lot DMNH 112102 was described in Part I (Bieler
& Bradford, 1991: 27); these are the paratypes mentioned (without
number) in the original description as in DMNH. The two lots
above were mentioned in the original description from the
collections of Eker and Salisbury.

Costellariidae. Considered a junior synonym of Vexillum (Pusia)
capricornea (Hedley, 1907) by Kay (1979).

scherer1, Helicina [as n. sp.]

F. Baker, 1913: 625-626, pl. 21, figs. 1-2.

4 paratypes: DMNH 135243; Ceara-Minm, Brazil. Leg. F. Baker; ex R.
W. Jackson (#2400). Dimensions (H x W): (1) 4.4 x 5.3 mm (with
operculum), (2) 4.4 x 5.1 mm, (3) 3.6 x 4.5 mm (with
operculum), (4) 2.6 x 3.6 mm (thin outer lip).

Remarks: The original description mentioned [holo]types of all new
species in ANSP, and that "cotypes of many will be" deposited in
the Leland Stanford Jr. University (Palo Alto, CA) and the Museu
Goeldi (Para, Brazil). This allows for additional type material
retained by Baker, thus these are here considered possible
paratypes (Baker's original "cotypes" can be considered paratypes
because he formally designated [holo]types). The [holo]type lot
was given as ANSP 109341a by Baker (1964).

Helicinidae.

serranum, Farcimen (Farcimen) auriculatum [as n. subsp.]

Alcalde y Ledon, 1945: 14, pl. 1, fig. 6.

Holo(?)type: DMNH 135223; Aguada de La Piedra, La Sierra,
Cienfuegos, Las Villas, Cuba. Ex O. Alcalde via R. W. Jackson
(#6562). Dimensions (H x W): 26.4 x 13.2 mm.

Remarks: The original description mentioned only one type specimen,
which would thus be the holotype, in the collection of the author
(no. 3467) and of dimensions close to those of this specimen (27
x 13 mm). The DMNH specimen was listed as a “paratype” in the
Jackson collection catalog. This could therefore be the holotype.

Megalomastomidae [originally Cyclophoridae: Megalomastominae].

24 NEMOURIA No. 41

sphoni, Mitra (Strigatella) [as spec. nov. ]

Shasky & Campbell, 1964: 118-119, pl. 22, figs. 13-14.

1 paratype: DMNH 197625; 30-40 ft, under rocks, between Punta
Colorada and Punta Lobos, Guaymas, Sonora, Mexico. Coll. D.
R. Shasky, 1964; ex W. O. Cernohorsky via H. Eker. Dimensions
(H x W): 16.9 x 6.0 mm.

Remarks: This specimen is listed here as a possible paratype. The
original description mentioned two paratypes, one each in the B.
Campbell and D. Shasky collections; we suspect the latter is this
specimen. The Cernohorsky index card in the DMNH collection
does not list this specimen.

Costellariidae [originally Mitridae]. Curated in the DMNH collection as
a junior synonym of Vexillum crenata (Broderip, 1836).

sprucecreekensis, Melongena [as new species]

Tucker, 1994: 197-202, figs. 6A-C.

10 paratypes: DMNH 155535; Spruce Creek, 4 miles north of New
Smyrna Beach, Florida. Coll. J. K. Tucker, 07 June 1983 [other
labels give 9-1983 and 6/17/83]. Dimensions (range): H = 89-169
mm, W = 46-90 mm.

Remarks: Formerly DMNH 161854. All specimens are with opercula.

Melongenidae.

superbum, Chondropoma [no status specified in original description]

Henderson & Simpson, 1902: 88-89, 1 text-fig.

4 syntypes: DMNH 043596; Thomazeau, Haiti. Coll. Henderson &
Simpson; ex USNM. Dimensions (range): H = 22.1-24.6 mm, W
= 12.2-13.4 mm.

Remarks: No type material was specifically cited in the original
description. These specimens are from the type locality and are
within the size range given. The source of this lot (USNM) is that
expected for type specimens of both J. B. Henderson and C. T.
Simpson, by virtue of their simultaneous association with USNM
(Abbott, et al., 1973). In spite of the lack of more concrete
evidence, these specimens are listed here as possible syntypes.

Annulariidae [none specified originally].

tagbilleranus, Geophorus (Diplopinax) [as n. sp.]
Bartsch, 1918: 655-656.
6 syntypes: DMNH 043615; Tagbileran, Bohol Island, Philippines. Ex
USNM 258760. Dimensions (range): H = 6.2-7.0 mm, W = 9.8-

November 30, 1997 Mikkelsen & Bradford 25

13 mim.

Remarks: Although the original description discussed the dimensions of
the "type," obviously referring to a single specimen, the original
label in this lot indicates that it was split from the cited type lot.
The type lot was listed by Ruhoff (1973) as USNM 258760.

Helicinidae [originally Helicinidae: Helicininae].

toroensis, Rissoa [as new species]

Olsson & McGinty, 1958: 26, pl. 4, fig. 5.

5 paratypes: DMNH 023673; Bocas Island, northeastern Panama. Coll.
Olsson and McGinty; ex ANSP 211886 (vd). Dimensions
(range): H = 1.3-2.1 mm, W = 1.0-1.2 mm.

Remarks: The ANSP source lot was cited as paratypic by the original
description; it 1s curated in ANSP in the genus Odostomia, and
contains ca. ten specimens.

Rissoidae.

torre1, Chondrothyrium (Plicathyrium) [as nueva especie]

Jaume & Sanchez de Fuentes, 1943c: 25-27, pl. 4, fig. 2.

1 paratype: DMNH 124280; Loma Ventana, San Blas, Cienfuegos, Las
Villas, Cuba. Ex O. Alcalde via R. W. Jackson [#6621, as C. (P.)
torret torret]|. Dimensions (H x W): 13.9 x 14.0 mm.

Remarks: "1460" is written on the shell. The original description
mentioned paratypes in the collections of the authors and O.
Alcalde.

Annulartidae [none specified originally].

vicdant, Scabricola [as sp. n.]

Cernohorsky, 1981: 193-195, figs. 1-3.

1 paratype: DMNH 197606; trawled by fishermen, Punta Enganfo,
Mactan Island, Cebu, Philippines. Coll. May 1981; ex H. Eker.
Dimensions (H x W): 36.1 x 13.9 mm (anterior end of outer lip
broken).

Remarks: The onginal description did not specifically allow for type
material of this species in the Eker collection, and Cernohorsky
(1991) did not list Eker among the repositories of type material.
However, the Cernohorsky index card at DMNH indicates
paratype #4 as in the H. Eker collection, from the type locality,
and measuring 37 mm; paratype #4 was deposited in the Auckland
Institute and Museum, according to the original description.
Because of this evidence plus the fact that Eker received several
other of her type specimens discussed here from the AIM, we

26 NEMOURIA No. 41

believe that this specimen is paratype #4.
Miutridae.

wolfe1, Drillia [as new species]
Tippett, 1995 b27,. 129sigs. 1,255.31.
1 paratype: DMNH 202362; 20 fms, 15 km east of Cape Lookout,
North Carolina, from the Elmer Dewey Willis scallop plant,
Williston, North Carolina. Coll. Dr. Douglas Wolfe, March-May
1971; ex D. Tippett. Dimensions (H x W): 14.9 x 5.2 mm.
Turridae.

wolfe1, Vexillum (Costellaria) [as new species]

Cernohorsky, 1978: 63-64, figs. 4-5.

1 paratype: DMNH 197623; 60-70 fms, Pokai Bay, Oahu, Hawaiian
Islands. Coll. R. Salisbury, 1975; ex Auckland Institute & Museum
via H. Eker. Dimensions (H x W): 5.0 x 2.2 mm.

Remarks: Paratypic lot DMNH 112103 was described in Part I (Bieler
& Bradford, 1991: 31). The original description mentioned
paratypes in "AIM and other institutions and collections."

Costellariidae.

yoshidat, Katayama nosophora [as new subspecies]

Bartsch, 1925: 72; 1936: 23-24, pl. 1, fig. 1, pl. 2, fig. 9, pl. 4, fig. 2.

6 paratypes: DMNH 043606; Kurume, Kyushu [published as Kuushiu]
Island, Japan. Coll. S. Yoshida, February 1919; ex USNM.
Dimensions (range): H = 4.7-6.6 mm, W = 2.2-2.6 mm (3
specimens, immature lip), 2.5-3.0 mm (3 specimens, mature lip).

Remarks: These specimens are listed here as possible paratypes. The
original description cited the [holo]type (USNM 362024) plus
"several hundred additional specimens from the type locality”
(USNM 340953, subsequently called paratypes; Bartsch, 1936:
23). Although the USNM number is not recorded in the DMNH
lot, the USNM label indicates "paratypes" and it is not
inconceivable that these specimens were originally part of USNM
340953. The USNM label indicates "= Oncomelania nosophora
Robson."

Pomatiopsidae [none listed originally]. Curated in the genus
Oncomelania in the DMNH collections.

November 30, 1997 Mikkelsen & Bradford 20

HETEROSTROPHA

weberi, Odostomia [as new species]

Morrison; 1965: 221, fig. 3.

2 paratypes: DMNH 108364; north of Bayou Chene Fleur, northern
part of Barataria Bay, Louisiana. Coll. J. P. E. Morrison; ex
USNM 635639 (with label noting DMNH donation, zd1).
Dimensions (H x W): (1) 1.3 x 0.7 mm (fresher shell), (2) 1.2 x
O.7 mm (very worn).

Pyramidellidae.

OPISTHOBRANCHIA
(ADDITIONS TO PART I; BIELER & BRADFORD, 1991)

carolynae, Platydoris [as n. sp.]

Mulliner & Sphon, 1974: 209-212, figs. 2-5.

1 paratype (in alcohol): DMNH 064524; SA-3, Long Beach, northern
Santa Cruz Island, Galapagos Islands, Ecuador. Leg. S. Andrews,
21 December 1972; ex S. Andrews. Dimensions (L x W,
preserved): 56 x 33 mm.

Remarks: The measurements of this specimen are larger than the largest
size given in the original description. The length of the highly
curled slug was crudely measured as the length of a string laid upon
the specimen from anterior to posterior.

Platydorididae [originally Dorididae].

BIVALVIA

acapulcensts, Leda (Saccella) [as new species]

Pilsbry & Lowe, 1932: 107, pl. 17, figs. 1-2.

1 paratype: DMNH 020361; 20 fms, Acapulco, Guerrero, Mexico. Coll.
H. N. Lowe, 1930; ex ANSP 155634 (vid7). Dimensions (L x H):
9, 0-x 5. 1 mim:

Remarks: The original figures (one inside view, one outside view) were
labelled "type" by Pilsbry & Lowe, constituting a holotype
designation. The DMNH specimen (split from the ANSP "type"
lot), therefore, can be considered a paratype.

28 NEMOURIA No. 41

Nuculanidae [originally Ledidae]. Curated as Nuculana in the DMNH
collection.

angas1, Corbicula [as nouvelle espece]

Prime, 1864: 151-152, pl. 7, fig. 6.

1 paralectotype: DMNH 020859; Murray River, South Australia. Ex
ANSP 123556 (Charles M. Wheatley Collection, deposited by
University of Pennsylvania, vidz). Dimensions (L x H): 10.3 x 7.9
mm.

Remarks: The original description did not cite a depository. The DMNH
and ANSP source lot were cited by Counts (1991: 8) as
“paratypes,” although a lectotype has been designated (MCZ
176917; Johnson, 1959). This therefore is a paralectotype. The
ANSP source lot is now incorrectly labelled as “lectotype,” with
four original specimens recataloged as “paralectotypes’ (ANSP
358422).

Corbiculidae [none specified originally].

awajiensis, Corbicula [as n. sp.]

Pilsbry, 1901b: 407; 1907: 159, pl. 7, figs. 13-14.

1 syntype: DMNH 020846; Noda, Awaji Island, Japan. Coll. Y. Hirase,
1901; ex ANSP 89378 (wid). Dimensions (L x H): 15.0 x 11.3
mm.

Remarks: No repository, catalog number, or original number of
specimens was provided in the original description, however, the
data match the stated type locality and source/collector. Both the
DMNH and ANSP source lots were cited as syntypic by Counts
(1991: 9). ANSP 89378 is now the labelled "holotype" lot in
ANSP, with three specimens recataloged as a "paratype" lot
(ANSP 358424); this statement 1s not a lectotype designation.

Corbiculidae [originally Cyrenidae].

brunnea, Tindaria [as n. sp.]

Dall, 1916: 401.

1 syntype: DMNH 020368; ALBATROSS Sta. 3604, between St. Paul
and Unnak Islands, Bering Sea, off Alaska. Ex USNM via ANSP
190943 (vidi). Dimensions (L x H): 7.35 x 5.8 mm.

4 syntypes: DMNH 043603; ALBATROSS Sta. 3604, midway between
St. Paul and Unnak Islands, Bering Sea, Alaska. Ex USNM.
Dimensions (Lx H): (1) 7.1 x 5.6 mm, (2) 7.0 x 5.5 mm, (3) 6.9
x 5.2 mm, (4) 5.5 x 4.1 mm.

Remarks: These specimens are listed here as possible syntypes. The

November 30, 1997 Mikkelsen & Bradford 29

original description specified USNM 226333, of length 7.5, height
6, diameter 4 mm (which could have been an average), but without
specifying number of specimens. USNM 226333 was also given as
the sole type lot by Boss, et al. (1968). USNM 226333, ANSP
190943, and both DMNH lots are from the same ALBATROSS
station, no. 3604.

Tindariidae [none specified originally].

bryanae, Congeria [as n. sp.]

Pilsbry, 192): 323.

4 syntypes (single unmatched valves): DMNH 020569; Mokuaula
Island, Hawaii. Coll. C. M. Cooke; ex ANSP 144011 (vrdz).
Dimensions (range): L = 6.9-8.3 mm, H = 3.8-5.0 mm.

Remarks: These specimens are listed here as possible syntypes. No type
specimens were specifically cited in the onginal description,
however, Pilsbry's types were generally deposited in ANSP; the
source ANSP lot is labelled as type material in the ANSP
collection.

Mytilidae [none specified originally]. Now placed in the genus Septifer
(Kay, 1979).

calcicola, Nucula [as new species]
Moore, 1977: 120-122, figs. 1-3.
7 paratypes (6 intact + 1 single unmatched valve, in paleoslide): DMNH
120581; 2m, Chancanab Lagoon, Cozumel, Mexico. Coll. D. R.
Moore, 29 November 1971; ex D. R. Moore. Dimensions (range):
L = 0.6-1.4 mm, H = 0.4-1.0 mm.
Nuculidae.

congo, Iphigenia [as new species]

Pilsbry & Bequaert, 1927: 373-374, text-figs. 83a-c.

2 paratypes: DMNH 021548; Malela, Belgian Congo [now Zaire],
Africa. Coll. J. Bequaert; ex ANSP 133884. Dimensions (Lx H):
21.5 x 14.2 mm.

Remarks: These specimens are listed here as possible paratypes. The
original description gave dimensions of the "type" (27.2 x 18.0
mm), one "paratype" (24.0 x 16.1 mm), and two other specimens
(22.0 x 14.3 and 20.3 x 14.0 mm). However, the ANSP source lot
(the type lot?) contains many more specimens; it has now been
split into holotype (2 valves, ANSP 133884) and paratypes (17
valves, ANSP 399290) (N. Gilmore, in litt., 1997).

Donacidae.

30 NEMOURIA No. 41

cranmerorum, Somalipecten [as new species]
Waller, 1986: 42-45, figs. 1-13.
1 paratype: DMNH 170849; taken by Taiwanese trawler, off Somalia
coast at 150-300 m. Coll. 1985; ex D. Dan. Dimensions (L x H):
67.6 x 62.7 mm.
Pectinidae.

cynthiae, Condylonucula [as new species]

Moore, 1977: 123-124, figs. 6-7.

2 paratypes (1 intact + 1 single unmatched valve, in paleoslide): DMNH
120580; 1.5 m, behind outer reef, Courtown Cays, western
Caribbean. Coll. D. R. Moore, 12 May 1966; ex D. R. Moore.
Dimensions (L x H): (1) 0.6 x 0.5 mm (single unmatched valve),
(2) 0-5 x 0-5 mim. (ntact).

Remarks: The original description listed these specimens as one 570 um
long, and one left valve 620 um long.

Nuculidae.

diegensis, Limopsts [as n. sp.]

Dall, 1908: 395, pl. 15, figs. 13, 15.

3 syntypes (1 intact + 2 single unmatched valves): DMNH 020487;
ALBATROSS Sta. 2923, off San Diego, California. Ex USNM
111422 (fide ANSP catalog, and A. Kabat, pers. comm., 1996) via
ANSP 190954. Dimensions (L x H): (1) 9.1 x 8.4 mm (intact),
(2) 8.3 x 7.9 mm (single unmatched valve), (3) 6.9 x 6.3 mm
(single unmatched valve).

4 syntypes: DMNH 043614; ALBATROSS Sta. 2923, 822 fms, mud,
39°F, off San Diego, California. Ex USNM. Dimensions (range):
L = 8.6-11.6 mm, H = 7.8-11.0 mm.

Remarks: Although the original description (also Boss, et al., 1968)
mentioned only lot USNM 122585, A. Kabat called these lots
"assuredly valid syntypes" as a result of his research into Dall's
ALBATROSS species (A. Kabat, pers. comm., 1996), noting that
"Drs. Morrison and Rosewater routinely split off "paratypes" for
various other institutions..." Both lots were cited as type material
by Kabat (1996). All specimens have thick intact periostracum.

Limopsidae. A synonym of L. panamensis Dall, 1902 (E. V. Coan, pers.
comm.).

diegensts, Thracia [as new species]
Dall, 1915: 443.

November 30, 1997 Mikkelsen & Bradford 5]

1 syntype: DMNH 022250; near Beacon 10, San Diego Bay, San Diego
County, California. Coll. Kelsey; ex ANSP 162047 [4 fms, Dr.
Fred Baker (Kelsey!), vidi]. Dimensions (Lx H): 6.5 x 5.1 mm.

Remarks: This specimen is listed here as a possible syntype. The type lot
is USNM 73604 (Boss, et al., 1968), with type locality San Diego
Bay, 1-5 fms, in sandy mud. The ANSP source lot is marked "part
of type lot," although no reference is made to USNM (N. Gilmore,
ia itt., 1997).

Thraciidae. Now placed in the genus Asthenothaerus (E. V. Coan, pers.
comm.).

ecuadoriana, Crassinella [as new species]

Olsson, 1961: 182-183, pl. 25, figs. 6-6e.

1 paratype: DMNH 020775; Puerto Calle [published as Callo],
Ecuador. Coll. A. A. Olsson, 1958; ex ANSP 218934 (see
Remarks, vzdz7). Dimensions (Lx H): 3.0 x 2.5 mm.

Remarks: ANSP 218934 is the holotypic lot, as stated in the original
description; ANSP 218934a (vdz) includes four paratypic
specimens and is the source of this specimen. Olsson (1961: 4)
stated "the holotypes ... have been deposited at the Academy of
Natural Sciences in Philadelphia, and where additional material is
available, the paratypes of the same species will be placed in other
museum collections."

Crassatellidae.

enstfera, Yoldia [as n. sp.]

Wally 1897: 9, pln2, he. 4.

1 syntype: DMNH 020398; ALBATROSS Sta. 3133, Monterey Bay,
California. Ex ANSP 190960 (vidz). Dimensions (L x H): 20.0 x
9.6 mm.

2 syntypes:s DMNH 043602; ALBATROSS Sta. 3133, 37 fms,
Monterey Bay, California. Ex USNM. Dimensions (L x H): (1)
20.5 x 9-5 mm, (2) 15:8.x 8.9 mm:

Remarks: These specimens are listed here as possible syntypes, although
no specific type locality or depository was published in the original
description (or by Boss, et a/., 1968). The distributional range was
originally given as "Vancouver Island and Puget Sound, southward
to Monterey, California, in from 0 to 135 fathoms." ANSP 190960
(the source of DMNH 020398) is considered type material at that
institution. Dall's types were generally deposited in USNM (the
source of DMNH 043602), and although not listed by Boss, er al.
(1968), the USNM collection includes six syntypes (USNM

52 NEMOURIA No. 41

107644). Onginal dimensions were listed as 35 x 16 mm,
substantially larger than the DMNH specimens.

Sareptidae [none specified originally]. This is a synonym of Y.
(Cnesterium) seminuda Dall, 1871 (E. V. Coan, pers. comm.).

exoptata, Leda (Adrana) [as new species]

Pilsbry & Lowe, 1932: 107-108, pl. 17, figs. 8-9.

1 paratype: DMNH 020378; 20 fms, Guaymas, west Mexico. Coll. H.
N. Lowe, 1930; ex ANSP 155633. Dimensions (L x H): 14.7 x
4.7 mm.

Remarks: Pilsbry & Lowe's fig. 8 was specifically called "type" in the
legend, while fig. 9 was labelled "paratype." The ANSP source lot
is labelled "type and paratypes” and includes two vials, one with a
single specimen (= holotype) and one (not recataloged) containing
three valves (= paratypes).

Nuculanidae [originally Ledidae]. Curated as Nuculana in the DMNH
collection.

gabb1, Strigilla [as new species]

Olsson & McGinty, 1958: 50-51, pl. 5, figs. 3, 3a.

1 paratype: DMNH 021741; Atlantic coast of Costa Rica. Coll. W. M.
Gabb; ex ANSP 53379 (vdz). Dimensions (L x H): 35.1 x 31.3
mm.

Remarks: The DMNH specimen has "53379" penned on the inside of
one valve. The original description cited ANSP 53379 as
paratypic, containing one specimen of 35 x 30.5 mm. ANSP
53379 originally contained two specimens, identified as S. sincera
Hanley; it now contains two unmatched valves [plus a note
indicating that the published exterior view of the holotype was
from this lot; this actually belongs with the holotypic lot, ANSP
218881 from Colon, Panama (N. Gilmore, in litt., 1997)]. The
DMNH lot is a confirmed matched pair.

Tellinidae.

gerrardt1, Transenella [as new species]

Abbott, 1958: 130-131, text-fig. 7, pl. 4a-c.

2 paratypes: DMNH 021194; 8 fms, 3/4 mi southwest of Low Point,
Grand Cayman Island. Coll. A. J. Ostheimer; ex ANSP 199509
(Sta. D13-14, vid). Dimensions (L x H): (1) 9.4 x 7.3 mm, (2)
8.3 x 6.6 mm.

Remarks: Both live-collected with dried soft parts inside shells.

Veneridae [originally Veneridae: Meretricinae].

November 30, 1997 Mikkelsen & Bradford 33

hawaiensts, Solecardia [as n. sp.]

Pilsbry, 1921: 324-325, text-figs. 6a-c.

1 syntype: DMNH 021111; Hilo, Hawaii. Coll. D. Thaanum; ex ANSP
83102 (vid). Dimensions (L x H): 6.2 x 4.3 mm.

Remarks: No type material was specifically mentioned in the original
description, however, Pilsbry's types were generally deposited in
ANSP. Although the ANSP source lot is labelled "type +
paratypes," no [holo]type is segregated and no _ lectotype
designation has been published to our Knowledge, therefore ail
specimens must be considered syntypes.

Kelliidae [none specified originally]. Considered a junior synonym of
Nesobornia bartschi Chavan, 1969 [nomen novum pro Erycina
(Poronia) ovata Gould, 1850, non Gray, 1825, nec Philippi, 1836;
Chavan, 1969: N525] according to Kay (1979).

tmpar, Leda [as new species]

Pilsbry & Lowe, 1932: 106, pl. 17, figs. 3-6.

1 paratype: DMNH 020360; Guaymas, west Mexico. Coll. H. N. Lowe,
1930; ex ANSP 155636 (20 fms, vid7). Dimensions (L x H): 11.7
% Oe7 inim.

Remarks: Pilsbry & Lowe's figs. 3-5 (three views) were specifically
labelled "type," which constitutes a holotype designation; fig. 6 was
simply called "young shell." ‘The ANSP source lot is labelled "type
+ paratypes" and includes two vials, one with two valves
(presumably one specimen, the holotype) and another (not
recataloged) containing three specimens plus three valves (=
paratypes).

Nuculanidae [originally Ledidae]. Curated as Nuculana in the DMNH
collection.

inflatula, Macoma [as n. sp.]

Dall, 1897: 11-12, pl. 1, figs. 19-20.

4 syntypes (3 single right valves + 2 fragments of a fourth valve):
DMNH 043610; Captain's Bay, Unalaska Island, Alaska, 12 fms.
Ex USNM. Dimensions (Lx H): (1) 15.7 x 12.2 mm, (2) 13.6 x
9.9 mm, (3) 12.8 x 10.0 mm; fragments not measured.

Remarks: The original description (and Boss, et al., 1968) did not
specifically mention type material, a type locality, or a depository;
Boss, et al. (1968: 166) cited only the distributional range
“Aleutian Islands to Puget Sound.” Dall's types, however, were
generally deposited in USNM, the source of this material, and

34 NEMOURIA No. 41

indeed, that collection was found to include one syntype (USNM
107643, from 5 fms) labelled “figured type.” The DMNH
specimens are listed here as possible syntypes.

Tellinidae [none specified originally]. In the DMNH collection, this is
considered a junior synonym of Macoma frigida (Hanley, 1844).

Krebsitana, Corbula [no status specified in original description]

C. B. Adams, 1852: 234.

1 paralectotype: DMNH 021941; Jamaica. Coll. C. B. Adams; ex ANSP
297148 (vidi). Dimensions (L x H): 4.0 x 3.3 mm.

Remarks: This specimen is listed here as a possible paralectotype. The
original description mentioned no type material or depository for
the orginal "about 60 specimens." The ANSP source lot (now with
two specimens) bears a red dot (indicating type status) but type
designation is absent from labels. Clench & Turner (1950: 400)
figured a lectotype (MCZ 155611, wd); another lot in MCZ
(155612, wd, labelled “paratypes”) contains another 60+ valves,
which are thus paralectotypes.

Corbulidae [none specified originally].

lermond1, Pisidium [as n. sp.]

Sterki, 1913: 138-139.

22 syntypes ("part of type lot"): DMNH 139931; Duck Pond, Warren,
Maine. Coll. V. Sterki, October 1909; ex N. W. Lermond
(#3059). Dimensions (range): L = 2.3-3.2 mm, H = 2.2-2.6 mm.

145 syntypes (143 intact + 2 single unmatched valves): DMNH
160045; ditch in Ellis Stahl's field, Warren, Knox County, Maine.
Ex N. W. Lermond. Dimensions (range): L = 1.3-2.6 mm, H =
1.0-2.2 mm.

Remarks: Types were mentioned in the original description as "No.
6364, and in the collection of Mr. Lermond." This catalog number
probably referred to "the special collection of Sphaeriidae" (Sterki,
1912: 6) [the Sterki Sphaeriidae collection of ca. 15,000 lots is
now housed at the Carnegie Museum of Natural History
(Pittsburgh) (A. Bogan, in litt., 1997), but P. lermondi was not
listed in Carnegie's type catalog (Parodiz & Tripp, 1988)]. The
type locality ("Hab.") was given as "Duck Pond, Warren, Me.," but
the author also stated "at other places in the vicinity of Warren,
Me." Because of this ambiguity, DMNH 160045 (labelled cotypes
in the Lermond collection) is retained here as possible syntypes.
"Cotypes" also were recognized at MCZ (20082, ex N. W.
Lermond/V. Sterki; Johnson, 1959).

November 30, 1997 Mikkelsen & Bradford 35

Sphaertidae. This species is a junior synonym of Pisidium nitidum Jenyns,
1832 (fide Herrington, 1962).

maya, Condylonucula [as new species]

Moore, 1977: 124-125, figs. 4-5.

4 paratypes (3 intact + 1 as separated valves, in two paleoslides):
DMNH 120579; 2 m, Chancanab Lagoon, Cozumel, Mexico.
Coll. D. R. Moore, 29 November 1971; ex D. R. Moore.
Dimensions (range): L = 0.4-0.5 mm, H = 0.3-0.4 mm.

Remarks: The original description indicated lengths of the four paratypes
as 380, 490, 500, and 510 um.

Nuculidae.

mediafricana, Eupera [as new species]

Pilsbry & Bequaert, 1927: 355-357, text-figs. 79a-f, 80a.

1 paratype: DMNH 021012; Stanleyville, Belgian Congo [now
Kisangani, Haut-Zaire, Zaire], Africa. Coll. Lang & Chapin, 1915;
ex ANSP 133826 (coll. February 1915, donor AMNH 1924, wid).
Dimensions (Lx H): 4.0 x 2.9 mm.

Remarks: No type material was listed in original description. ‘The ANSP
lot is labelled "type and paratypes"; ANSP's source is recorded as
AMNH,, but the latter contains no type lot of this species (Boyko
& Sage, 1996; C. B. Boyko, pers. comm., 1997). According to
Boyko (pers. comm., 1997), the ANSP specimens most likely
originated in an AMNH expedition, not from the registered
AMNH collections themselves.

Sphaeriidae.

micella, Pleuromeris [as new species]

Olsson & McGinty, 1958: 45, pl. 5, figs. 7, 7a.

4 paratypes (single unmatched valves, 3 right + 1 left): DMNH 020822;
Bocas Island, northeastern Panama. Coll. Olsson and McGinty,
1953; ex ANSP 211907 (vdz). Dimensions (L x H): (1) 1.7 x 1.7
ini, (2) 1-6:'x 1.5 mm, (3) 1.5 x 1.6 mm, 4) 1.5 x 1.6 mm.

Remarks: The original description cited ANSP 211907 as the holotypic
lot and 211908 as the "paratype"; according to the ANSP

collection catalog, 211908 originally contained 35 specimens.
Carditidae.

montereyensis, Yoldia [as n. s.]
Dall, 1893: 29-30.
2 syntypes (1 intact + 1 single unmatched valve): DMNH 043608;

36 NEMOURIA No. 41

ALBATROSS Sta. 3202, 382 fms, 41°F, Monterey Bay,
California. Ex USNM. Dimensions (L x H): (1) 21.3 x 13.2 mm
(intact), (2) 28.2 x 16.3 mm (single unmatched valve).

Remarks: These specimens are from the original lot (ALBATROSS sta.
3202) and are ex USNM, the location of the known type material
(USNM 106972; Boss, et al., 1968).

Sareptidae [none specified originally]. Now placed in the subgenus
Megayoldia (fide E. V. Coan, pers. comm.).

mulawana, Tivela [as n. sp.]

Biggs, 1969: 205-206, figs. 1-2.

4 paratypes (single unmatched valves, 3 left, 1 right): DMNH 098136;
Masirah Island, South Arabia. Ex P. Cambridge. Dimensions
(range): L = 28.7-33.8 mm, H = 17.2-21.6 mm.

Remarks: The original description cited paratypes "in collection of Mr.
Philip Cambridge."

Veneridae.

nonuranus, Modiolus (Modiolus) [as n. sp.]

Pilsbry & Olsson, 1935: 16, pl. 1, fig. 3.

1 paratype: DMNH 020572; beach of Nonura Bay, near Punta Agua,
Peru. Coll. A. A. Olsson; ex ANSP 164613 (South Bayovar near
Punta Aguja, vidz). Dimensions (L x H): 27.4 x 15.0 mm.

Remarks: The onginal description listed the [holo]type as ANSP 164612
plus "paratypes in the Olsson collection"; the source lot (ANSP
164613) 1s listed in the ANSP collection catalog as containing 12
paratypes (valves), collected and donated by A. A. Olsson.

Mytilidae. Curated in the DMNH and ANSP collections as a junior
synonym of Semimytilus algosus (Gould, 1850).

nux, Egeria [as new species]

Pilsbry & Bequaert, 1927: 367-368, pl. 29, figs. 4-7.

1 paratype: DMNH 021488; Malela, Congo River, Belgian Congo [now
Zaire], Africa. Coll. H. Lang; ex ANSP 133889. Dimensions (L x
Pe 350x297 mim:

Remarks: The original description listed three specimens, consisting of
the type plus two paratypes. The ANSP source lot presently
consists of 66 valves labelled as paratypes (N. Gilmore, in litt.,
1997), and was listed as paratypic by Boyko & Sage (1996).

Donacidae. Curated in the genus Galatea in the DMNH collection.

November 30, 1997 Mikkelsen & Bradford eM |

omussa, Macrocallista (Chionella) [as new species]

Pilsbry & Lowe, 1932: 102, pl. 17, figs. 13-16.

1 paratype: DMNH 021196; San Juan del Sur, Nicaragua. Coll. H. N.
Lowe; ex ANSP 155597 (see Remarks, vid7). Dimensions (L x H):
6.1 X51 mini:

Remarks: Pilsbry & Lowe's (1932: pl. 17, figs. 13-16) figure legend
considered the specimen in figs. 13-14 as "type" and the remainder
"paratypes." The ANSP source lot (ANSP "155590," corrected on
the label to 155597, in the ANSP collection catalog as 155596) is
labelled "type + 2 paratypes." The collection catalog entry (ANSP
155596) indicates "type + cotypes." The lot now consists of two
vials, one with one specimen, and another (as paratypes, not
recataloged) containing two valves (one specimen?). The
[holo]type has in all these instances been segregated; the remaining
(ancluding the DMNH specimen) can be considered paratypes.

Veneridae. Curated as Transenella in the DMNH and ANSP collections.

pacis, Macoma [as new species]

Pilsbry & Lowe, 1932: 95, pl. 10, figs. 1, la, 2-3.

1 paratype: DMNH 021695; La Paz, Baja California Sur, Mexico. Coll.
H. N. Lowe, 1930; ex ANSP 152070 (vidz). Dimensions (L x H):
33.5 X 20.5 mim.

Remarks: The original description cited the ANSP source lot specifically
as paratypes. This specimen corresponds to the "long form"
illustrated in pl. 10, fig. 3.

Tellinidae. A junior synonym of Psammotreta viridotincta (Carpenter,
1856) according to Keen (1971).

panamensts, Cyrenoida [as new species]

Pilsbry & Zetek, 1931: 69, pl. 3, fig. 4.

1 paratype: DMNH 020969; near Punta Paitilla, close to Panama City,
Republic of Panama. Coll. J. Zetek; ex ANSP 155419 (wd1).
Dimensions (L x H): 17.8 x 16.5 mm.

Remarks: The type locality was given merely as "near Panama City."
The specimen is dirt-filled.

Cyrenoididae [none specified originally].

patagonicum, Pisidium [as sp. nov.]
Pilsbry, 1911: 607-608, pl. 47, figs. 8-10.
5 paratypes (3 intact + 2 single unmatched valves): DMNH 021008;
spring on Rio Chico, 25 mi [same on ANSP label; published as 15
mu] above Sierra Oveja, Patagonia, Argentina. Coll. J. B. Hatcher;

38 NEMOURIA No. 41

ex ANSP 88808. Dimensions (range): L = 5.3-6.8 mm, H = 4.4-
5.4 mm.

Remarks: These specimens are listed here as possible paratypes. No
specific type lot was cited in the original description, but Pilsbry's
types are generally considered to be at ANSP. The "type lot" at
ANSP is 88810 (two valves). There are nine other lots with similar
data, including ANSP 88807 and 88809 labeled as syntypes and
the source lot 88808 (10+ specimens) labeled "co-types" (N.
Gilmore, in litt., 1997).

Sphaeriidae. Curated in the DMNH collection in the genus Sphaerium
(Musculium); curated in the ANSP collection as Musculium.

pentston1, Cytherea [as new species (in title) ]

Heilprin, 1889: 142, pl. 8, fig. 4.

2 syntypes (single unmatched valves): DMNH 021176; Bermuda. Coll.
A. Heilpnin; ex ANSP 69644 (vdz). Dimensions (L x H): (1) 14.3
x 11.7 mam, (2) 12.9 11.4 mm.

Remarks: No type material or depository was designated in the original
description. The ANSP source lot is labelled "types."

Veneridae [none specified originally]. Curated as a junior synonym of
Pitar fulminatus (Menke, 1828) in the DMNH collection; curated
as Pitar (Pitar) penistoni in the ANSP collection.

platyptycha, Tapes [as n. sp.]

Pilsbry, 1901a: 206-207; 1901b: 400, pl. 19, fig. 6.

1 syntype: DMNH 021363; Hirado Island, Hizen, Japan. Coll. Y.
Hirase, 1901; ex ANSP 81218 ("1196," one segregated pair + two
pairs of valves, vid). Dimensions (L x H): 48.2 x 32.2 mm.

Remarks: No repository, catalog number, or original number of
specimens was provided in the original description, however, the
data match the stated type locality and source/collector. Pilsbry's
(1901b) subsequent text cited ANSP 81218 [also Hirase's
collection no. 1196].

Veneridae.

pontchartrainensis, Mulinia [as new species]
Morrison, 1965: 222-223, figs. 5-9.
6 paratypes: DMNH 108367; Middle Ground, eastern part of Lake
Pontchartrain, Louisiana. Coll. J. P. E. Morrison; ex USNM
635644. Dimensions (range): L = 5.4-7.4 mm, H = 3.9-5.1 mm.
Mactridae.

November 30, 1997 Mikkelsen & Bradford 39

redondoensts, Aligena [as sp. nov.]

T. Burch, 1941: 50-51, pl. 4, figs. 5-7.

1 paratype (right valve broken): DMNH 021105; Redondo Beach, Los
Angeles County, California, 75 fms. Coll. T. and J. Q. Burch,
1938; ex ANSP 178027 (#3833, coll. 7-31-38, vidz). Dimensions
Cox Be 1.9 x71. 7mm.

Remarks: The original description mentioned paratypes in several
repositories, including ANSP (without number). The source lot is
considered paratypic in the ANSP collection.

Thyasiridae [none specified originally]. Now placed in the genus
Axinodon (Coan & Scott, 1997).

redondoenstis, Nuculana [as new species]

J. Q. Burch, 1944: 9-10, fig.; 1945: 32 [as N. pendent redondoensis).

2 paratypes (single unmatched valves): DMNH 020353; off Redondo
Beach, California. Coll. B. R. Bales; ex ANSP 222225 (vidi).
Dimensions (L x H): (1) 6.0 x 3.5 mm (left valve), (2) 5.1 x 3.1
mm (right valve).

Remarks: These specimens are listed here as possible paratypes. Burch
(1945) stated that "a type specimen" had been deposited in the
USNM, and that "paratypes will be available for all western
museums wishing them." The source lot, ANSP 222225 (ex B. R.
Bales Coll. #16501, J. S. Schwengel, donor, 1958), is considered
a paratypic lot at that institution.

Nuculanidae. Considered a junior synonym of Nuculana (Saccella) hindsu
(Hanley, 1860) according to Keen (1971).

Reiniana, Corbicula [as n. sp.]

Clessin in Kuster, 1878: 196-197, pl. 39, figs. 8-9.

2 "paratypes" (1 intact + 1 single unmatched valve): DMNH 056392;
E-416, [Yokohama according to original description], Japan. Ex E.
Doremus. Dimensions (L x H): (1) 19.5 x 16.4 mm (single
unmatched valve), (2) 17.2 x 14.7 mm (intact, one valve broken).

Remarks: This lot was cited as paratypic by Counts (1991: 31). Both
specimens are smaller than the size range given in the original
description (length 27-32 mm, width 24-28 mm).

Corbiculidae [originally Cycladea].

rhypts, Pandora (Kennerlia) [as new species]
Pilsbry & Lowe, 1932: 105, pl. 16, figs. 8-11.
1 paratype: DMNH 021989; La Union, Gulf of Fonseca, San Salvador,
E] Salvador. Coll. H. N. Lowe; ex ANSP 155503a (coll. 1931,

40 NEMOURIA No. 41

vid1). Dimensions (L x H): 18.6 x 16.1 mm.

Remarks: Pilsbry & Lowe's original figures (figs. 8-11) were labelled
"type and paratypes." The ANSP source lot is the paratypic portion
of the onginal lot ANSP 155503, the latter number now applied to
the segregated holotype.

Pandoridae.

Rushi, Thracia [as n. sp.]

Pilsbry, 1897: 292-293, pl. 7, fig. 30.

1 syntype: DMNH 022252; Maldonado Bay, Uruguay. Coll. W. H.
Rush; ex ANSP 70502 ("types,” vidi). Dimensions (L x H): 18.6
x 12.3 mm,

Remarks: The original description gave length and height of three
specimens, without type designation or catalog number.

Thraciidae [none specified originally].

sablensis, Aequipecten (Plagioctenium) trradians [as ssp. n.]

A. H. Clarke, 1965: 175-176, pl. 1, figs. 1-6.

1 paratype (1 valve): DMNH 020650; Sable Island, Nova Scotia. Coll.
Mrs. F. Andrewschuk, July 1962; ex ANSP 290900 (ex National
Museum of Canada 14153, approx. 43.59° N, 60.00° W, widz).
Dimensions (L x H): 41.5 x 40.2 mm.

Remarks: The original description mentioned paratypes in USNM,
ANSP, and MCZ (without numbers). The ANSP source lot is
considered paratypic in that collection.

Pectinidae [none specified originally]. Curated in the genus Argopecten
in the ANSP collection.

sadoensts, Corbicula [as n. sp.]

Pilsbry, 1901la: 206-207; 1907: 158, pl. 7, figs. 15-16.

1 paralectotype: DMNH 020855; Sado Island, Japan. Coll. Y. Hirase,
1901; ex ANSP 89379 (vidi). Dimensions (L x H): 30.7 x 25.5
mm.

Remarks: No repository, catalog number, or original number of
specimens was provided for this species in the original description,
however, the data match the stated type locality and
source/collector. Counts (1991: 32) called ANSP 89379 (the
source of the DMNH specimen) the "holotype," resulting in a
lectotype designation [this lot presently contains one segregated
valve pair (= the lectotype) plus three additional sets (=
paralectotypes, although not mentioned by Counts, 1991)].
Counts (1991) called the DMNH specimen a paratype, but it is

November 30, 1997 Mikkelsen & Bradford 4]

actually a paralectotype.
Corbiculidae [originally Cyrenidae].

sororcula, Transenella [as new species]

Pilsbry & Lowe, 1932: 102, pl. 9, figs. 12-16.

1 figured paratype: DMNH 021190; San Juan del Sur, Nicaragua. Coll.
H. N. Lowe, 1931; ex ANSP 155599 (zidz). Dimensions (L x H):
o>. romee vetacy

Remarks: The legend to Pilsbry & Lowe's figs. 12-16 (as Macrocallista
sorocula [sic]) designated figs. 13 and 13a as "type," and the
remaining figured specimens as "paratypes." The source lot in the
ANSP collection was originally labelled "type + 9 paratypes"; a
single specimen (= holotype) 1s now segregated under the original
number, and the remaining specimens (two intact specimens and
11 valves) have been separated (but not recataloged) as paratypes.
The DMNH specimen can be considered a paratype. From the
color pattern, this specimen appears to be that illustrated by Pilsbry
& Lowe in fig. 12.

Veneridae. Curated as a junior synonym of T. modesta (Sowerby, 1835)
in the DMNH collection.

Sterkianum, Pisidium [as n. sp.]

Pilsbry, 1897: 291-292, pl. 6, figs. 1-4.

2 syntypes: DMNH 020994; creek in "Prado," Montevideo, Uruguay.
Coll. W. H. Rush; ex ANSP 70490 ("types," vidz). Dimensions (L
x HH): (14.5 x 3.6 mm, (2) 4.5 x 3.5 mim:

Remarks: "Many specimens were collected," but none were designated
specifically as types in the orginal description. This lot matches the
original locality and collector. "Cotypes" from this locality also
were recognized at MCZ (78076, ex B. Walker; Johnson, 1959).

Sphaeriidae [none specified originally].

sulfurea, Pitar [as n. sp.]

Pilsbry, 1904: 553, pl. 39, figs. 7-9.

1 syntype: DMNH 021167; Fukura, Awaji Island, Hyogo Prefecture,
Hyogo, Japan. Coll. Y. Hirase, 1901; ex ANSP 82135 (vid).
Dimensions (L x H): 25.8 x 22.8 mm.

Veneridae [none specified originally].

Swiftiana, Corbula [no status specified in original description]
C. B. Adams, 1852: 236-237.
1 paralectotype: DMNH 021936; east part of Kingston Harbor,

42 NEMOURIA No. 41

Kingston, Jamaica. Coll. C. B. Adams; ex Adams Collection via
MCZ via ANSP 182627 (3-4 fms, vidi). Dimensions (L x H): 7.7
xX 5.d omni,

Remarks: This specimen is listed here as a possible paralectotype. No
type material or depository were given in the original description
for the original “about 250 specimens’; the ANSP source lot is
considered type material in that collection. Clench & Turner
(1950: 400) figured a lectotype (MCZ 186103) and a
paralectotype (MCZ 155602, as “paratype”).

Corbulidae [none specified originally]. Considered a junior synonym of
C. caribaea Orbigny, 1846.

thaanum1, Cardium [as n. sp.]

Pilsbry, 1921: 323-324, pl. 12, fig. 24.

1 paratype: DMNH 020918; off Waikiki, Oahu Id., Hawani. Coll. D.
Thaanum, 1916; ex D. B. Kuhns via ANSP 47179 (35-50 fms,
vid1). Dimensions (L x H): 9.8 x 9.1 mm.

Remarks: No type material was designated in the original text
description, although the legend to the original figure (a
photograph) indicates "Type. 47179." This indication consititutes
a holotype designation. The ANSP source lot is now labelled "type
+ paratypes," and the DMNH specimen is from the paratypic
portion of this lot.

Cardiidae [none specified originally]. Now placed in the genus
Nemocardium (Kay, 1979; Moretzsohn & Kay, 1995).

troglodytes, Divariscintilla [as sp. nov.]

Mikkelsen & Bieler, 1989: 186-187, figs. 2, 4, 7, 12-15, 27.

3 paratypes (in alcohol): DMNH 175517; Sta. PMM-930, mud flat,
< 0.5 m by yabby pump from stomatopod burrow, 1100-1130 hrs,
Sebastian Inlet, Brevard County, Florida. Coll. P. M. Mikkelsen,
et al., 30 December 1987; ex P. M. Mikkelsen. Dimensions (L x
H; preserved mantle): (1) 5.1 x 3.9 mm [published as 4.9 mm], (2)
4.6 x 4.1 mm [published as 4.8 mm], (3) 4.5 x 3.9 mm.

Galeommatidae.

Uruguayenstis, Semele (Abra ?) [as n. sp.]
Pilsbry, 1897: 293, pl. 7, figs. 27-29.
2 syntypes (single left valves): DMNH 021617; Maldonado Bay,
Uruguay. Coll. W. H. Rush; ex ANSP 70493 (vidz). Dimensions
(Lx H):-@): 8.6 x 7.0 mm, (2)'7.4 x 6.5 mm.
Remarks: No type material or depository was designated in the original

November 30, 1997 Mikkelsen & Bradford 43

description although Maldonado Bay (in 3-6 fms) was given as the
original locality; original dimensions were listed as 9.5 x 8 mm.
The source ANSP lot is not labelled as a type lot but has a red dot
on the box indicative of same; it consists of one segregated
specimen plus others. In the absence of a formal lectotype
designation, all specimens must continue to be considered as
syntypes.

Semelidae [none specified originally]. Curated in the DMNH collection
in the genus Abra.

viola, Corbicula [as n. sp.]

Pilsbry, 1907: 158, pl. 7, figs. 7-10.

1 syntype: DMNH 020858; Lake Biwa, Japan. Coll. T. Isakawa, 1905;
ex ANSP 89371 (see Remarks, vidz7). Dimensions (L x H): 24.0 x
2159 mm: |

Remarks: No type specimens were listed specifically in the original
description. Both DMNH and ANSP lots were mentioned and
called syntypic by Counts (1991: 38). In the ANSP collection, lot
89371 1s now labelled as "holotype"; six valves were removed and
recataloged as "paratype" lot 358423. This does not here constitute
a lectotype designation.

Corbiculidae [none specified originally].

vulgaris, Leda [as n. sp.]

Brown & Pilsbry, 1913: 499, fig. 3.

232 syntypes (44 intact + 188 single unmatched valves): DMNH
135065; Pleistocene, Mt. Hope, Panama Canal Zone. Coll. W. B.
Scott; ex J. Parker. Dimensions (range): L = 3.8-12.0 mm, H =
2.3-6.2 mm.

Remarks: These specimens are listed here as possible syntypes. No type
material or repository was mentioned in the original description.

Nuculanidae [none specified originally]. Curated as Nuculana in the
DMNH collection.

webbianus, Unio (Elliptio) [as sp. nov.]
Wright, 1934: 94-95, pl. 10, figs. 1-2.
1 paratype: DMNH 150106; Citrus County, Lake Consuelo, Florida.
Ex W. F. Webb via M. L. Walton (#4195). Dimensions (L x H):
42.2 x 27.4 mm.
Unionidae [none specified originally].

44 NEMOURIA No. 41

yokohamensts, Raeta [as n. sp.]
Pilsbry, 1895: 119, pl. 3, figs. 4-5.
1 syntype: DMNH 021479; Yokohama, Kanagawa, Honshu Island,
Japan. Coll. F. Stearns, 1895; ex ANSP 70873 (zd). Dimensions
(Lx AH): 16.7 x 12.3:
Remarks: The ANSP source lot is labelled "types" in that collection.
Mactridae.

yoyo, Divariscintilla [as sp. nov.]

Mikkelsen & Bieler, 1989: 177-186, figs. 1, 3, 5-6, 8-11, 26.

2 paratypes (in alcohol): DMNH 175516; Sta. PMM-924, sand flat,
< 0.5 m, by yabby pump from stomatopod burrow, Ft. Pierce
Inlet, St. Lucie County, Florida. Coll. P. M. Mikkelsen, et al., 31
August 1987; ex P. M. Mikkelsen. Dimensions (L x H, preserved
mantle): (1) 4.8 x 2.5 mm [published as 5.7 mm], (2) 3.7 x 2.8
mm [published as 4.7 mm].

Galeommatidae.

SCAPHOPODA

teliger, Cadulus [as ""n. sp." for Cadulus spretus Suter non Tate & May]

C. J. Finlay, 1927: 444, figs. 52-54.

1 paratype: DMNH 022370; north of Auckland Islands, New Zealand
Subantarctic Islands. Ex A. W. B. Powell via ANSP 232249 (10 mi
N of Auckland Island, 86677, 51° S, 168° E, 95 fms, wd).
Dimensions (L x W): 4.3 x 0.7 mm.

Remarks: This specimen is listed here as a possible paratype. The
original description stated that "types of all new species described
are in the Finlay collection; paratypes wherever possible have been
deposited in the Australian Museum, Sydney" (Finlay, 1927: 330).
This would seem to exclude these specimens from type status,
however, the ANSP source lot was listed as paratypic by Spamer
& Bogan (1992).

Gadilidae [none specified originally]. This species is curated as the genus
Gadila in the ANSP collection.

November 30, 1997 Mikkelsen & Bradford 45

ALPHABETIC INDEX BY AUTHOR
(PARTS I AND II: BIELER & BRADFORD, 1991, AND THIS PAPER)
Families reflect current placement.

Abbott, R. T.
arcas, Terebra (Strioterebrum), Terebridae (Part I)
caribaea, Coralhophila, Muricidae
eloiseae, Acteon, Acteonidae (Part I)
gerrardi, Transenella, Veneridae
habet, Assiminea habet, Assimineidae (Part I)
hoffmeyert, Terebra (Strioterebrum), Terebridae (Part I)
patae, Conus, Conidae (Part I)
stegert, Terebra (Myurella) floridana, Terebridae (Part I)

Abbott, R. T. & C. J. Finlay

cosmani, Chicoreus, Muricidae (Part I)

Abbott, R. T. & H. Lewis
boschi, Cymatium, Ranellidae (Part I)

Adams, C. B.
Krebsiana, Corbula, Corbulidae
scalaroides, Rissoa, Rissoidae (Part I)
Swiftiana, Corbula, Corbulidae

Aguayo, C. G. & M. L. Jaume

jaumei, Eutrochatella (Troschelviana), Helicinidae

Alcalde y Ledon, O. (see also under Jaume)
fuentesi1, Chondrothyrium crenimargo, Annulariidae
jaumet, Chrondrothynum violaceum, Annulariidae
serranum, Farcimen (Farcimen) auriculatum, Megalomastomidae

Baker, F.

scherert, Helicina, Helicinidae

Bales, B. R.

nanus, Strombus raninus, Strombidae (Part I and this paper)

Barnard, K. H.
boswellae, Pirenella, Potamididae (Part I)

46 NEMOURIA

Bartsch, P.
smithi, Prodalha, Volutidae (Part I)
tagbilleranus, Geophorus (Diplopinax), Helicinidae
yoshidai, Katayama nosophora, Pomatiopsidae

Bayer, F. M.
aurora, Columbarium (Peristarium), Turbinellidae (Part I)
electra, Columbarium (Peristarium), Turbinellidae (Part I)
merope, Columbarium (Peristarium), Turbinellidae (Part I)

Bequaert, J. C. (see also under Pilsbry)
veracruzensis, Cerithidea pliculosa, Cerithideidae (Part I)

Bieler, R. (see under Mikkelsen)

Biggs, H. E. J.

mulawana, Tivela, Veneridae
Bratcher, T. (see under Cernohorsky)

Bratcher, T. & R. D. Burch
allym, Terebra, Terebridae (Part I)
dorothyae, Terebra, Terebridae (Part I)
hancocki, Terebra, Terebridae (Part I)

Brown, A. P. & H. A. Pilsbry
vulgaris, Leda, Nuculanidae

Burch, J. Q.

redondoensis, Nuculana, Nuculanidae

Burch, J. Q. & R. L. Burch
rejecta, Oliva, Olividae (Part I)

Burch, R. D. (see also under Bratcher)
betsyae, Hastula (Punctoterebra), Terebridae (Part I)
crake1, Duplicaria (Duplicaria), Terebridae (Part I)
maryleeae, Hastula (Hastula), Terebridae (Part I)

Burch, R. L. (see under J. Q. Burch)

No. 41

November 30, 1997 Mikkelsen & Bradford

Burch, T.
redondoensis, Aligena, Thyasiridae

Burgess, C. M.
mauiensis, Cypraea, Cypraeidae (Part I)

Campbell, G. B. (see under Shasky)

Cardin, C. & J. G. Walls

lenoreae, Phenacovolva (Subsimnia), Ovulidae (Part I)

Cate, C.N.
eugeniae, Notadusta, Cypraeidae
lahainaensis, Volva (Phenacovolva), Ovulidae (Part I)

Cate, J. M.

filiareginae, Vexillum regina, Costellariidae

Cernohorsky, W. O. (see also under Emerson)
adamsianum, Vexillum (Costellaria), Costellariidae (Part I)
backae, Scabricola (Scabricola), Mitridae
clovert, Cancilla (Ziba), Mitridae
crosniert, Cancilla scrobiculata, Mitridae (Part I and this paper)
dondam, Mitra, Mitridae
earlet, Mitra (Nebularia), Mitridae
ekerae, Scabricola (Swainsonia) ocellata, Mitridae
hansenae, Vexillum (Pusia), Costellaridae
marrow, Vexillum (Pusia), Costellaridae
martinorum, Vexillum (Costellaria), Costellariidae
nodospiculum, Vexillum (Costellaria), Costellariidae
orrt, Muricopsis, Muricidae
salisbury, Mitrolumna, Turridae (Part I)
salisburyi, Vexillum (Pusta), Costellariidae (Part I and this paper)
vicdam, Scabricola, Mitridae
wolfe1, Vexillum (Costellaria), Costellariidae (Part I and this paper)

Cernohorsky, W. O. & T. Bratcher
parkinsoni, Terebra, Terebridae (Part I)

Clarke, A. H., Jr.

sablensis, Aequipecten (Plagioctenium) irradians, Pectinidae

47

48 NEMOURIA No. 41

Clench, W. J.
finlayn, Murex (Murex), Muricidae (Part I)
haysae, Thats floridana, Muricidae (Part I)
trujilloi, Oliva, Olividae (Part I)

Clench, W. J. & R. D. Turner
johnstonet, Melongena corona, Melongenidae (Part I, as Buccinidae)
langi, Thats (Thaisella), Muricidae (Part I)
riost, Adelomelon (Weaveria), Volutidae (Part I)

Clessin, S. in H. C. Kiister
Reiniana, Corbicula, Corbiculidae

Coelho, A. C. dos Santos (see under Matthews)

Cotton, B. C.
rosseli, Zoila, Cypraeidae (Part I)

Dall, W. H.
atkana, Littorina sitkana, Littorinidae (Part I)
brunnea, Tindaria, Tindaridae
diegensis, Limopsis, Limopsidae
diegensis, Thracia, Thraciidae
ensifera, Yoldia, Sareptidae
inflatula, Macoma, Tellinidae
major, Buccinum pemphigus, Buccinidae (Part I)
montereyensis, Yoldia, Sareptidae
vernicosum, Mangilia (Calhotectum), Volutidae (Part I, as Turridae)

Dall, W. H. zn C. R. Orcutt & W. H. Dall
orcutti, Caecum, Caecidae (Part I)

D'Attilio, A. (see under Emerson)

DuShane, H.
textimattum, Epitonium (Asperiscala), Epitoniidae (Part I)

Emerson, W. K. & W. O. Cernohorsky
hadart, Drupa ricinus, Muricidae

Emerson, W. K. & A. D'Attilio
dalh, Latiaxis (Babelomurex), Muricidae (Part I)

November 30, 1997 Mikkelsen & Bradford

guamensis, Favartia, Muricidae
mactanensis, Murexiella, Muricidae

Emerson, W. K. & B. J. Piech
jenniernestae, Distorsio (Distorsio), Personidae

Ferreira, A. J.
portobelensis, Callistochiton, Ischnochitonidae

Finlay, C. J. (see also under Abbott)
teliger, Cadulus, Gadilidae

Finlay, H. J.

cuvierensis, Microvoluta, Volutomitridae (Part I)

Ford, J.
blanest, Ohivella, Olividae (Part I)
gaylordi, Olivella gracilis, Olividae (Part I)
greegori, Cypraea cruenta, Cypraeidae (Part I)
michaeli, Ocinebra, Muricidae (Part I)

Harasewych, M. G.
ecphoroides, Fulgurofusus (Fulgurofusus), ‘Vurbinellidae (Part I)
germonae, Tractolira, Volutidae (Part I)
harrisae, Columbarium, Turbinellidae (Part I)
timor, Fulgurofusus (Peristarium), Turbinellidae (Part I)
xenismatis, Fulgurofusus (Histricosceptrum), Turbinellidae (Part I)

Harasewych, M. G. & R. H. Jensen
guesti, Natica (Natica), Naticidae (Part I)
guesti, Pterynotus (Pterynotus), Muricidae (Part I)
hghtbournt, Pterynotus (Pterynotus), Muricidae (Part I)
radwini, Pterynotus (Pterynotus), Muricidae (Part I)

Hedley, C.
perplicata, Voluta, Volutidae (Part I)

Heilprin, A.
penistoni, Cytherea, Veneridae

Henderson, J. B., Jr. & C. T. Simpson
superbum, Chondropoma, Annulariidae

50 NEM OURIA No. 41

Hoerle, R. C.
finlayi, Neritopsis, Neritopsidae (Part I)

Howard, F. B.
aureola, Pyrene = aureomexicana, Pyrene, Columbellidae (Part I and
this paper)

Hutton, F. W.

corticata, Acmaea, Lottiidae (Part I, as Acmaeidae)

Iredale, T.
kellnert, Aulica, Volutidae (Part I)

Jaume, M. L. (see under Aguayo)

Jaume, M. L. & O. Alcalde y Ledon
fuentest, Chondropoma (Chondropoma), Annularidae

Jaume, M. L. & L. Sanchez de Fuentes
alcaldet, Chondropoma (Chondropoma), Annulariidae
alcaldet, Chondrothyrium (Plicathyrium), Annulariidae
britot, Eutudora limbifera, Annulariidae
morenoi, Chondrothyrium (Chondrothyrium) violaceum, Annulariidae
torret, Chondrothyrium (Phcathyrium), Annularidae

Jensen, R. H. (see under Harasewych)

Keen, A. M.
fayae, Anachis (?Costoanachis), Columbellidae (Part I)

Klappenbach, M. A.
teaguet, Olivancillaria, Olividae (Part I)

Krieger, K. A.
alcoviensis, Somatogyrus, Hydrobiidae

Lewis, H. (see also under Abbott)
burgessi, Distorsio, Personidae (Part I, as Ranellidae)
habet, Distorsio constricta, Personidae (Part I, as Ranellidae)

Little, M. D. (see under Malek)

November 30, 1997 Mikkelsen & Bradford
Lowe, H. N. (see under Pilsbry)

Lyons, W. G.
barbarae, Brachycythara, Turridae (Part I)
girardi, Cerodrillia, ‘Turridae (Part I)
inglet, Oceanida, Eulimidae (Part I)
lineata, Acanthochitona, Acanthochitonidae

Malek, E. A. & M. D. Little
colombiensis, Aroapyrgus, Hydrobiidae

Marcus, Ernst (see under Marcus, Eveline)

Marcus, Eveline & Ernst Marcus
vina, Assiminea succinea, Assimineidae (Part I)

Matthews, H.R. & A. C. dos Santos Coelho
lopest, Mitra (Cancilla), Mitridae

McGinty, T. L. (see also under Olsson)
stegert, Daphnella, ‘Yurridae (Part I)
thompsont, Eudolium, ‘Yonnidae (Part I)

Melvill, J. C.

jacksoni, Scissurella, Scissurellidae

Mikkelsen, P. M. & R. Bieler
troglodytes, Divariscintilla, Galeommatidae
yoyo, Divanscintilla, Galeommatidae

Mitchell-Tapping, H. J.

donmooret, Caecum, Caecidae (Part I)

Moore, D. R.
calcicola, Nucula, Nuculidae
cynthiae, Condylonucula, Nuculidae
maya, Condylonucula, Nuculidae

Morrison, J. P. E.
barretti, Odostomiae = booneae, Hydrobia, Hydrobiidae
chocolatum, Incidostoma, Poteriidae
diminutum, Incidostoma, Poteriidae

52 NEM OURIA No. 41

jackson, Incidostoma, Poteriidae
louisianae, Vioscalba, Hydrobiidae
pontchartrainensis, Mulinia, Mactridae
weber, Odostomia, Pyramidellidae

Mulliner, D. K. & G. G. Sphon
carolynae, Platydoris, Platydorididae

Oliver, W. R. B.

kermadecensis, Haurakia, Rissoidae (Part I)

Olsson, A. A. (see also under Pilsbry)
ecuadoniana, Crassinella, Crassatellidae
stegert, Olivella (Olivella), Olividae (Part I)

Olsson, A. A. & T. L. McGinty
elegans, Vitrinella (Stnovitrinella), Vitrinellidae (Part I)
gabbi, Strigilla, Tellinidae
marmosa, Ohivella minuta, Olividae (Part I)
micella, Pleuromens, Carditidae
toroensis, Rissoa, Rissoidae

Orcutt, C. R. (see under Dall)

Pease, W. H.
dispar, Neritina, Neritidae (Part I)
rubra, Alcyna, Trochidae (Part I)

Peile, A. J.
bermudensis, Rissoina, Rissoidae (Part I)

Petit, R. E. (see also under Roth)
darwint, Cancellaria, Cancellariidae (Part I)

Petuch, E. J.
abrolhosensis, Acmaea (Collisella), Lottiidae (Part I, as Acmaeidae)
aphrodite, Conus, Conidae (Part I)
baileyi, Oliva, Olividae (Part I)
boholensis, Conus, Conidae (Part I)
eccentricus, Zelippistes, Capulidae (Part I)
echinophilus, Africonus, Conidae (Part I)
exquisita, Pseudocypraea, Ovulidae (Part I)

November 30, 1997 Mikkelsen & Bradford

fragilissimus, Conus, Conidae (Part I)

kaicherae, Vexillum (Costellaria), Costellariidae (Part I)
kurzi, Guildfordia, Turbinidae (Part I)

kurzt, Morum (Oniscidia), Harpidae (Part I)

lightbourni, Conus (Floraconus), Conidae (Part I)

nereis, Conus, Conidae (Part I)

nodosivaricosa, Agatnix (Olssonella), Cancellaridae (Part I)
philippinensis, Axymene, Muricidae (Part I)

zulu, Conus, Conidae (Part I)

Piech, B. J. (see under Emerson)

Pilsbry, H. A. (see also under Brown)
awajiensis, Corbicula, Corbiculidae
bryanae, Congeria, Mytilidae
dalhana, Acmaea, Lottiidae (Part I, as Acmaeidae)
floridana, Stenacme, Epitoniidae (Part I)
hawaiensis, Solecardia, Kelliidae
patagonicum, Pisidium, Sphaeriidae
platyptycha, Tapes, Veneridae
Rushit, Thracia, Vhraciidae
sadoensis, Corbicula, Corbiculidae
Sterkianum, Pisidium, Sphaeridae
sulfurea, Pitar, Veneridae
thaanumi, Cardium, Cardiidae
tokyoensis, Rissoa, Rissoidae (Part I)
Uruguayensis, Semele (Abra?), Semelidae
viola, Corbicula, Corbiculidae
yokohamensis, Raeta, Mactridae

Pilsbry, H. A. & J. C. Bequaert
congo, Iphigenia, Donacidae
mediafnicana, Eupera, Sphaeriidae
nux, Egeria, Donacidae

Pilsbry, H. A. & H. N. Lowe
acapulcensis, Leda (Saccella), Nuculanidae
exoptata, Leda (Adrana), Nuculanidae
hilh, Anachis (Glyptanachis), Columbellidae
impar, Leda, Nuculanidae
omissa, Macrocallista (Chionella), Veneridae
pacis, Macoma, Tellinidae

54 NEM OURIA

rhypis, Pandora (Kennerlia), Pandoridae
sororcula, Transenella, Veneridae

Pilsbry, H. A. & A. A. Olsson
nonuranus, Modiolus (Modiolus), Mytilidae

Pilsbry, H. A. & J. Zetek
panamensis, Cyrenoida, Cyrenoididae

Powell, A. W. B.
ahiparana, Turritella (Zeacolpus), Turritellidae (Part I)
bouvetia, Margarella, Trochidae (Part I)
delh, Penion benthicola, Buccinidae (Part I)
kingae, Xenoturris, Turridae (Part I)
macquaniensis, Trophon, Muricidae (Part I)
mauganuica, Austrodiaphana, Diaphanidae (Part I)
mawsoni, Parmaphorella, Fissurellidae
nivosa, Ovirissoa, Rissoidae (Part I)
pseudanaloga, Puncturella, Fissurellidae (Part I)
queenslandica, Leucosyrinx, Turridae (Part I)
scotiana, Sinuber sculpta, Naticidae (Part I)
signeyana, Chlanidota, Buccinidae (Part I)
tenerum, Parvaplustrum, Aplustridae (Part I, as Hydatinidae)
venustula, Pareuthria, Buccinidae (Part I)
weaverl, Turridrupa, Turridae (Part I)
wormaldi, Columbarium (Coluzea), Turbinellidae (Part I)
zebrina, Zediloma (Fractarmilla) corrosa, Trochidae (Part I)

Prime, T.
angas!, Corbicula, Corbiculidae

Quinn, J. F., Jr.
acanthus, Mirachelus, Trochidae
bermudense, Calliostoma, Trochidae
carinatus, Lamellitrochus, ‘Vrochidae
guesti, Calliostoma, Trochidae

Rehder, H. A.
barnardi, Fusivoluta, Volutidae (Part I)
boswellae, Volutocorbis, Volutidae (Part I)
clarkei, Fusivoluta, Volutidae (Part I)
disparilis, Volutocorbis, Volutidae (Part I)

No. 41

November 30, 1997 Mikkelsen & Bradford a5

floridanus, Microdochus, Elachisinidae (Part I)
gracihor, Benthovoluta, Turbinellidae (Part I)

Richards, H. G.
edgerlyi, Gyrineum perca, Ranellidae
inspinata, Melongena corona, Melongenidae

Roth, B. & R. E. Petit
mastirana, Persicula, Marginellidae (Part I)

Sanchez de Fuentes, L. (see under Jaume)

Schilder, F. A.
summersi, Bistolida (Blasicrura) palhdula, Cypraeidae (Part I)

Schilder, F. A. & R. Summers
casta, Notocypraea, Cypraeidae
hesperina, Blasicrura coxent, Cypraeidae (Part I)

Schwengel, J. S.
koto, Lamellaria, Lamellariidae (Part I)
meginty1, Eubela, Turridae (Part I)

Shasky, D. R.
berry1, Anachis, Columbellidae (Part I)
maclean, Coralliophila, Muricidae (Part I)
mazatlanica, Ruthia, Columbellidae (Part I)
mendozana, Strombina (Cotonopsts), Columbellidae (Part I)

Shasky, D. R. & G. B. Campbell
macleant, Calliostoma, Trochidae (Part I)
sphon, Mitra (Strigatella), Costellariidae

Simpson, C. T. (see under Henderson)

Smith, A. G.
stilest, Neptunea, Buccinidae (Part I)

Smith, E. A.
huttonu, Trochus (Cantharidus), Trochidae (Part I)

56 NEMOURIA No. 41

Snyder, M. A.
lightbourn, Fusinus, Fasciolaridae (Part I, as Buccinidae)
martini, Latirus, Fasciolariudae (Part I, Buccinidae)

Sphon, G. (see under Mulliner)

Sterki, V.
lermondi, Pisidium, Sphaeriidae

Stokes, A. J.
randalh, Cymbiola, Volutidae (Part I)

Summers, R. (see under Schilder)

Suter, H.
lyalliana, Rissota (Alvania) plicata, Rissoidae (Part I)

Test, A. R.
cona, Acmaea, Lottiidae (Part I, as Acmaeidae)

Thompson, F. G.
rhecta, Meganipha, Annulariidae

Tippett, D. L.
brasiliana, Borsonia, Turridae (Part I)
petiti, Fenimorea, Turridae
wolfet, Drillia, Turridae

Tucker, J. K.
sprucecreekensis, Melongena, Melongenidae

Turner, R. D. (see under Clench)

Vanatta, E. G.
amosi, Metula, Fasciolariidae (Part I, as Buccinidae)

Waller, T. R.

cranmerorum, Somalipecten, Pectinidae

Walls, J. G. (see also under Cardin)
axelrodi, Conus, Conidae (Part I)
clovert, Conus, Conidae (Part I)

November 30, 1997 Mikkelsen & Bradford 57

kerstitchi, Conus, Conidae (Part I)

meyert, Conus biliosus, Conidae (Part I)
parvatus, Conus musicus, Conidae (Part I)
reductaspiralis, Conus nielsenae, Conidae (Part I)
sandwichensis, Conus suturalis, Conidae (Part I)
tribblei, Conus, Conidae (Part I)

wittingt, Conus, Conidae (Part I)

Watters, G. T.
andersont, Acanthochitona, Acanthochitonidae

Weaver, C. S.
cloveriana, Lyria, Volutidae (Part I)
daisyae, Volutoconus hargreavest, Volutidae (Part I)
dampieria, Amoria (Amoria), Volutidae (Part I)
dupont, Festilyria, Volutidae (Part I)

Wheat, S. C.
fergusont Wheat, 1913, Acmaea, Lottiidae

Wright, B. H.
webbianus, Unio (Elhiptio), Unionidae

Zetek, J. (see under Pilsbry)

TAXONOMIC INDEX
(PARTS I AND II: BIELER & BRADFORD, 1991, AND THIS PAPER)
(numbers of lots/specimens in parentheses; * = includes holotype)
Listed by original epithet within current families.

POLYPLACOPHORA:
Acanthochitonidae
andersoni Watters, 1981, Acanthochitona (1/1)
lineata Lyons, 1988, Acanthochitona (1/1)
Ischnochitonidae
portobelensis Ferreira, 1976, Callistochiton (1/1)

GASTROPODA: PROSOBRANCHS:
Acmaeidae - see Lottiidae
Annulariidae
alcalde: Jaume & Sanchez de Fuentes, 1943, Chondropoma

58 NEMOURIA No. 41

(Chondropoma) (1/4)
alcaldei Jaume & Sanchez de Fuentes, 1943, Chondrothyrium
(Phicathyrium) (1/1)
britoi Jaume & Sanchez de Fuentes, 1943, Eutudora limbifera (1/6)
fuentest Jaume & Alcalde y Ledon, 1944, Chondropoma (Chondropoma)
Gly)
fuentest Alcalde y Ledon, 1943, Chondrothyrium crenimargo (1/1)
jaumet Alcalde y Ledon, 1943, Chrondrothyrium violaceum (1/4)
morenoi Jaume & Sanchez de Fuentes, 1943, Chondrothyrium
(Chondrothyrium) violaceum (1/2)
rhecta Thompson, 1978, Meganipha (1/3)
superbum Henderson & Simpson, 1902, Chondropoma (1/4)
torre: Jaume & Sanchez de Fuentes, 1943, Chondrothyrium
(Phicathyrium) (1/1)
Assimineidae
habet Abbott, 1958, Assitminea habet, Part I (1/2)
vina Marcus & Marcus, 1965, Assiminea succinea, Part I (1/31)
Buccinidae
deli Powell, 1971, Penion benthicola, Part I (1/1)
major Dall, 1919, Buccinum pemphigus, Part I (1/1)
signeyana Powell, 1951, Chlanidota, Part I (1/1)
stilest Smith, 1968, Neptunea, Part I (1/1)
venustula Powell, 1951, Pareuthria, Part I (1/1)
Caecidae
donmooret Mitchell-Tapping, 1979, Caecum, Part I (*1/1)
orcutti Dall in Orcutt & Dall, 1885, Caecum, Part I (1/2)
Cancellariidae
darwint Petit, 1970, Cancellaria, Part I (1/1)
nodosivaricosa Petuch, 1979, Agatrix (Olssonella), Part I (*1/1)
Capulidae
eccentricus Petuch, 1979, Zelippistes, Part I (*1/1)
Cerithideidae
veracruzensis Bequaert, 1942, Centhidea pliculosa, Part I (1/1)
Columbellidae
aureola Howard, 1963, Pyrene = aureomexicana Howard, 1963, Pyrene,
Part I. (1/1)
berryt Shasky, 1970, Anachis, Part I (1/2)
fayae Keen, 1971, Anachis (2?Costoanachis), Part I (1/2)
hill Pilsbry & Lowe, 1932, Anachis (Glyptanachis) (1/2)
mazatlanica Shasky, 1970, Ruthia, Part I (1/1)
mendozana Shasky, 1970, Strombina (Cotonopsis), Part I (1/1)

November 30, 1997 Mikkelsen & Bradford 59

Conidae
aphrodite Petuch, 1979, Conus, Part I (*2/2)
axelrodi Walls, 1978, Conus, Part I (*1/1)
boholensis Petuch, 1979, Conus, Part I (*2/3)
clovert Walls, 1978, Conus, Part I (*1/1)
echinophilus Petuch, 1975, Africonus, Part I (1/1)
fragilissimus Petuch, 1979, Conus, Part I (*2/4)
kerstitchi Walls, 1978, Conus, Part I (*1/1)
lightbourni Petuch, 1986, Conus (Floraconus), Part I (*2/4)
meyert Walls, 1979, Conus biliosus, Part I (*1/1)
nerets Petuch, 1979, Conus, Part I (*2/2)
parvatus Walls, 1979, Conus musicus, Part I (*2/7)
patae Abbott, 1971, Conus, Part I (*4/7)
reductaspiralis Walls, 1979, Conus nielsenae, Part I (*1/1)
sandwichensis Walls, 1978, Conus suturalis, Part I (*1/1)
tribble: Walls, 1977, Conus, Part I (*1/1)
wittigs Walls, 1977, Conus, Part I (*1/1)
zulu Petuch, 1979, Conus, Part I (*3/6)
Costellariidae
adamsianum Cernohorsky, 1978, Vexillum (Costellaria), Part I (1/1)
filiareginae Cate, 1961, Vexillum regina (2/2)
hansenae Cernohorsky, 1973, Vexillum (Pusia) (2/3)
kaicherae Petuch, 1979, Vexillum (Costellaria), Part I (1/2)
marrow Cernohorsky, 1973, Vexillum (Pusia) (4/5)
martinorum Cernohorsky, 1986, Vexillum (Costellaria) (1/1)
nodospiculum Cernohorsky, 1970, Vexillum (Costellaria) (2/2)
salisburyt Cernohorsky, 1976, Vexillum (Pusia), Part I (1/3) and this
paper (2/3)
sphonit Shasky & Campbell, 1964, Mitra (Strigatella) (1/1)
wolfet Cernohorsky, 1978, Vexillum (Costellaria), Part I (1/1) and this
paper (1/1)
Cypraeidae
casta Schilder & Summers, 1963, Notocypraea (1/1)
eugeniae Cate, 1975, Notadusta (2/2)
greegori Ford, 1893, Cypraea cruenta, Part I (1/1)
hesperina Schilder & Summers, 1963, Blasicrura coxeni, Part I (1/1)
mauiensis Burgess, 1967, Cypraea, Part I (1/2)
rosseli Cotton, 1948, Zoila, Part I (1/1)
summerst Schilder, 1958, Bistolida (Blasicrura) pallidula, Part I (1/1)
Elachisinidae
floridanus Rehder, 1943, Microdochus, Part I (1/78)

60 NEMOURIA No. 41

Epitoniidae
floridana Pilsbry, 1945, Stenacme, Part I (1/1)
textimattum DuShane, 1977, Epitonium (Asperiscala), Part I (1/1)
Eulimidae
ingle: Lyons, 1978, Oceanida, Part I (1/1)
Fasciolariidae
amost Vanatta, 1913, Merula, Part I (1/2)
lightbournt Snyder, 1984, Fusinus, Part I (*2/4)
martini Snyder, 1988, Latirus, Part I (1/1)
Fissurellidae
mawsont Powell, 1958, Parmaphorella (1/1)
pseudanaloga Powell, 1957, Puncturella, Part I (2/5)
Harpidae
kurzi Petuch, 1979, Morum (Oniscidia), Part I (*1/1)
Helicinidae
jaumet Aguayo & Jaume, 1957, Eutrochatella (Troscheluviana) (1/4)
scherert Baker, 1913, Helicina (1/4)
tagbilleranus Bartsch, 1918, Geophorus (Diplopinax) (1/6)
Hydrobiidae
alcoviensis Krieger, 1972, Somatogyrus (2/24)
barrett: Morrison, 1965, Odostomia = booneae Morrison, 1973,
Hydrobia (1/2)
colombiensis Malek & Little, 1971, Aroapyrgus (*2/92)
louisianae Morrison, 1965, Vioscalba (1/23)
Lamellariidae
koto Schwengel, 1944, Lamellaria, Part I (2/2)
Littorinidae
atkana Dall, 1886, Littorina sitkana, Part I (1/2)
Lottiidae
abrolhosensis Petuch, 1979, Acmaea (Collisella), Part I (1/4)
cona Test, 1945, Acmaea, Part I (1/1)
corticata Hutton, 1880, Acmaea, Part I (1/1)
dalhana Pilsbry, 1891, Acmaea, Part I (1/1)
fergusont Wheat, 1913, Acmaea (1/1)
Marginellidae
masirana Roth & Petit, 1972, Persicula, Part I (*1/1)
Megalomastomidae
serranum Alcalde y Ledon, 1945, Farcimen (Farcimen) auriculatum
CV)
Melongenidae
inspinata Richards, 1933, Melongena corona (*1/1)
johnstonet Clench & Turner, 1956, Melongena corona, Part I (4/9)

November 30, 1997 Mikkelsen & Bradford 61

sprucecreekensis Tucker, 1994, Melongena (1/10)
Mitridae
backae Cernohorsky, 1973, Scabricola (Scabricola) (2/2)
clover’ Cernohorsky, 1971, Cancilla (Ziba) (1/1)
crosniert Cernohorsky, 1970, Cancilla scrobiculata, Part I (*2/3) and
this paper (1/1)
dondani Cernohorsky, 1985, Mitra (1/1)
earle: Cernohorsky, 1977, Mitra (Nebularia) (1/1)
ekerae Cernohorsky, 1973, Scabricola (Swainsonta) ocellata (1/1)
lopest Matthews & Coelho, 1969, Mitra (Cancilla) (1/3)
vicdant Cernohorsky, 1981, Scabricola (1/1)
Muricidae
caribaea Abbott, 1958, Coralliophila (1/3)
cosmam Abbott & Finlay, 1979, Chicoreus, Part I (1/1)
dalli Emerson & D’ Attilio, 1963, Latiaxis (Babelomurex), Part I (1/2)
finlayi Clench, 1955, Murex (Murex), Part I (1/1)
guamensis Emerson & D’ Attilio, 1979, Favartia (1/1)
guestit Harasewych & Jensen, 1979, Prerynotus (Pterynotus), Part I
(*1/1)
hadari Emerson & Cernohorsky, 1973, Drupa ricinus (1/2)
haysae Clench, 1927, Thais floridana, Part I (2/4)
langi Clench & Turner, 1948, Thais (Tharsella), Part I (1/6)
lightbournt Harasewych & Jensen, 1979, Prerynotus (Pterynotus), Part I
Cyt)
macleant Shasky, 1970, Coralliophila, Part I (1/2)
mactanensis Emerson & D’ Attilio, 1979, Murextella (1/1)
macquariensis Powell, 1957, Trophon, Part I (1/3)
michael Ford, 1888, Ocinebra, Part I (1/2)
orrt Cernohorsky, 1976, Muricopsis (1/2)
philippinensis Petuch, 1979, Axymene, Part I (*1/1)
radwint Harasewych & Jensen, 1979, Prerynotus (Pterynotus), Part I
CD)
Naticidae
guesti Harasewych & Jensen, 1984, Natica (Natica), Part I (2/8 [7])
scotiana Powell, 1951, Sinuber sculpta, Part I (1/2)
Neritidae
dispar Pease, 1867, Neritina, Part I (1/6)
Neritopsidae
finlayi Hoerle, 1974, Neritopsis, Part I (1/1)
Olividae
baileyi Petuch, 1979, Oliva, Part I (*2/2)
blanest Ford, 1898, Olivella, Part I (1/2)

62 NEMOURIA No. 41

gaylordi Ford, 1895, Olivella gracilis, Part I (1/2)
marmosa Olsson & McGinty, 1958, Olivella minuta, Part I (1/1)
rejecta Burch & Burch, 1962, Oliva, Part I (1/1)
stegert Olsson, 1956, Olivella (Ohivella), Part I (1/4)
teaguei Klappenbach, 1964, Olivancillaria, Part I (1/1)
trujillo: Clench, 1938, Oliva, Part I (1/7)
Ovulidae
exquisita Petuch, 1979, Pseudocypraea, Part I (*1/1)
lahainaensis Cate, 1969, Volva (Phenacovolva), Part I (1/2)
lenoreae Cardin & Walls, 1980, Phenacovolva (Subsimnia), Part I
‘Golee
Personidae
burgesst Lewis, 1972, Distorsio, Part I (2/2)
haber Lewis, 1972, Distorsio constricta, Part I (1/1)
jenniernestae Emerson & Piech, 1992, Distorsio (Distorsio) (1/2)
Pomatiopsidae
yoshidai Bartsch, 1925, Katayama nosophora (1/6)
Potamididae
boswellae Barnard, 1963, Pirenella, Part I (*1/1)
Poteriidae
chocolatum Morrison, 1955, Incidostoma (1/5)
diminutum Morrison, 1955, Incidostoma (2/13)
jacksom Morrison, 1955, Incidostoma (2/18)
Ranellidae
boschi Abbott & Lewis, 1970, Cymatium, Part I (*1/1)
edgerlyi Richards, 1933, Gyrineum perca (*1/1)
Rissoidae
bermudensis Peile, 1926, Rissoina, Part I (1/2)
kermadecensis Oliver, 1915, Haurakia, Part I (1/3)
lyalhana Suter, 1898, Rissoia (Alvania) plicata, Part I (1/1)
mivosa Powell, 1957, Ovirissoa, Part I (1/3)
scalaroides Adams, 1850, Rissoa, Part I (1/7)
tokyoensis Pilsbry, 1904, Rissoa, Part I (1/1)
toroensis Olsson & McGinty, 1958, Rissoa (1/5)
Scissurellidae
jacksont Melvill, 1904, Scissurella (1/1)
Strombidae
nanus Bales, 1942, Strombus raninus, Part I (1/5) and this paper (1/2)
Terebridae
allyni Bratcher & Burch, 1970, Terebra, Part I (1/1)
arcas Abbott, 1954, Terebra (Strioterebrum), Part I (2/3)
betsyae Burch, 1965, Hastula (Punctoterebra), Part I (2/6)

November 30, 1997 Mikkelsen & Bradford 63

craket Burch, 1965, Duplicaria (Duplicaria), Part I (1/2)
dorothyae Bratcher & Burch, 1970, Terebra, Part I (1/1)
hancocki Bratcher & Burch, 1970, Terebra, Part I (1/1)
hoffmeyeri Abbott, 1952, Terebra (Strioterebrum), Part I (1/1)
maryleeae Burch, 1965, Hastula (Hastula), Part I (1/2)
parkinsonit Cernohorsky & Bratcher, 1976, Terebra, Part I (1/1)
steger. Abbott, 1954, Terebra (Myurella) floridana, Part I (1/1)
Tonnidae
thompsont McGinty, 1955, Eudolium, Part I (1/1)
Trochidae
acanthus Quinn, 1991, Mirachelus (*2/4)
bermudense Quinn, 1992, Calliostoma (*2/10)
bouvetia Powell, 1951, Margarella, Part I (1/2)
carinatus Quinn, 1991, Lamellitrochus (*3/54)
guesti Quinn, 1992, Calliostoma (*2/4)
huttonu Smith, 1876, Trochus (Canthiridus), Part I (1/2)
macleant Shasky & Campbell, 1964, Calliostoma, Part I (1/1)
rubra Pease, 1860, Alcyna, Part I (1/2)
zebrina Powell, 1946, Zediloma (Fractarmilla) corrosa, Part I (2/6)
Turbinellidae
aurora Bayer, 1971, Columbarium (Peristarium), Part I (1/1)
ecphoroides Harasewych, 1983, Fulgurofusus (Fulgurofusus), Part I (1/1)
electra Bayer, 1971, Columbarium (Peristarium), Part I (1/1)
gracilior Rehder, 1967, Benthovoluta, Part I (1/1)
harrisae Harasewych, 1983, Columbarium, Part I (1/1)
merope Bayer, 1971, Columbarium (Peristarium), Part I (1/1)
tumor Harasewych, 1983, Fulgurofusus (Peristarium), Part I (1/1)
wormaldi Powell, 1971, Columbarium (Coluzea), Part I (1/1)
xenismatis Harasewych, 1983, Fulgurofusus (Histricosceptrum), Part I
GD)
Turbinidae
kurzi Petuch, 1979, Guildfordia, Part I (*2/5)
Turridae
barbarae Lyons, 1972, Brachycythara, Part I (2/2)
brasilana Tippett, 1983, Borsonia, Part I (1/2)
girardi Lyons, 1972, Cerodnillia, Part I (1/1)
kingae Powell, 1964, Xenoturris, Part I (1/2)
mcgintyt Schwengel, 1943, Eubela, Part I (1/1)
petit Tippett, 1995, Fenimorea (1/1)
queenslandica Powell, 1969, Leucosyrinx, Part I (1/2)
salisburyi Cernohorsky, 1978, Mitrolumna, Part I (1/1)
steger’ McGinty, 1955, Daphnella, Part I (1/1)

64 NEMOURIA No. 41

weaver Powell, 1967, Turndrupa, Part I (1/1)

wolfe: Tippett, 1995, Drillia (1/1)
Turritellidae

ahiparana Powell, 1927, Turntella (Zeacolpus), Part I (1/6)
Vitrinellidae

elegans Olsson & McGinty, 1958, Vitrinella (Striovitrinella), Part |

(1/1)

Volutidae

barnardi Rehder, 1969, Fuszvoluta, Part I (*2/2)

boswellae Rehder, 1969, Volutocorbis, Part I (*1/1)

clarket Rehder, 1969, Fusivoluta, Part I (*3/4)

cloverrana Weaver, 1963, Lyna, Part I (1/1)

daisyae Weaver, 1967, Volutoconus hargreavesi, Part I (*3/3)

dampieria Weaver, 1960, Amoria (Amoria), Part I (1/1)

disparilis Rehder, 1969, Volutocorbis, Part I (1/3)

duponti Weaver, 1968, Festilyria, Part I (*1/1)

germonae Harasewych, 1987, Tractolira, Part I (1/2)

kellnert Iredale, 1957, Aulica, Part I (1/1)

perplicata Hedley, 1902, Voluta, Part I (1/1)

randalli Stokes, 1961, Cymbiola, Part I (1/1)

riost Clench & Turner, 1964, Adelomelon (Weaveria), Part I (1/1)

smith Bartsch, 1942, Prodallia, Part I (1/1)

vernicosum Dall, 1889, Mangilia (Calliotectum), Part I (1/1)
Volutomitridae

cuvierensis Finlay, 1930, Microvoluta, Part I (1/1)

GASTROPODA: HETEROBRANCHIA: HETEROSTROPHA:
Pyramidellidae
webert Morrison, 1965, Odostomia (1/2)

GASTROPODA: HETEROBRANCHIA: OPISTHOBRANCHIA:
Acteonidae

eloiseae Abbott, 1973, Acteon, Part I (*2/3)
Aplustridae

tenerum Powell, 1951, Parvaplustrum, Part I (1/6)
Diaphanidae

mauganuica Powell, 1952, Austrodiaphana, Part I (3/38)
Hydatinidae - see Aplustridae
Platydorididae

carolynae Mulliner & Sphon, 1974, Platydorts (1/1)

November 30, 1997 Mikkelsen & Bradford 65

BIVALVIA:
Cardiidae

thaanumt Pilsbry, 1921, Cardium (1/1)
Carditidae

micella Olsson & McGinty, 1958, Pleuromerts (1/4)
Corbiculidae

angasi Prime, 1864, Corbicula (1/1)

awajiensis Pilsbry, 1901, Corbicula (1/1)

Reiniana Clessin in Kuster, 1878, Corbicula (1/2)

sadoensis Pilsbry, 1901, Corbicula (1/1)

viola Pilsbry, 1907, Corbicula (1/1)
Corbulidae

Krebsiana Adams, 1852, Corbula (1/1)

Swiftiana Adams, 1852, Corbula (1/1)
Crassatellidae

ecuadoriana Olsson, 1961, Crassinella (1/1)
Cyrenoididae

panamensis Pilsbry & Zetek, 1931, Cvrenorda (1/1)
Donacidae

congo Pilsbry & Bequaert, 1927, Iphigenia (1/2)

nux Pilsbry & Bequaert, 1927, Egeria (1/1)
Galeommatidae

troglodytes Mikkelsen & Bieler, 1989, Divartscintilla (1/3)

yoyo Mikkelsen & Bieler, 1989, Divartscintilla (1/2)
Kelliidae

hawaiensis Pilsbry, 1921, Solecardia (1/1)
Limopsidae

diegensis Dall, 1908, Limopsis (2/7)
Mactridae

pontchartrainensts Morrison, 1965, Mulinia (1/6)

yokohamensis Pilsbry, 1895, Raeta (1/1)
Mytilidae

bryanae Pilsbry, 1921, Congeria (1/4)

nonuranus Pilsbry & Olsson, 1935, Modiolus (Modtolus) (1/1)
Nuculanidae

acapulcensis Pilsbry & Lowe, 1932, Leda (Saccella) (1/1)

exoptata Pilsbry & Lowe, 1932, Leda (Adrana) (1/1)

impar Pilsbry & Lowe, 1932, Leda (1/1)

redondoensis Burch, 1944, Nuculana (1/2)

vulgaris Brown & Pilsbry, 1913, Leda (1/232)
Nuculidae

calcicola Moore, 1977, Nucula (1/7)

66 NEM OURIA

cynthiae Moore, 1977, Condylonucula (1/2)

maya Moore, 1977, Condylonucula (1/4)
Pandoridae

rhypis Pilsbry & Lowe, 1932, Pandora (Kennerlia) (1/1)
Pectinidae

cranmerorum Waller, 1986, Somalipecten (1/1)

sablensis Clark, 1965, Aequipecten (Plagioctenium) irradians (1/1)
Semelidae

Uruguayensis Pilsbry, 1897, Semele (Abra ?) (1/2)
Sphaeriidae

lermondi Sterki, 1913, Pisidium (2/167)

mediafricana Pilsbry & Bequaert, 1927, Eupera (1/1)

patagonicum Pilsbry, 1911, Prstdium (1/5)

Sterkianum Pilsbry, 1897, Prsidium (1/2)
Tellinidae

gabbi Olsson & McGinty, 1958, Strigilla (1/1)

inflatula Dall, 1897, Macoma (1/4)

pacis Pilsbry & Lowe, 1932, Macoma (1/1)
Tindariidae

brunnea Dall, 1916, Tindaria (2/5)
Thraciidae

diegensis Dall, 1915, Thracia (1/1)

Rushi Pilsbry, 1897, Thracia (1/1)
Thyasiridae

redondoensis Burch, 1941, Aligena (1/1)
Unionidae

webbianus Wright, 1934, Unio (Elhptio) (1/1)
Veneridae

gerrardi Abbott, 1958, Transenella (1/2)

mulawana Biggs, 1969, Tivela (1/4)

omissa Pilsbry & Lowe, 1932, Macrocallista (Chionella) (1/1)

penistoni Heilprin, 1889, Cytherea (1/2)

platyptycha Pilsbry, 1901, Tapes (1/1)

sororcula Pilsbry & Lowe, 1932, Transenella (1/1)

sulfurea Pilsbry, 1904, Pitar (1/1)
Sareptidae

ensifera Dall, 1897, Yoldia (2/3)

montereyensis Dall, 1893, Yoldia (1/2)

SCAPHOPODA:
Gadilidae
teliger Finlay, 1927, Cadulus (1/1)

No. 41

November 30, 1997 Mikkelsen & Bradford 67

LITERATURE CITED

Abbott, R. T. 1958. The marine mollusks of Grand Cayman Island, British
West Indies. Monographs of the Academy of Natural Sciences of
Philadelphia, no. 11, 138 pp., 5 pls.

Abbott, R. T., M. E. Young, A. H. Clarke, W. J. Clench, R. W. Dexter, A.
LaRocque, W. E. Old, Jr., A. G. Smith, & M. C. Teskey. 1973.
American Malacologists, a national register of professional and amateur
malacologists and private shell collectors, and biographies of early
American mollusk workers born between 1618 and 1900. First edition,
1973-1974. American Malacologists, Falls Church, Virginia, iv + 494
pp.

Adams, C. B. 1852. Descriptions of new species of Corbula from Jamaica.
Contributions to Conchology, 12: 233-241.

Aguayo, C. G. & M. L. Jaume. 1957. Adiciones a la fauna malacologica
Cubana - I. Memorias de la Sociedad Cubana de Historia Natural
"Felipe Poey," 23(2): 117-142, pls. 1-6.

Alcalde y Ledon, O. 1943. Dos nuevos Chondrothyrium. Revista de la Sociedad
Malacologica "Carlos de la Torre," 1(1): 11-16, pl. 1.

Alcalde y Ledon, O. 1945. Estudio y revision de los moluscos Cubanos del
genero Farcimen, Parte I. Revista de la Sociedad Malacologica "Carlos
de la Torre," 3(1): 5-18, pl. 1.

Baker, F. 1913. The land and fresh-water mollusks of the Stanford Expedition
to Brazil. Proceedings of the Academy of Natural Sciences of
Philadelphia, December 1913: 618-672, pls. 21-27.

Baker, H. B. 1964. Type land snails in the Academy of Natural Sciences of
Philadelphia. Part III. Limnophile and thalassophile Pulmonata. Part IV.
Land and fresh-water Prosobranchia. Proceedings of the Academy of
Natural Sciences of Philadelphia, 116(4): 149-193.

Bales, B. R. 1942. A new subspecies of Strombus raninus Gmelin. The
Nautilus, 56(1): 18-19, pl. 4.

Bartsch, P. 1918. Classification of the Philippine operculate land shells of the
family Helicinidae, with a synopsis of the species and subspecies of the
genus Geophorus. Journal of the Washington Academy of Sciences, 8:
643-657.

Bartsch, P. 1925. Some new intermediate hosts of the Asiatic human blood
fluke. Journal of the Washington Academy of Sciences, 15(4): 71-73.

Bartsch, P. 1936. Molluscan intermediate hosts of the Asiatic blood fluke,
Schistosoma japonicum, and species confused with them. Smithsonian
Miscellaneous Collections, 95(5): 1-60, pls. 1-8.

Bieler, R. 1992. Gastropod phylogeny and systematics. Annual Review of

68 NEMOURIA No. 41

Ecology and Systematics, 23: 311-338.

Bieler, R. & A. Bradford. 1991. Annotated catalog of type specimens in the
malacological collection of the Delaware Museum of Natural History.
Gastropoda (Prosobranchia and Opisthobranchia). Nemouria, (36): 1-
48.

Biggs, H. E. J. 1969. Marine Mollusca of Masirah I., South Arabia. Archiv fir
Molluskenkunde, 99: 201-207.

Boss, K. J., J. Rosewater, & F. A. Ruhoff. 1968. The zoological taxa of
William Healey Dall. United States National Museum Bulletin, no. 287,
427 pp.

Bouchet, P. & A. Warén. 1986 (1985). Mollusca Gastropoda: taxonomical
notes on tropical deep water Buccinidae with descriptions of new taxa.
Memoires Museum Nationale d'Histoire Naturelle, ser. A, Zoologie,
133: 457-499, pls. 1-18.

Boyko, C. B. & W. E. Sage III. 1996. Catalog of Recent type specimens in the
Department of Invertebrates, American Museum of Natural History. II.
Mollusca Part 1 (Classes Aplacophora, Polyplacophora, Gastropoda
[Subclass Opisthobranchia], Bivalvia, and Scaphopoda). American
Museum Novitates, no. 3170, 50 pp.

Brown, A. P. & H. A. Pilsbry. 1913. Two collections of Pleistocene fossils
from the Isthmus of Panama. Proceedings of the Academy of Natural
Sciences of Philadelphia, 65: 493-500.

Burch, J. Q. 1944. Check list. West North American marine Mollusca from
San Diego, Calif. to Alaska. Minutes of the Conchological Club of
Southern California, no. 33: 7-14.

Burch, J. Q. 1945. Nuculana penderi redondoensis Burch, 1944. Minutes of the
Conchological Club of Southern California, no. 47: 32, plate figs. 32-37.

Burch, T. 1941. A survey of the west American Aligenas with a description of
a new species. The Nautilus, 55(2): 48-51, pl. 4.

Burgess, C. M. 1985. Cowries of the World. Gordon Verhoef, Seacomber
Publications, Cape Town, xvi + 289 pp.

Cate, C. N. 1975. New cypraeacean species (Mollusca: Gastropoda). The
Veliger, 17(3): 255-2611, 2 pls:

Cate, J. M. 1961. A discussion of Vexillum regina (Sowerby, 1825) and related
species, with description of a new subspecies. The Veliger, 4(2): 76-85,
pls. 18-20, table 1.

Cernohorsky, W. O. 1970a. New Mitridae and Volutomitridae. The Nautilus,
83(3): 95-104.

Cernohorsky, W. O. 1970b. Systematics of the families Mitridae &
Volutomitridae (Mollusca: Gastropoda). Bulletin of the Auckland
Institute and Museum, no. 8, iv + 190 pp., 18 pls.

Cernohorsky, W. O. 1971. New molluscan species of Strombus (Strombidae)

November 30, 1997 Mikkelsen & Bradford 69

and Cancilla (Mitridae) from New Britain and Taiwan. Records of the
Auckland Institute and Museum, 8: 131-135.

Cernohorsky, W. O. 1973. Description of new West Australian Mitridae and
Vexillidae (Mollusca: Gastropoda). Records of the Auckland Institute
and Museum, 10: 133-142.

Cernohorsky, W. O. 1976. The taxonomy of some Indo-Pacific Mollusca. Part
4. With descriptions of new taxa and remarks on Nassarius coppingert
(Smith). Records of the Auckland Institute and Museum, 13: 111-129.

Cernohorsky, W. O. 1977. The taxonomy of some Indo-Pacific Mollusca. Part
5. With descriptions of new taxa and remarks on Nassarius albus (Say).
Records of the Auckland Institute and Museum, 14: 121-132.

Cernohorsky, W. O. 1978. New species of Mitridae, Costellariidae and
Turridae from the Hawaiian Islands with notes on Mitra sphoni in the
Galapagos Islands. ‘The Nautilus, 92(2): 61-67.

Cernohorsky, W. O. 1981. The taxonomy of some Indo-Pacific Mollusca. Part
9. Records of the Auckland Institute and Museum, 18: 193-202.
Cernohorsky, W. O. 1985. The taxonomy of some Indo-Pacific Mollusca. Part
12. With remarks on two American gastropod species. Records of the

Auckland Institute and Museum, 22: 47-67.

Cernohorsky, W. O. 1986. The taxonomy of some Indo-Pacific Mollusca. Part
13. With description of a new species. Records of the Auckland Institute
and Museum, 23: 45-57.

Cernohorsky, W. O. 1991. The Mitridae of the world. Part 2. The subfamily
Mitrinae concluded and subfamilies Imbricarinae and Cylindromitrinae.
Monographs of Marine Mollusca, no. 4, 164 pp.

Chavan, A. 1969. Leptonacea, Cyamiacea. In: Moore, R. C., ed., Treatise on
Invertebrate Paleontology, Part N, Volume 2, Mollusca 6, Bivalvia.
Geological Society of America and University of Kansas, pp. 518-543.

Clarke, A. H., Jr. 1965. The scallop superspecies Aeguipecten irradians
(Lamarck). Malacologia, 2(2): 161-188.

Clench, W. J. & R. D. Turner. 1950. The western Atlantic marine mollusks
described by C. B. Adams. Occasional Papers on Mollusks, Museum of
Comparative Zoology, Harvard University, 1(15): 233-403.

Clessin, S. 1878. Die Familie der Cycladeen. In Abbildungen nach der Natur
mit Beschrebungen. In: Kuster, H. C., Systematisches Conchylien-
Cabinet von Martini und Chemnitz. Verlag von Bauer und Raspe,
Nurnberg, 9(3): 1-238.

Coan, E. V. & P. H. Scott. 1997. Checklist of the marine bivalves of the
northeastern Pacific Ocean. Santa Barbara Museum of Natural History,
Contributions in Science, no. 1, 28 pp.

Coomans, H. E., R. G. Moolenbeek & E. Wils. 1982. Alphabetical revision of
the (sub)species in recent Conidae. 5. baccatus to byssinus, including

70 NEMOURIA No. 41

Conus brettinghami nomen novum. Basteria, 46: 3-67.

Counts, C. L., HI. 1991. Corbicula (Bivalvia: Corbiculidae). Part 1. Catalog
of fossil and Recent nominal species. Part 2. Compendium of
zoogeographic records of North America and Hawaii, 1924-1984.
Tryonia, no. 21, 134 pp.

Dall, W. H. 1893. On a new species of Yoldia from California. The Nautilus,
7(3): 29-30.

Dall, W. H. 1897. Notice of some new or interesting species of shells from
British Columbia and the adjacent region. Natural History Society of
British Columbia, Bulletin, no. 2, art. 1, 18 pp., 2 pls.

Dall, W. H. 1908. Reports on the dredging operations off the west coast of
Central America to the Galapagos, to the west coast of Mexico, and in
the Gulf of California, in charge of Alexander Agassiz, carried on by the
U. S. Fish Commission Steamer "Albatross," during 1891, Lieut.
Commander E. L. Tanner, U. S. N., commanding. XX XVII. Reports on
the scientific results of the expedition to the eastern tropical Pacific, in .
charge of Alexander Agassiz, by the U. S. Fish Commission Steamer _
"Albatross," from October, 1904, to March, 1905, Lieut. Commander
L. M. Garrett, U. S. N., commanding. XIV. The Mollusca and the
Brachiopoda. Bulletin of the Museum of Comparative Zoology at
Harvard College, 43(6): 205-487, pls. 1-22.

Dall, W. H. 1915. A review of some bivalve shells of the group Actinacea from
the west coast of America. Proceedings of the United States National
Museum, 49(2116): 441-456.

Dall, W. H. 1916. Diagnoses of new species of marine bivalve mollusks from
the northwest coast of America in the collection of the United States
National Museum. Proceedings of the United States National Museum,
52(2183): 393-417.

Emerson, W. K. & W. O. Cernohorsky. 1973. The genus Drupa in the Indo-
Pacific. Indo-Pacific Mollusca, 3(13): 1-40.

Emerson, W. K. & A. D'Attilio. 1979. Six new living species of muricacean
gastropods. The Nautilus, 93(1): 1-10.

Emerson, W. K. & B. J. Piech. 1992. Remarks on Distorsio constricta (Broderip,
1833) and related species in the eastern Pacific Ocean, with the
description of a new species (Gastropoda: Personidae). The Veliger,
35(2): 105-116.

Emerson, W. K. & W. E. Sage III. 1986. On the rediscovery of Teramachia
mirabilis (Clench and Aguayo, 1941), and its relationship to other
calliotectine volutes. The Nautilus, 100(4): 147-151.

Ferreira, A. J. 1976. A new species of Callistochiton in the Caribbean. The
Nautilus, 90(1): 46-49.

Finlay, C. J. 1927 (1926). A further commentary on New Zealand molluscan

November 30, 1997 Mikkelsen & Bradford 71

systematics. Transactions of the New Zealand Institute, 57: 320-485,
pls. 18-23.

Heilprin, A. 1889. On some new species of Mollusca from the Bermuda
Islands. Proceedings of the Academy of Natural Sciences of Philadelphia,
1889: 141-142, pl. 8.

Henderson, J. B., Jr. & C. T. Simpson. 1902. A new Haitien Chondropoma.
The Nautilus, 16(8): 88-89.

Henning, T. & J. Hemmen. 1993. Ranellidae and Personidae of the World.
Verlag Christa Hemmen, Wiesbaden, Germany, 263 pp.

Herrington, H. B. 1962. A revision of the Sphaeriidae of North America
(Mollusca: Pelecypoda). Miscellaneous Publications, Museum of
Zoology, University of Michigan, no. 118, 75 pp., 8 pls.

Houart, R. 1994. Illustrated Catalogue of Recent Species of Muricidae named
since 1971. Verlag Christa Hemmen, Wiesbaden, Germany, 179 pp.

Howard, F. B. 1963. New name for a Pyrene from Mexico (Mollusca -
Gastropoda). Santa Barbara Museum of Natural History, Occasional
Papers, no. 7 (suppl.): (1 p.).

International Commission on Zoological Nomenclature [ICZN]. 1985.
International Code of Zoological Nomenclature, 3rd ed. International
Trust for Zoological Nomenclature, London, xx + 338 pp.

Jacobson, M. K. & W. K. Emerson. 1971. Shells from Cape Cod to Cape
May, with Special Reference to the New York City Area. Dover
Publications, New York, 152 pp.

Jaume, M. L. & O. Alcalde y Ledon. 1944. Nuevo Chondropoma de
Cienfuegos, Cuba Central. Revista de la Sociedad Malacologia "Carlos
de la Torre," 2(2): 39-40, pl. 6.

Jaume, M. L. & L. Sanchez de Fuentes. 1943a. Nuevo molusco del genero
Chondropoma. Revista de la Sociedad Malacologia "Carlos de la Torre,"
LOe 17-18, pli 2.

Jaume, M. L. & L. Sanchez de Fuentes. 1943b. Dos nuevas subespecies de
Eutudora limbifera. Revista de la Sociedad Malacologia "Carlos de la
Torre,” 1(1)2 19-21, pl. 3.

Jaume, M. L. & L. Sanchez de Fuentes. 1943c. Dos subgeneros y tres nuevas
especies de Chondrothyrium. Revista de la Sociedad Malacologia "Carlos
de la Torre," 1(1): 22-29, pl. 4.

Jaume, M. L. & L. Sanchez de Fuentes. 1943d. Nueva subespecie de
Chondrothynum violaceum. Revista de la Sociedad Malacologia "Carlos
de la Torre," 1(2): 50-51, pl. 8.

Johnson, R. I. 1959. The types of Corbiculidae and Sphaeriidae (Mollusca:
Pelecypoda) in the Museum of Comparative Zoology, and a bio-
bibliographic sketch of Temple Prime, an early specialist of the group.
Bulletin of the Museum of Comparative Zoology at Harvard College,

te NEMOURIA No. 41

120(4): 429-479.

Kabat, A. R. 1996. Molluscan types of the Albatross expeditions to the eastern
Pacific described by W. H. Dall (1908). Bulletin of the Museum of
Comparative Zoology, 155(1): 1-31.

Kay, E. A. 1979. Hawaiian Marine Shells. Bishop Museum Press, Honolulu,
652/pp:

Keen, A. M. 1971. Sea Shells of Tropical West America; marine mollusks
from Baja California to Peru, 2nd ed. Stanford University Press,
Stanford, California, 1064 pp., 22 pls.

Krieger, K. A. 1972. Somatogyrus alcoviensis, new gastropod species from
Georgia (Hydrobiidae). The Nautilus, 85(4): 120-125, figs. 1-4.
Lindberg, D. R. 1986. Name changes in the “Acmaeidae.” The Veliger, 29:

142-148.

Lyons, W. G. 1988. A review of Caribbean Acanthochitonidae (Mollusca:
Polyplacophora) with descriptions of six new species of Acanthochitona
Gray, 1821. American Malacological Bulletin, 6(1): 79-114.

Malek, E. A. & M. D. Little. 1971. Aroapyrgus colombiensis n. sp. (Gastropoda:
Hydrobiidae), snail intermediate host of Paragonimus caliensis in
Colombia. The Nautilus, 85(1): 20-26.

Matthews, H. R. & A. C. dos Santos Coelho. 1969. Uma nova espécie da
familia Mitridae do Brasil (Mollusca: Gastropoda). Boletim do Museu
Nacional, nova série, Zoologia, no. 272: 1-8. [The first author's last
name is incorrectly spelled on the title page as Mattews. |

Melvill, J. C. 1904. Description of twenty-eight species of Gastropoda from
the Persian Gulf, Gulf of Oman, and Arabian Sea, dredged by Mr. F. W.
Townsend, of the Indo-European Telegraph Service, 1900-1904.
Proceedings of the Malacological Society of London, 6: 158-169, pl. 10.

Mikkelsen, P.M. & R. Bieler. 1989. Biology and comparative anatomy of
Divariscintilla yoyo and D. troglodytes, two new species of Galeommatidae
(Bivalvia) from stomatopod burrows in eastern Florida. Malacologia,
31(1)2 175-195;

Moore, D. R. 1977. Small species of Nuculidae (Bivalvia) from the tropical
western Atlantic. The Nautilus, 91(4): 119-128.

Moretzsohn, F. & E. A. Kay. 1995. Hawaiian marine molluscs - an update to
Kay, 1979. University of Hawai at Manoa, Honolulu, 23 pp.

Morrison, J. P. E. 1955. Notes on American cyclophoroid land snails, with
two new names, eight new species, three new genera, and the family
Amphicyclotidae, separated on animal characters. Journal of the
Washington Academy of Sciences, 45(5): 149-162.

Morrison, J. P. E. 1965. New brackish water mollusks from Louisiana.
Proceedings of the Biological Society of Washington, 78:217-224.

Morrison, J. P. E. 1973. New name for a Texas Hydrobia. The Nautilus,

November 30, 1997 Mikkelsen & Bradford 73

87(1): 28.

Mulliner, D. K. & G. G. Sphon. 1974. A new Platydoris (Gastropoda;
Nudibranchia) from the Galapagos Islands. Transactions of the San
Diego Society of Natural History, 17(15): 209-216.

Olsson, A. A. 1961. Mollusks of the tropical eastern Pacific particularly from
the southern half of the Panamic-Pacific faunal province (Panama to
Peru). Panamic-Pacific Pelecypoda. Paleontological Research Institution,
Ithaca, New York, 574 pp., 86 pls.

Olsson, A. A. & T. L. McGinty. 1958. Recent marine mollusks from the
Caribbean coast of Panama with the description of some new genera and
species. Bulletins of American Paleontology, 39(177): 1-59.

Parodiz, J. J. & J. J. Tripp. 1988. Types of Mollusca in the collection of the
Carnegie Museum of Natural History. Part I. Bivalvia and Gastropoda
(Prosobranchia and Opisthobranchia). Annals of Carnegie Museum,
57(5): 111-154.

Pilsbry, H. A. 1895. Catalogue of the marine mollusks of Japan with
descriptions of new species and notes on others collected by Frederick
Stearns. F. Stearns, Detroit, viii + 196 pp., 11 pls.

Pilsbry, H. A. 1897. New species of mollusks from Uruguay. Proceedings of
the Academy of Natural Sciences of Philadelphia, 49: 290-298, pls. 6-7.

Pilsbry, H. A. 1901a [March]. New Mollusca from Japan, the Loo Choo
Islands, Formosa and the Philippines. Proceedings of the Academy of
Natural Sciences of Philadelphia, 53: 193-210.

Pilsbry, H. A. 1901b [July]. New Japanese marine, land and fresh-water
Mollusca. Proceedings of the Academy of Natural Sciences of
Philadelphia, 53: 385-408, pls. 19-21.

Pilsbry, H. A. 1904. New Japanese marine Mollusca: Pelecypoda. Proceedings
of the Academy of Natural Sciences of Philadelphia, 56: 550-561, pls.
39-41.

Pilsbry, H. A. 1907. On Japanese species of Corbicula. Annotationes
Zoologicae Japonenses, 6(3): 153-160, pl. 7.

Pilsbry, H. A. 1911. Non-marine Mollusca of Patagonia. Reports of the
Princeton University Expeditions to Patagonia, 1896-1899. Vol. 3 -
Zoology. Part 5: 513-633, pls. 38-47.

Pilsbry, H. A. 1921 (1920). Marine mollusks of Hawaii, VIII-XIII.
Proceedings of the Academy of Natural Sciences of Philadelphia, 72:
296-328, pl. 12.

Pilsbry, H. A. & J. Bequaert. 1927. The aquatic mollusks of the Belgian
Congo. With a geographical and ecological account of Congo
malacology. Bulletin of the American Museum of Natural History, 53:
69-602, pls. 10-77.

Pilsbry, H. A. & H. N. Lowe. 1932. West Mexican and Central American

74 NEMOURIA No. 41

mollusks collected by H. N. Lowe, 1929-31. Proceedings of the
Academy of Natural Sciences of Philadelphia, 84: 33-144, pls. 1-17.

Pilsbry, H. A. & A. A. Olsson. 1935. New mollusks from the Panamic
Province. The Nautilus, 49(1): 16-19, pl. 1.

Pilsbry, H. A. & J. Zetek. 1931. A Panamic Cyrenoida. The Nautilus, 45(2):
69, pl. 3.

Ponder, W. F. & A. Warén. 1988. Appendix: Classification of the
Caenogastropoda and Heterostropha - a list of the family-group names
and higher taxa. Malacological Review, Suppl. 4 (Prosobranch
Phylogeny): 288-326.

Poppe, G. T. & Y. Goto. 1992. Volutes. Mostra Mondiale Malacologia,
Cupra Marittima, Italy, 348 pp.

Powell, A. W. B. 1958. Mollusca from the Victoria-Ross Quadrants of
Antarctica. Reports of the B. A. N. Z. Antarctic Research Expedition,
1929-1931, Ser. B, 6(9): 165-215, pls. 1-3.

Prime, T. 1864. Description d'une nouvelle espéce de Corbicula - C. angast.
Journal de Conchyliologie, 12: 151-152, pl. 7, fig. 6.

Quinn, J. F., Jr. 199la. Lamellitrochus, a new genus of Solariellinae
(Gastropoda: Trochidae), with descriptions of six new species from the
western Atlantic Ocean. The Nautilus, 105(3): 81-91.

Quinn, J. F., Jr. 1991b. New species of Gaza, Mirachelus, Calliotropis, and
Echinogurges (Gastropoda: Trochidae) from the northwestern Atlantic
Ocean. The Nautilus, 105(4): 166-172.

Quinn, J. F., Jr. 1992. New species of Calhostoma Swainson, 1840
(Gastropoda: Trochidae), and notes on some poorly known species from
the western Atlantic Ocean. The Nautilus, 106(3): 77-114.

Richards, H. G. 1933. New varieties of Melongena and Gyrineum. The
Nautilus, 47(2): 57, pl. 6.

Robertson, R., [C.] L. Richardson, G. M. Davis & A. E. Bogan. 1981.
Catalog of the types of neontological Mollusca of the Academy of
Natural Sciences of Philadelphia. Part 2. Gastropoda,
Archaeogastropoda: Pleurotomariacea, Fissurellacea, Patellacea.
Tryonia, no. 5, 48 pp.

Robertson, R., C. L. Richardson, G. M. Davis & A. E. Bogan. 1986. Catalog
of the types of Recent Mollusca of the Academy of Natural Sciences of
Philadelphia. Part 4. Gastropoda, Archaeogastropoda: Trochacea
(concluded), Neritacea (sensu lato). Tryonia, no. 14, 1 + 153 pp.

Ruhoff, F. A. 1973. Bibliography and zoological taxa of Paul Bartsch.
Smithsonian Contributions to Zoology, no. 143, 166 pp.

Schilder, F. A. & R. Summers. 1963. One more new species of Notocypraea.
The Cowry, 1(5): 65-66.

Shasky, D. R. & G. B. Campbell. 1964. New and otherwise interesting species

November 30, 1997 Mikkelsen & Bradford Us

of mollusks from Guaymas, Sonora, Mexico. The Veliger, 7(2): 114-
120, pls. 21-22.

Spamer, E. E. & A. E. Bogan. 1992. Catalog of the types of neontological
Mollusca of the Academy of Natural Sciences of Philadelphia. Part 7:
Cephalopoda and Scaphopoda. Tryonia, no. 24, vi + 23 pp.

Sterki, V. 1912. Sphaeriidae, old and new. The Nautilus, 26(1): 6-9.

Sterki, V. 1913. Sphaeriidae, old and new, IV. The Nautilus, 26(12): 136-
140.

Thompson, F. G. 1978. A new genus of operculate land snails from
Hispaniola with comments on the status of family Annulariidae. The
Nautilus, 92(1): 41-54.

Tippett, D. L. 1995. Taxonomic notes on the western Atlantic Turridae
(Gastropoda: Conoidea). The Nautilus, 109(4): 127-138.

Trew, A. 1987. James Cosmo Melvill’s New Molluscan Names. National
Museum of Wales, Cardiff, 84 pp.

Tucker, J. K. 1994. The crown conch (Melongena: Melongenidae) in Florida
and Alabama with the description of Melongena sprucecreekensis, n. sp.
Bulletin of the Florida Museum of Natural History, 36(7): 181-203.

Turgeon, D. D., A. E. Bogan, E. V. Coan, W. K. Emerson, W. G. Lyons, P.
M. Mikkelsen, J. F. Quinn, Jr., C. F. E. Roper, G. Rosenberg, B. Roth,
M. J. Sweeney, A. H. Scheltema, F. Thompson, M. Vecchione & J. D.
Williams. In press. Common and Scientific Names of Aquatic
Invertebrates from the United States and Canada: Mollusks, 2nd ed.
American Fisheries Society, Bethesda, Maryland.

Vaught, K. C. 1989. A Classification of the Living Mollusca. American
Malacologists, Melbourne, Florida, x11 + 195 pp.

Waller, T. R. 1986. A new genus and species of scallop (Bivalvia: Pectinidae)
from off Somalia, and the definition of a new tribe Decatopectinini. The
Nautilus, 100(2): 39-46.

Watters, G. T. 1981. Two new species of Acanthochitona from the New World
(Polyplacophora: Cryptoplacidae). The Nautilus, 95(4): 171-177.

Weaver, C. S. & J. E. duPont. 1970. Living Volutes, a monograph of the
Recent Volutidae of the world. Delaware Museum of Natural History,
Greenville, 375 pp.

Wells, F. E. 1977. Type specimens in the Department of Molluscs, Western
Australian Museum. Records of the Western Australian Museum, 6(1):
33-61.

Wheat, S. C. 1913. Long Island fauna and flora - II. A Long Island Acmaea,
and a new variety of Urosalpinx cinerea. Science Bulletin, The Museum
of The Brooklyn Institute of Arts and Sciences, 2(2): 17-20, pl. 1.

Wnight, B. H. 1934. New Florida pearly mussels. The Nautilus, 47(3): 94-95,
pi 10:

76 NEMOURIA No. 41

ACKNOWLEDGMENTS

We thank DMNH curatorial staff and volunteers, especially Dr. Timothy
A. Pearce, Dr. Albert F. Chadwick, Mr. Russell H. Jensen, Mr. Geoffrey
Halfpenny, and Ms. Elizabeth Shea, for technical assistance and for providing
background information on collection acquisitions. Former DMNH Curator,
Dr. Rudiger Bieler (FMNH), provided valuable advice on the collection and
on interpretation of presumed type specimens. Dr. Gary Rosenberg and Mr.
Ned Gilmore assisted greatly with this project in a number of ways at ANSP.
Assistance with literature and type material of various taxa was provided by A.
Bogan (North Carolina State Museum of Natural Science, Raleigh), C. B.
Boyko (AMNH), D. Dan (Ft. Myers, Florida), R. Germon (USNM), A.
Kabat (USNM), R. E. Petit (North Myrtle Beach, South Carolina), B. J.
Piech (DMNH), J. Slapcinsky (FMNH), and D. Tippett (USNM). Foreign
language translations were provided by V. Ovtsharenko (AMNH). C. B.
Boyko (AMNH), M. G. Harasewych, and one anonymous reviewer improved
the manuscript through their comments.

age
Fhe. =
; -
: r 5 7
~
: -
a :
5 ‘ ee
+
oS
' 4 a
a ppees _
‘
7 6
* oO":
- 1
a - =
:
» a -
; a) ae -
ar a Si = aan
we ’ 7 --
- oes
_ ¢
ca
1 1,
ra . 1
" 1 ; 7
y 7 7
= =i 7
A
fl ae ; . . a
ae a - a
c a ‘ ~
ars - ae
‘
a oe
1 -
‘ i an
: =
- -
‘ ‘ oe
[& <i { 7
1 7 I =
att } -
i b>
i s: -
, = =
q
,

*

¥ - -
oe
+; ’
ay
: i eh)
‘ . - 3
ae
- —
2 se ee
eS ) ee
i oe -

r .- '
; i 7 bo _ : a ~ 1
eas i =
. a ¢ 2
A, 2 ra i
2 - .
— -
ee a
“ =
a 7 :
<
ea -_ ee ‘
/ a o -
1
: 7 7 ae , i -
asf
i a = 7 7 = eee

; :
a -
= ou _
=
ue
oe =
: -
cay
oe
— ~
=
i
-
=
i
= ”
{
.

=

Founding editor Managing editor
JOHN E. DU PONT TIMOTHY A. PEARCE

Consulting Editors

WILLIAM K. EMERSON, American Museum of Natural History, New York,
NY.

M. G. HARASEWYCH, National Museum of Natural History, Smithsonian
Institution, Washington, DC.

KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh, PA.

RICHARD W. THORINGTON, JR., National Museum of Natutal History,
Smithsonian Institution, Washington, DC.

Published by the

DELAWARE MUSEUM OF NATURAL HISTORY
P. O. Box 3937, Wilmington,
Delaware 19807, U.S.A.

(NOV 1 6 20 )

LIBRARIES
NEMOURIA

Occasional Papers of the Delaware Museum of Natural History

NUMBER 42 OCTOBER 25, 2000

THREE NEW CHITONS OF THE GENUS LEPIDOZONA
PILSBRY, 1892 (POLYPLACOPHORA:
ISCHNOCHITONIDAE) FROM THE ALEUTIAN ISLANDS

Roger N. Clark”

ABSTRACT. The genus Lepidozona Pilsbry, 1892 in Aleutian waters is discussed. In
addition to the three species already recorded from the area, L. trifida (Carpenter,
1864), L. ima Sirenko, 1975, and L. allyni (Ferreira, 1977), three new species, L.
baxteri, L. beringiana and L. attuensis are described, and the range of one additional
species, L. abyssicola (Smith @ Cowan, 1966) is extended west into the region. The
new species are compared to known members of the genus from the Aleutians, as
well as to L. lindbergi (Yakovleva, 1952) from the Russian, Kurile Islands.

Key words: Chiton, Ischnochitonidae, Lepidozona, Aleutian Islands.

INTRODUCTION

The genus Lepidozona Pilsbry, 1892 has a worldwide distribution, and
contains more than 50 known species. Kaas and Van Belle (1987) reviewed the
genus globally, and divided its members into two subgenera based on the number
of slits in the intermediate valves. The members of the subgenus Lepidozona (sensu
stricto) have a single slit per side. Members of the subgenus Tripoplax Berry, 1919
have two or more slits per side. In their review, Kaas and Van Belle recorded five
species from Alaska, including two, L. (T.) allyni and L. (T.) ima from the
Aleutians.

The distributions of Alaskan taxa, particularly those of the Aleutian Islands,
have remained poorly defined. Berry (1917) first recorded three species of

*Field Associate in Malacology, Natural History Museum of Los Serial Publication
Angeles County, 900 Exposition Boulevard, Los Angeles, ISSN 0085-3887
California 90007, U.S.A., email: insignis@cdsnet.net

Mailing Address: 1839 Arthur Street, Klamath Falls, Oregon 97603-4617

2 NEMOURIA No. 42

Lepidozona (as a subgenus of Ischnochiton Gray, 1847) in Alaskan waters, but placed
two other species in the genera Ischnochiton Gray, 1847 and _ Ischnoradsia
Shuttleworth, 1853 (again as subgenera of Ischnochiton). Smith and Cowan (1966)
described Ischnochiton abyssicola from the Gulf of Alaska and remarked on its
affinities with Ischnochiton trifidus, suggesting that both species should perhaps be
placed in the subgenus Tripoplax. Ferreira (1977) described Ischnochiton allyni from
Amchitka Id. in the Aleutians and suggested that his new species might be a
member of Lepidozona, or perhaps Gurjanovillia Yakovleva, 1952 (now recognized
as a synonym of Tripoplax). Baxter (1987) recorded only L. allyni (as Gurjanovillia)
from the Aleutians, but used Lepidozona, Tripoplax, Ischnochiton, and Stenosemus
Middendorff, 1847 for other taxa presently assigned to this genus. Sirenko (1975)
described L. ima from the vicinity of the Commander Is., west of the Aleutians.
Clark (1991) reported L. scabricostata (Carpenter, 1864) from the Gulf of Alaska,
and L. ima and L. trifida from the Aleutians.

Presently seven species of Lepidozona are known from the Aleutian Islands, all
of them members of Tripoplax: L. (Tripoplax) abyssicola (Smith & Cowan, 1966), L.
(T.) allyni (Ferreira, 1977), L. (T.) ima Sirenko, 1975, L. (T.) trifida (Carpenter,
1864), L. (T.) beringiana new species, L. (T.) baxteri new species, and L. (T.) attuensis
new species.

MATERIALS AND METHODS

Chitons were collected intertidally by hand and to a depth of 18 m by
SCUBA. Deep-water specimens (85-350 m) were collected during National
Marine Fisheries Service triennial Aleutian Islands Trawl Surveys (1994 & 1997)
carried out by the chartered fishing Vessels F/V Vesteraalen and F/V Dominator.
Collections were made by otter trawl and by small dredges consisting of a heavy
mesh bag (open at the tail end), with a small (12.5 mm) mesh liner, tied at the tail
end and held open by a heavy chain at the mouth hung from the foot rope of the
otter trawl, just behind the 35 cm “roller gear” (rock rollers).

National Marine Fisheries Service data are recorded in the material sections
by a series of numbers: vessel number, cruise number, and haul number. For
example 23-971-199 would be 23, F/V Dominator, Cruise 971 (1997-1) (some
vessels do more than one cruise per year), Haul (or trawl) number 199. Vessel
number 23 is the F/V Dominator, vessel 94 is the F/V Vesteraalen, and vessel 57
is the F/V Morning Star. Additional survey data are available upon request to Dr.
Gary Stauffer, division director, Resource Assessment and Conservation
Engineering Division, National Marine Fisheries Service, Alaska Fisheries Science
Center, 7600 Sand Point Way NE, Bin C-15700, Bldg. 4, Seattle, Washington
98115, U.S.A.

October 25, 2000 Clark 3

The specimens were relaxed in cool seawater, tied flat and preserved in 75%
ethanol. The valves of disarticulated specimens were cleaned in household bleach,
rinsed in distilled water and dried. Radulae and girdle fragments were mounted
on scanning electron microscope (SEM) stubs, sputter coated with gold-palladium
for 2 minutes and examined at 10 ky, with a Hitachi S-2100 scanning electron
microscope at the Department of Biology at Southern Oregon University.

Abbreviations used in the text are: CAS, California Academy of Sciences;
DMNH, Delaware Museum of Natural History; LACM, Los Angeles County
Museum of Natural History; NMFS, National Marine Fisheries Service; NMFS-
AB, Auke Bay Laboratory; OS&S, Of Sea and Shore Museum; RMNH, National
Museum of Natural History, Leiden; RNC, the author’s collection; SBMNH,
Santa Barbara Museum of Natural History; SOU, Southern Oregon University;
UAF, University of Alaska, Fairbanks; USNM, United States National Museum
(Smithsonian); and ZISP, Zoological Institute, Saint Petersburg.

SYSTEMATICS

Class: POLYPLACOPHORA Gray, 1821
Family: ISCHNOCHITONIDAE Dall, 1889
Genus: Lepidozona Pilsbry, 1892

Chitons of small to medium size (10-60 mm), terminal valves and lateral areas
of intermediate valves with radial ribs or rows of pustules; central areas of valves
with longitudinal ribs, often crossed by similar but smaller riblets forming a
lattice; interior of valves (articulamentum) with sutural plate, separated (often
obscurely) from the sutural laminae by small notches or slits; insertion plate slit
formula is many/one or more/many; dorsal surface of girdle with convex, usually
ribbed or striated scales, often with a nipple-like process at the apices.

Aleutian Islands species: Lepidozona (Tripoplax) trifida, L. (T.) abyssicola, L. (T.)
ima, L. (T.) allyni, L. (T.) beringiana n. sp., L. (T.) baxteri n. sp., L. (T.) attuensis n. sp.

Lepidozona (Tripoplax) beringiana new species
(Figures 1-7)
Lepidozona (Tripoplax) ima Sirenko; Clark, 1991: 93 (in part) non L. ima Sirenko,
1975.

Diagnosis: Chitons of medium size (to 4.5 cm), oval to elongate-oval in outline,
Tegmentum brick red, often with paler terminal and lateral areas, and darker
specks. Girdle with small (275 tm x 225 ym) scales, slightly bent and striated
along the upper half, topped with a long mammillation about 35 pm in length.

4 NEMOURIA No. 42

Figures 1-7. Lepidozona (Tripoplax) beringiana new species. 1. Whole animal,
paratype, RNC 2993; 30.0 mm x 14.5 mm. 2-4. Paratype, RNC 3010. 2. Head
valve, width 10.2 mm. 3. Intermediate valve five, width 12.0 mm. 4. Tail valve,

width 9.0 mm. 5-7. Holotype, LACM 2798. 5,6. Dorsal girdle scales. 7. Radula.

October 25, 2000 Clark 5

Description: Body (Fig. 1) oval in outline, valves microgranular, carinated,
unbeaked, side slopes slightly convex; tegmentum brick red often with darker
specks, terminal and lateral areas often paler; girdle light brown to reddish.

Head valve (Fig. 2) semicircular and bearing about 40 low, faint, rounded
radial ribs capped by a row of very minute (0.1 mm in diameter), low, rounded
granules (often obsolete), especially on older portion of shell; posterior margin
widely V-shaped; insertion plate with 15-17 slits.

Intermediate valves (Fig. 3) oblong in outline, central areas with very fine
pitting; lateral areas well-defined, slightly raised, cut by two or three shallow sulci
into three or four low, rounded (often bifurcated) ribs frequently bearing a few
very minute granules like those on the head valve; insertion plates with 2-4 slits.

Tail valve (Fig. 4) semicircular, slightly elongated posteriorly; mucro central,
post-mucronal slope straight to slightly concave; insertion plate with 15 slits.

Interior of valves (articulamentum) white; sutural laminae short, slightly
rounded, connected across the jugal sinus by a narrow, concave jugal plate
notched at the sides; insertion teeth short and sharp, those on the head valve
smooth, those on the tail valve slightly rugose.

Girdle about 1/4 as wide as fifth valve at intermediate valves, much narrower
at ends; dorsal side covered with overlapping scales (Figs. 5 & 6) slightly longer
than wide, up to about 275 pm x 225 um, with about 18 striations along the
upper half and capped by a long nipple about 35 um in length; ventral surface
covered by radiating rows of elongated, rectangular, transparent scales of about
100 x 20 pm; margin of girdle with long, slender spicules up to 500 pm long.

Radula (fig. 7, holotype) 8 mm in length and bearing about 30 mature rows
of teeth; rachidian tooth spatulate, distally dilated, distal edge about 90 um wide;
blade of major lateral teeth relatively large and heavy, bicuspid, the inner denticle
much longer than the outer one.

Gills holobranchial, adanal, 30 per side.

Type locality: South of Semisopochnoi Island, Rat Islands, Aleutian Islands,
Alaska (51°53.34'N, 179°45.58'E), 121 m.

Material examined: Type material: Holotype, LACM 2798 (leg. Rae Baxter, 7
September 1986); twenty paratypes: 1, LACM 2799 (disarticulated valves only),
from type locality; 1, LACM 2800, W of Amatignak Island, Andreanof Islands,
Aleutians (51°14.46'N, 179°11.84'W), 241 m (leg. Rae Baxter, 17 August 1986);
1, ZIAS 2149, W of Kiska Island, Rat Islands, Aleutians (52°04.29'N,
177°15.33'E), 91 m (leg. RNC, 31 July 1997) (NMFS 23-971-199); 1 USNM
880313, (also from W of Kiska); 1, CAS 120751, S of Tanaga Island, Andreanof
Islands, Aleutians (51°44.50'N, 178°07.68'W), 95 m (leg. RNC, 15 July 1997)
(NMFS 23-971-137); 1, DMNH 210608, Petrel Bank (near type locality)
(52°19.18'N, 179°50.08'E), 116 m (leg. RNC, 21 July 1997) (NMFS 23-971-162);

6 NEMOURIA No. 42

1, SBMNH 345415, S of Tanaga (as above) (NMFS 23-971-137); 1, RNC 1216,
Petrel Bank, N of Semisopochnoi Island, Rat Islands, Aleutian Islands, Alaska
(52°10.60'N, 179°43.77'E), 92 m (leg. RNC, 10 July 1994) (NMFS 94-941-151); 4,
RNC 2992, S of Tanaga (as above) (NMFS 23-971-137); 2, RNC 3010, Petrel
Bank (near type locality) (52°19.18'N, 179°50.08'E), 116 m (leg. RNC, 21 July
1997) (NMFS 23-971-162); 3, RNC 3016, W of Kiska Island, Rat Islands,
Aleutians (52°04.29'N, 177°15.33'E), 91 m (leg. RNC, 31 July 1997) (NMFS 23-
971-199).

Additional material: 2, RNC 2993, S of Tanaga Island, Andreanof Islands,
Aleutians (51°46.69'N, 178°10.59'W), 89 m (leg. RNC, 17 July 1997) (NMFS 23-
971-141); 1, RNC 2994, SSW of Tanaga Island, Andreanof Islands, Aleutians
(51°43.17'N, 178°17.23'W), 215 m (leg. RNC, 17 July 1997) (NMFS 23-97 1-142),
1, RNC 3005, SE of Kiska Island, Rat Islands, Aleutians (51°53.86'N,
178°20.82'E) 143 m (leg. RNC, 28 July 1997) (NMFS 23-971-185); 1, RNC 3006,
E of Kiska Island, Rat Islands, Aleutians (52°03.44'N, 178° 18.62'E), 352 m (leg.
RNC, 29 July 1997) (NMFS 23-971-187); 1, RNC 3021, W of Buldir Island,
Aleutians (52°19.77'N, 175°48.56'E), 226 m (leg. RNC, 9 August 1997) (NMFS
23-971-241).

Distribution: Endemic to the central Aleutian Islands, Alaska [178°W (near
Tanaga Island) to 175°E (W of Buldir Island)] (Fig. 30).

Habitat: Lives at 85-352 m on cobbles, bivalve shells (Chlamys sp.) and bryozoans.

Remarks: This species was originally thought to be Lepidozona (Tripoplax) ima
(Clark, 1991: 93). Close examination revealed it instead to be distinct, and
morphologically more similar to L. (T.) allyni (Figs. 25 & 26). These two species
are characterized by the very long mammillation on the girdle scales. L. (T.)
beringiana is also similar to L. (T.) lindbergi (Yakovleva, 1952) of the Kurile Is. (Fig.
27), but may be distinguished by the fine pitting of the central areas and the
differently proportioned girdle scales.

Lepidozona (Tripoplax) baxteri new species

(Figs. 8-14)

Diagnosis: Chitons of medium size (to 3.5 cm), oval to elongate-oval in outline,
valves whitish or tan with speckles or concentric rings of reddish-brown or (rarely)
pink. Girdle with small (325 pm x 250 pm) scales bearing 22-25 striations, with
slight, if any mammillation.

Description: Body (Fig. 8) (holotype) oval in outline; valves microgranular,
carinated, unbeaked (juveniles may be slightly beaked), side slopes convex;
tegmentum whitish to tan or light brown, with dark brown, reddish-brown or

October 25, 2000 Clark a

oy

Figures 8-14. Lepidozona (Tripoplax) baxteri new species. 8. Whole animal,
holotype, LACM 2796; 27.5 mm x 16.0 mm. 9-11. Paratype, LACM 2797. 9.
Head valve, width 9.0 mm. 10. Intermediate valve five, width 10.5 mm. 11. Tail
valve, width 7.6 mm. 12-14. Paratype, RNC1230. 12, 13. Dorsal girdle scales. 14.
Radula, RNC 1230 (25 mm specimen).

8 NEMOURIA No. 42

(rarely) pink-lavender speckles and/or concentric rings; girdle unicolored whitish
to light brown, or with alternating bars of reddish-brown and tan.

Head valve (Fig. 9) semicircular, bearing 21-27 low, faint ribs capped by an
often incomplete or obsolete row of minute (about 80-100 pm in diameter),
widely-spaced granules, three to eight in a series; interstices are granular, posterior
margin widely V-shaped; insertion plate with 15 slits.

Intermediate valves (Fig. 10) oblong in outline, central areas with fine pitting,
lateral areas with three or four radiating, sub-triangular ribs capped by an often
incomplete or obsolete row of minute granules of the same nature and number as
those on the head valve; insertion plates with 2 slits.

Tail valve (Fig. 11) semicircular; mucro post-central, post-mucronal area
flattened and bearing about 18 faint or obsolete radiating ribs of the same nature
as those on the head and lateral areas; insertion plate with 15 slits (teeth prone to
splitting).

Interior of valves (articulamentum) white, translucent; sutural laminae short,
thin, sharp, connected across the jugal sinus by a narrow, concave jugal plate,
slightly notched at the sides in juveniles, but not in adults; insertion teeth short,
solid, smooth; those of head valve blunt, rugose on the anterior side, those of the
tail valve still rather blunt, but somewhat sharper.

Girdle nearly one half as wide as fifth valve at sides, much narrower at ends;
dorsal side covered with overlapping, bent, near-pentagonal scales, wider than
long (Figs. 12-13), (imperceptibly mammillated) and bearing 22-25 striations,
reaching about 325 pm x 250 pm; ventral surface covered with radiating rows of
elongated, rectangular, transparent scales of about 100 x 20 pm; marginal spicules
slender, cylindrical or nearly cylindrical to about 220 pm long.

Radula (Fig. 14) 8 mm long and bearing about 33 mature rows of teeth;
rachidian tooth relatively small, oblong, about 108 um long, working edge about
70 ym wide; blade of major lateral teeth bicuspid, the inner cusp much longer
than the outer one.

Gills holobranchial, adanal, about 30 per side in specimens 24 mm or greater
in length.

Range of morphological variation: In some specimens the granules on the ribs
and sometimes the ribs themselves are obsolete. Although most specimens are
whitish to tan with speckles and/or concentric rings of reddish-brown, in a few
specimens the reddish-brown is replaced with pinkish or pale lavender; some
specimens are uniform whitish.

Type locality: Eider Point, west side of entrance to Unalaska Bay, Bering Sea side
of Unalaska Island, Aleutian Islands, Alaska (53°57'40"N, 166°35'30"W).

Material Examined: Type material: Holotype, LACM 2796 and 100+ paratypes
(leg. RNC, 21 June 1993); other paratypes: 10, LACM 2797; 4, USNM 880161;

October 25, 2000 Clark 9

4, CAS 105915; 4, SBMNH 143146; 4, ZISP 1935; 3, UAF Mo-5570; 60+, RNC
1230; 1, H.L. Strack Coll.; 2, B. Dell Angelo Coll.; 4, H. Saito Coll.; 3, R-A. Van
Belle Coll.; 2, D.J. Eernisse Coll.; 2, OS&S Coll.

Additional material: 2, LACM 152676; and 6, RNC 3160, Channel at south
end of Amacknak Island (Dutch Harbor), Unalaska Bay, Unalaska Island, Alaska,
10-15 m.

Distribution: Known so far from only two localities within Unalaska Bay,

Unalaska Island, Aleutian Islands, Alaska (Fig. 30).

Habitat: Lives at 0-15 m on cobbles encrusted with bryozoans, resting on or
slightly buried in sand.

Remarks: This species somewhat resembles Lepidozona ima (Figs. 23-24) but may
be distinguished by the unique color patterns of the valves, larger (dorsal) girdle
scales, which bear many more striations (22-25 compared to 12-14 in L. ima), and
relatively smaller (central and minor lateral) teeth of the radula.

Lepidozona (Tripoplax) attuensis new species

(Figs. 15-21)

Diagnosis: Chitons of small size (to 2.5 cm), oval in outline; valves whitish with a
few faint, rusty-orange, longitudinal markings; head valve, lateral areas, and post-
mucronal area of tail valve with numerous, low, rounded, radiating ribs, lacking
pustules or granules. Girdle with relatively large, smooth, imperceptibly
mammnillated scales.

Description: Body (Fig. 15) oval in outline; valves granular, subcarinated,
unbeaked, side slopes slightly convex; tegmentum whitish (holotype) or whitish
with pale rust sub-jugal stripe and a few faint rust mottlings on the central areas;
girdle uniformly whitish.

Head valve (Fig. 16) semicircular, posterior margin widely V-shaped; bearing
23-26 low, rounded, radiating ribs; insertion plate with 15-17 slits.

Intermediate valves (Fig. 17) oblong, central areas with fine pitting; lateral
areas slightly raised, defined by three to four low, rounded ribs (nearly obsolete on
some valves of the holotype) crossed by concentric growth lines; insertion plates
with 2 slits.

Tail valve (Fig. 18) semicircular, mucro central, post-mucronal slope slightly
convex; post-mucronal area bearing about 14 low, rounded, well spaced radiating
ribs or undulations; insertion plate with 14-15 slits.

Interior of valves (articulamentum) white; sutural laminae fairly short,
rounded at edges, connected across the jugal sinus by a concave jugal plate,
sometimes notched at sides; insertion teeth very short and blunt, those on the
head valve smooth, the ones on the tail valve rugose on the anterior side.

10 NEMOURIA No. 42

ee

Figures 15-21. Lepidozona (Tripoplax) attuensis new species. 15-18. Paratype RNC
317; 12.0 mm x 12.0 mm. 15. Whole animal (note: valve five is lost, this is not an
aberrant, seven-plated specimen). 16. Head valve, width 8.0 mm. 17. Intermediate
valve four, width 9.0 mm. 18. Tail valve, width 6.5 mm. 19-21. Holotype, LACM
2801. 19, 20. Dorsal girdle scales. 21. Radula.

October 25, 2000 Clark 11

Girdle about 1/4 as wide as fifth valve at sides, much narrower at ends;
dorsal side covered with smooth, bent, proximally rounded scales (Figs. 19 & 20),
about 275 pm long and 150 pm wide; ventral surface covered with radiating,
rectangular, transparent scales of about 100 x 20 um; marginal spicules slender,
bluntly pointed, about 150 pm in length.

Radula (Fig. 21) 7.8 mm long (in the 21 mm holotype) and bearing about 30
mature rows of teeth; rachidian tooth oblong, about 265 pm long, constricted in
the middle to about 60 tm and broadening again at the base, working edge about
130 um wide; blade of major lateral teeth bicuspid, the inner cusp about twice as
long as the outer one.

Gills holobranchial, adanal, 26 per side in the holotype (curled length ca. 23
mm) and 25 per side in the paratype (21 mm long).

Type locality: Murder Point, Massacre Bay, Pacific side of Attu Island, Near
Islands, Aleutian Islands, Alaska (52°47.6'N, 173°11.0'E).

Material Examined: Type material: Holotype, LACM 2801 and one paratype,
RNC 317 (leg. David R. Lindberg, 20 July 1979; intertidal).

Distribution: Known so far only from the type material.

Remarks: This species is very similar to Lepidozona ima (Figs. 23-24), especially in
the characters of the radula. L. attuensis differs from L. ima by the prominent
radial ribs on the valves, which lack pustules, the larger, smooth, differently
proportioned girdle scales, and fewer gills. Also L. ima is usually found much

deeper (100-250 m).
DISCUSSION

The chiton fauna of the Aleutian Islands is very rich and diverse, containing
35 known species [including one as yet undescribed species (unpublished notes)].
The genus Lepidozona with seven species makes up 20% of this fauna. Five of
these (71%) are endemic to the Aleutian faunal province. Two of these L.
(Tripoplax) attuensis and L. (T.) baxteri appear to be short range endemics, restricted
to a single island or island group. These species appear to be divisible into two
(natural?) groups based on their dorsal girdle scales. Group one contains species
with mammillated scales, L. (T.) allyni and L. (T.) beringiana. Group two contains
species with non-mammillated scales, L. (T.) abyssicola, L. (T.) ima, L. (T.) baxteri, L.
(T.) attuensis, and L. (T.) trifida.

The radula of Lepidozona (Tripoplax) allyni (Fig. 25) described from Amchitka
Island, in the Aleutians was described and illustrated (drawn) as having unicuspid
major lateral teeth. However, a study of the paratypes as well as another specimen

iz NEMOURIA No. 42

Figures 22-26. Lepidozona spp. 22-24 Lepidozona (Tripoplax) ima Sirenko, 1975. 22.
Whole animal, RNC 3172 (Stalemate Bank, W of Attu Id., 114 m); 18.5 mm x
10.5 mm. 23-24. Specimen from near Cooper Id., Commander Is., Russia, 130-
200 m; RNC 2363. 23. Dorsal girdle scales. 24. Radula. 25-26. Lepidozona
(Tripoplax) allyni (Ferreira, 1977). 25. Whole animal. Amchitka Id., 25 m
(topotype), AB 73-25. 26. Radula. Paratype, CAS 016564.

October 25, 2000 ig

Y
Yyy

Y
Wyn
yy

Oy
MON

29

Figures 27-29. Lepidozona spp. 27. L. (Tripoplax) lindbergi (Yakovleva, 1952), RNC
120; whole animal, Simushir Id., Kurile Is., Russia, 50-60 m; curled, ca. 20.0 mm.
28. L. (T.) abyssicola (Smith & Cowan, 1966), RNC 3106; whole animal, near
Farallon Is., California, 2750 m; curled, ca. 50 mm. 29. L. (T.) trifida (Carpenter,
1864); RNC 2417; whole animal, Ketchikan, Alaska, 25 m; 35 mm x 24 mm.

taken with the types (but apparently not seen by Ferreira) in the collection of the
National Marine Fisheries Service, Auke Bay, Alaska Laboratory (AB 73-25)
revealed that the major lateral teeth of this species are in fact bicuspid (Fig. 26). It
is not known whether Ferreira erred in his description, or if the radula of the
holotype was aberrant, because the slide containing the holotype’s radula could

14 NEMOURIA No. 42

1 ALASKA 2”

s y —
or
f gv"

COMMANDER.
Be LN LAND:

Hie I "a eS D S Linaias - an
: AN 1st S Wmnak “ps Sanak
_ Yunaska ve

on Seguam | 4 7%
3 \ Sha@nya eae |x
Nizki @2@. te Sage os Chuginadak
Kiska ®e Atka
P ACU SE Cm OCB AUN.

Unimak has A
\ suv hy.
A

e.
Amchitka Atha
Oe 170° 160° eae

Distribution of Lepidozona (Tripoplax )
in the Aleutian Islands.

[a] L. abyssicola (o] L. trifida fx) L. allyni fe L. ima {e] L. beringiana
+4] L. baxteri fe] L. attuensis

Figure 30. Map of Aleutian Islands showing distributions of Lepidozona species.

not be found (E. Kools, pers. comm., 18 March, 1996). However, due to the
bicuspid condition of the paratypes and other material, Ferreira’s description
must be amended.

Three specimens of Lepidozona (T.) allyni were collected at a depth of 15 m off
the point at the east side of the entrance to Crescent Bay, Atka Island, Aleutian
Islands (52°02'15"N, 175°14'00"W), on the bottom of stones resting on sand (leg.
RNC, 8 July 1997). One of these has been deposited as a voucher (LACM
152584); the remaining two were retained (RNC 3026). Additionally a single
curled juvenile (length ca. 6.0 mm) taken at 18 m at Kiska Island, Aleutian
Islands, was found in the collection of the United States National Museum
(USNM 208587). The new records extend the known range of the species
approximately 125 km to the west, and 375 km to the east.

As already discussed in the species descriptions, Lepidozona (Tripoplax) ima is
morphologically similar to both L. (T.) baxteri, and L. (T.) attuensis, but differs in
the details of the valve sculpture, girdle scales and radula. L. (T.) ima and its girdle
scales and radula are illustrated in Figs. 22-24.

Smith G@ Cowan (1966: 8) described the tegmental coloration of Lepidozona
(Tripoplax) abyssicola (Fig. 28) as dark reddish-brown. However this description is
misleading, because the reddish-brown (or blackish) appearance is due to deep sea
deposits present on most specimens. The tegmentum of this species is white. The
range of L. (T.) abyssicola is here extended west into the eastern Aleutian Islands,
to south of Samalga Pass (west end of Umnak Island) (52°36.2'N, 169°25.2'W)
based on a single specimen (CAS 120748), trawled, F/V Morning Star, 199 m on
rocks (NMFS Gulf of Alaska Trawl Survey, 57-991-3). The new record extends the

known range approximately 825 km to the west.

October 25, 2000 Clark 15

Lepidozona (Tripoplax) trifida is also found in the (eastern) Aleutians, and
although it probably would not be confused with any of the other members the
genus due to the unique sculpturing of its valves [broad, flattened radial ribs
(three on lateral areas) separated by distinct sulci], it is illustrated for comparison

(Fig. 29).
ACKNOWLEDGMENTS

For their help in various aspects of this work I thank James H. McLean
(LACM), Darlene H. Southworth (SOU), Boris I. Sirenko (ZISP), Elizabeth Kools
(CAS), Bruce L. Wing (NMFS-AB), Nick Hindman (Dutch Harbor, Alaska), Eric
Brown, Robin Harrison, Robert Lauth, Mike MacEwan and Michael Martin
(NMFS, Seattle), the captains and crew of the F/V Dominator and the F/V
Vesteraalen, and Douglas J. Eernisse (California State University at Fullerton) for
critically reading the manuscript.

LITERATURE CITED

Baxter, R. 1987. Mollusks of Alaska. Shells and Sea Life Inc., Bayside, California.
163 pp.

Berry, S.S. 1917. Notes on west American chitons-I. Proceeding of the California
Academy of Sciences, Fourth Series, 7(10): 229-248.

Berry, $.S. 1919. Notes on west American chitons II. Proceedings of the California
Academy of Sciences, Fourth Series, 9(1): 1-36, pls. 1-8.

Carpenter, P.P. 1864. Supplementary report on the present state of our
knowledge with regard to the Mollusca of the west coast of North America.
Report British Association Advancement Science, 517-686.

Clark, R.N. 1991. Notes on the distribution, taxonomy and natural history of
some north Pacific chitons (Mollusca: Polyplacophora). The Veliger, 34(1): 91-
96.

Ferreira, A.J. 1977. A new species of chiton from the Aleutian Islands (Mollusca:
Polyplacophora). The Veliger, 20(1): 27-29, figs. 1-5.

Gray, J.E. 1847. Additional observations on chitons. Proceedings of the Zoological
Society of London, 15: 126-127.

Kaas, P. & R.A. Van Belle. 1987. Monograph of Living Chitons. Vol. II. Dr.
Backhuys Publisher, Rotterdam. 302 pp.

Middendorff, A.T. von. 1847. Beitrage zu einer Malacozoologia Rossica. Mem. Sci.
nat. Acad. imp. Saint Petersburg, 6(1): 1-151, pls. 1-14.

Pilsbry, H.A. 1892-1894. Polyplacophora. In: G.W. Tryon, Jr. (ed.) Manual of
Conchology. 14(53-54): 1-128, pls. 1-30 (1892); 14(55-56): 129-350 + ixxxiv,

16 NEMOURIA No. 42

pls. 31-68 (1893); 15(57): 1-64, pls. 1-10 (1893); 15(58): 65-132, pls. 11-17
(1894).

Shuttleworth, R.J. 1853. Diagnosen neur Mollusken. 4, Ueber den Bau der
Chitoniden, mit Aufzahlung der die Antillen und die Canarischen Inseln
bewohnenden Arten. Mitt. naturf. Ges. Bern, 286-291: 169-207.

Sirenko, B.I. 1975. On the taxonomy of the genus Lepidozona Pilsbry. Biologya
Belogo Morya Moskva, 3: 13-28, figs. 1-6 (in Russian).

Smith, A.G. @ IMcT. Cowan. 1966. A new deep-water chiton from the
northeastern Pacific. Occasional Papers of the California Academy of Sciences, No.
56, 15 pp.

Yakovleva, A.M. 1952. Shell bearing mollusks (Loricata) of the seas of USSR. Keys
to the fauna of USSR. Academy of Sciences, USSR, no. 45, 127 pp. [English

translation, Jerusalem, 1965].

Founding Editor Managing Editor
JOHN E. DU PONT TIMOTHY A. PEARCE

Consulting Editors

M.G. HARASEWYCH, National Museum of Natural History, Smithsonian
Institution, Washington, DC.

KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh, PA.

RICHARD W. THORINGTON, JR., National Museum of Natural History,
Smithsonian Institution, Washington, DC.

Published by the

DELAWARE MUSEUM OF NATURAL HISTORY
P.O. Box 3937, Wilmington,
Delaware 19807-0937, U.S.A.

(_ Nov 1 6 200

LIBRARIES

Occasional Papers of the Delaware Museum of Natural History

NUMBER 43 OCTOBER 30, 2000

THE CHITON FAUNA OF THE GULF OF CALIFORNIA
RHODOLITH BEDS (WITH THE DESCRIPTIONS OF FOUR
NEW SPECIES)

Roger N. Clark*

ABSTRACT. The shallow water rhodolith beds of the Gulf of California have a rich
and diverse fauna. However, there are no published studies the chiton fauna of
this unique habitat. The present paper reports on eight species of chitons (four
new to science) found living on the rhodoliths: Stenoplax mariposa (Bartsch MS,
Dati, 1919), Callistochiton sp., cf. C. elenensis (Sowerby, 1832), Lepidochitona beanii
Carpenter, 1857, Acanthochitona avicula (Carpenter, 1864), Lepidochitona corteziana
new species, Ischnochiton rhodolithaphilus new species, Ischnochiton tomhalei new
species, and Acanthochitona burghardtae new species.

Key words: Chiton, Gulf of California, rhodoliths.
INTRODUCTION

Rhodoliths are free-living, often branched, nongenticulate coralline red algae
found in dense concentrations (or beds) worldwide, from the low intertidal zone
to over 100 m depth (Bosence, 1983). Extensive shallow water (2-12 m) rhodolith
beds of the southwestern Gulf of California were discussed by Bosence (1983) and
their ecological parameters defined. Steller G Foster (1995) and Foster et al.
(1997) further documented the rich biodiversity associated with these specialized
habitats.

* Associate, Los Angeles County, Museum of Natural History Serial Publication
900 Exposition Boulevard, Los Angeles, California ISSN 0085-3887
90007, U.S.A., email: insignis@cdsnet.net

Mailing Address: 1839 Arthur Street, Klamath Falls, Oregon 97603-4617

4,
|

2 NEMOURIA No. 43

In April of 1995, Mr. Barry F. Putman of Cuesta College in San Luis Obispo,
California sent me his rough manuscript, which described an apparently new
species of chiton from rhodolith beds at Isla El Requeson, Bahia Conception,
Baja California Sur. Mr. Putman requested that I re-write the manuscript, and
submit it for publication. He also put me in contact with Dr. Michael S. Foster of
Moss Landing Marine Laboratories (California). Dr. Foster provided 30+
additional chiton specimens collected at various localities between Punta Bajo and
La Paz, from November 1994 to April 1995 by Mr. Marco Medina-Lopez, a
student at Universidad Autonoma de Baja California Sur (La Paz). All specimens
were among the often closely spaced branches of individual rhodoliths as part of
their diverse cryptofauna. This paper reports the study of these specimens, as well
as three additional specimens collected at Punta Chivato in March of 1998, and
received subsequently from Dr. Foster. The map in figure 26 shows the collection
sites.

MATERIALS AND METHODS

Numerous very small chitons (2-10 mm) were collected from rhodoliths taken
with SCUBA at depths of 4-12 m at several locations along the SE shore of Baja
California Sur, and preserved in 70% ethanol. The specimens were studied using
a light and a scanning electron microscope (SEM), and their characters compared
with species known to occur in the region (Skoglund, 1997; Kaas and Van Belle,
1985, 1990, 1994).

Specimens were prepared for SEM examination by boiling whole in 10%
KOH and separating out the radula, valves, and epidermal layers of the girdle.
The valves were air dried, and the radula and girdle fragments were dehydrated in
an acetone series, rinsed, dried and mounted on SEM viewing stubs. The
specimens were then sputter coated for 2 minutes with gold-palladium and
examined with at 10-15 kv with a Hitachi 2100 S SEM at the Department of
Biology at Southern Oregon University, Ashland, Oregon.

Whole animals were photographed using a Nikon SLR camera with macro
lens and bellows attachment.

Abbreviations used in the text are LACM, Los Angeles County Museum of
Natural History; CAS, California Academy of Sciences; SBMNH, Santa Barbara
Museum of Natural History; RNC, Private collection of the author.

RESULTS

Eight species of chitons were identified, representing four families and four
genera, including four species new to science. The species are described and

October 30, 2000 Clark 3

illustrated herein, and are compared to similar species that co-occur in the Gulf of
California region.

SYSTEMATICS
The systematic arrangement in this paper follows the recent system of
revisions proposed by Sirenko (1993,1997).

Class: POLYPLACOPHORA Gray, 1821
Order: CHITONIDA Theile, 1910
Suborder: CHITONINA Theile, 1910
Superfamily: CHITONOIDEA Rafinesque, 1815
Family: ECHNOCHITONIDAE Dall, 1889
Genus: Ischnochiton Gray, 1847
I. rhodolithophilus new species
I. tomhalei new species
Genus: Stenoplax Carpenter in Dall, 1879
S. mariposa (Bartsch MS, Dall, 1919)
Family: CALLISTOPLACIDAE Pilsbry, 1892
Genus: Callistochiton Carpenter in Dall, 1879
C. sp. cf. elenensis (Sowerby, 1832)
Suborder: ACANTHOCHITONINA Bergenhayn, 1930
Superfamily: MOPALIOIDEA Dall, 1889
Family: TONICELLIDAE Simroth, 1894
Genus: Lepidochitona Gray, 1841
L. beanii Carpenter, 1857
L. corteziana new species
Superfamily: CRYPTOPLACOIDEA H. & A. Adams, 1858
Family: ACANTHOCHITONIDAE Pilsbry, 1893
Genus: Acanthochitona Gray, 1821
A. avicula (Carpenter, 1864)
A. burghardtae new species

Stenoplax mariposa (Bartsch MS, Dall, 1919)
(Figs. 1 G 2)

Three specimens (2.8-10 mm in length) (RNC 3130) of this common Gulf
species were taken at El Pardito (leg. Marco Medina-Lopez, SCUBA,12 m, 7 April,
1995). All specimens appear typical, and although the tiny juveniles are nearly
identical in color pattern (variegated with brown, black, pale green and white,
with brilliant blue spots on the pleural areas) with Ischnochiton rhodolithophilus,

4 NEMOURIA No. 43

yy

yj
Wi’
Y

Figures 1-5. Chiton species. 1-2. Stenoplax mariposa (Barstch MS, Dall, 1919), RNC
3130. 1. Whole animal, length 4.2 mm. El Pardito. 2. Valve fragment from 10
mm specimen. El Pardito. 3. Callistochiton sp., cf. C. elenensis (Sowerby, 1832),
RNC 3131. First, fourth and eighth valves. Valve eight width 3.8 mm. Canal de
San Lorenzo. 4. Lepidochitona beanii Carpenter, 1857, RNC 3132. Whole animal,
length 3.0 mm. Canal de San Lorenzo. 5. Acanthochitona avicula (Carpenter,

1864), RNC 3133. Whole animal, length 7.0 mm. Coronados Id.

October 30, 2000 Clark 5

they are easily distinguished by the ribbed central areas, and rugose lateral areas of
the valves.

Ischnochiton rhodolithophilus Putman MS, Clark, new species
(Figs. 6-10)

Diagnosis: Very small chitons (to 6.0 mm), elongate-oval in outline, elevated;
valves roundly arched, moderately beaked; tegmentum mostly smooth, but with
some quincuntial pitting on pleural areas, color variegated, dark-brown and cream
with blue spots on terminal and pleural areas; sutural laminae short, bluntly
triangular; girdle medium wide, with short, wide scales (to 100 pm x 60 pm) with
12-18 deep incisions on middle 1/3; radula with heavy, tricuspid major lateral
teeth.

Description: Very small chitons (to 6.0 mm), elongate-oval in outline, moderately
elevated; valves roundly arched, moderately beaked (Fig. 6); tegmentum mostly
smooth, with some pitting (Fig. 7); articulamentum thin, white, slit formula 8-1-8;
color, variegated with dark brown and cream, terminal valves and pleural areas of
intermediate valves with brilliant blue spots.

Head valve semicircular, slightly convex, posterior margin widely v-shaped;
insertion teeth short, sharp, fairly thick; intermediate valves rectangular, about
1/3 as long as wide; weakly beaked, slightly indented at jugal sinus; lateral areas
poorly defined, slightly if at all raised, smooth; jugum smooth, pleural areas
quincuntially punctate (to varying degrees, depending on size of specimen)
especially on the lower portions; sutural laminae fairly long, narrow, bluntly
triangular; jugal sinus very wide; insertion teeth short, thin, sharp; slit ray
grooved, porous. Tail valve about 3/4 the width of the head valve, anterior
margin slightly convex; mucro central, post-mucronal slope concave; ante-
mucronal area quincuntially punctate, terminal area smooth; sutural laminae
fairly long, rounded; jugal sinus moderately wide; insertion teeth short, relatively
thick, sharp.

Girdle moderately wide, about 1/4 as wide as valve 5, clothed dorsally with
relatively wide, bluntly rounded, bent scales 70-100 tm wide x 40-60 pm long,
with 12-18 narrow ribs on the central 1/3, deeply incised between (Figs. 8 & 9);
ventral scales rectangular, about 38 pm x 18 pm (Fig. 9); marginal spicules broad,
bluntly pointed, 50 pm x 20 pm, with 5-6 oblique striations (Fig. 9).

Radula (Fig. 10, paratype 1) curled, ca. 5.0 mm long, 1.2 mm long with about
35 mature rows of teeth; central tooth very small, slender, elongated, slightly
dilated distally, about 20 pm long, distal end about 5 um wide; minor lateral teeth
wing shaped, about 40 pm x 15 pm; major lateral teeth heavy, rugged, tridentate,
deeply (and rather widely) indented between, about 30 pm x 25 jm; denticles very

6 NEMOURIA No. 43

1G0um

O

7

Mili:

Y

y

Figures 6-10. Ischnochiton rhodolithaphilus Clark, spec. nov. 6. Whole animal,
length 3.9 mm. Paratype, RNC 3134, Isla San Jose. 7. Intermediate valve five
(scale on figure), Paratype, CAS 098787, Bahia Concepcion. 8. Dorsal girdle
scales (scale on figure), Paratype, RNC 3138, Punta Bajo. 9. Girdle scales (scale on
figure), Paratype, RNC 3138. 10. Radula (scale on figure), Paratype, RNC 3138.

October 30, 2000 Clark 7

blunt, finger-like in appearance, inner one the longest, central one slightly shorter,
the outer one about 1/2 as long as the inner one.
Ctenidia about 12-15 per side in specimens 2.8-5.0 mm in length.

Type Locality: Gulf of California, Isla El Requeson, Bahia Conception, Baja
California Sur, Mexico (26°43'N, 111°46'W); 4-9 m.

Type Material: Holotype (CAS 098786) and 22 Paratypes (leg. Michael S. Foster,
SCUBA, January 1990) (10, CAS 098787); (10, Barry Putman, private collection);
(2,RNC 3134).

Additional Material: 10 additional specimens, 3, RNC 3148, Canal de San
Lorenzo, 12 m (leg. Marco Medina-Lopez, SCUBA, 2 February, 1995); 1, RNC
3149, S side of SW tip of Isla San Jose, 12 m (leg. Marco Medina Lopez,
SCUBA, 10 February, 1995); 4, RNC 3150, Punta Bajo, 12 m (leg. Marco Medina-
Lopez, SCUBA, 11 November, 1994); 1, RNC 3151, El Pardito, N of Isla San
Francisco, 12 m (leg. Marco Medina-Lopez, SCUBA, 7 April, 1995); 1, RNC
3152, N side of SW tip of Isla San Jose, 5 m (leg. Marco Medina-Lopez, SCUBA,
10 February 1995).

Distribution: Gulf of California, SE coast of Baja California Sur from the type
locality, S to Canal de San Lorenzo, about 40 km N of La Paz (24°2I1'N,
111°15'W); 4-12 m.

Discussion: Ischnochiton rhodolithophilus is very similar to Stenoplax mariposa
(Bartsch MS, Dall, 1919), and might be mistaken for a juvenile of that species,
however it may be distinguished by 1) lack of ribbing or lirae on valves; 2) less
numerous, heavier ribs on the girdle scales, 12-18 instead of 45-50. Both species
may be found together in the rhodolith habitat.

Etymology: The specific name is derived from the Latin rhodo (red) + the Greek
lithos (stone) and philia (love); the lover of red stones (rhodoliths).

Ischnochiton tomhalei new species

(Figs. 11-14)

Diagnosis: Very small chitons (to 4.0 mm), oval in outline, valves roundly arched,
not carinated, only slightly beaked; tegmentum reticula-granular; mucro of tail
valve at ante-central, post-emucronal slope straight to slightly concave; girdle
medium wide, about 1/3 of valve 5 width, dorsal scales with 12-16 strong ribs;
radula with relatively heavy, tricuspid major lateral teeth.

Description: Very small chitons (to 4.0 mm), oval in outline, valves roundly
arched, not carinated, barely if at all beaked (Fig. 11); tegmentum reticula-
granular; slit formula 9-1-8; color reddish-brown to light brown or cream, with
white flame-like markings, especially at the valve apices.

8 NEMOURIA No. 43

Figures 11-14. Ischnochiton tomhalei Clark, spec. nov. 11. Whole animal, length 4.0
mm, RNC 3135, Isla San Jose. 12. Intermediate valve five (scale on figure),
Holotype, LACM 2877. 13. Dorsal girdle scales (scale on figure), Holotype. 14.
Radula (scale on figure), Holotype.

Head valve semicircular, slightly convex, posterior margin widely v-shaped;
sculpture, evenly reticula-granular; insertion teeth fairly short, wide and thick.
Intermediate valves (Fig. 12) rectangular, about 1/3 as long as wide; lateral areas

October 30, 2000 Clark 9

poorly defined, slightly raised, often with faint growth costae; central areas with
reticula-granular sculpture, becoming obsolete at valve apices; articulamentum
thin, white; sutural laminae rather short, narrow, rounded, jugal sinus very wide;
insertion teeth short, thick, triangular. Tail valve slightly wider than long, inflated,
anterior margin convex, mucro ante-central, post-mucronal slope straight to
slightly concave; ante-mucronal area reticula-granular, posterior area granular;
insertion teeth short, thick.

Girdle moderately wide, about 1/3 the width of valve 5, clothed dorsally with
broad, short, bent scales about 60-75 um x 32 pm, with 12-16 fairly narrow ribs,
deeply incised between (Fig. 13); ventral scales very small, rectangular about 25
uum x 12 pm; marginal spicules relatively short, broad, flattened, bluntly pointed,
about 40 um x 12 um, with three oblique striations.

Radula (Fig. 14) about 800 pm long, with 20-25 mature rows of teeth; central
tooth slender, elongated about 25 pm long, distal end dilated, working edge about
4 um wide; wing shaped, elongate about 25 pm x 10 pm; major laterals fairly
heavy, tricuspid, central cusp slightly longer than lateral two; cusps rather short
and broad.

Ctenidia 12-16 per side in specimens 2.8-4.0 mm in length.

Type Locality: Gulf of California, Punta Bajo (about 10 km N of Loreto), Baja
California Sur, Mexico (approx. 26°05'N, 112°42'W); 12 m.

Type Material: Holotype (LACM 2877), and one paratype (RNC 3142) (leg.,
Marco Medina-Lopez, SCUBA, 11 November, 1994).

Additional Material: 1, RNC 3139, Punta Chivado, 10-12 m (leg. Michael S.
Foster, SCUBA, 18 March, 1998); 3, RNC 3140, Canal de San Lorenzo, 12 m
(leg. , Marco Medina-Lopez, 2 February, 1995); 1, RNC 3141, W side of Isla
Coronados, 7 m (leg. , Marco Medina-Lopez, SCUBA, "December, 1994"); 1,
RNC 3153, S side of SW tip of Isla San Jose, 12 m (leg., Marco Medina-Lopez,
SCUBA, 10 February, 1995).

Distribution: Gulf of California, east coast of Baja California Sur, Mexico, from
Punta Chivado (about 25 km N of Mulege)(27°05'N, 112°05'W) to Canal de San
Lorenzo, about 40 km N of La Paz (24°21'N, 111°15'W); 7-12 m.

Discussion: The only eastern Pacific iscnochitonid that Ischnochiton tomhalei is
likely to be confused with is juveniles of I. tridentatus Pilsbry, 1893, which (rarely)
may be similarly colored. However, the coarser sculpture and fewer slits in the
insertion plates (8-1-9 instead of 12-18/ 2/ 11-13 in I. tridentatus) will distinguish
them.

Etymology: It is with great pleasure that I name this species after Mr. Thomas S.
Hale of Portland, Oregon, a long time shell collector who for many years has

10 NEMOURIA No. 43

educated and inspired a great many people in the sciences of conchology and
marine biology.

Callistochiton sp. cf. C. elenensis (Sowerby, 1832)
(Big. 3)

A single small (9 mm) specimen (RNC 3131) taken at Canal de San Lorenzo
(leg. Marco Medina-Lopez, SCUBA, 12 m, 2 February 1995) is tentatively
identified as a juvenile of this species, but may represent an undescribed species.
The valves are more highly elevated and the side-slopes more convex than in
specimens of C. elenensis that I have examined. Also I can detect no jugal laminae,
another character present in (adult) C. elenensis , but this may be due to the size of
the specimen. The valves are rust colored with dark brown specks. More material
is required before a solid conclusion can be made on its identification.

The genus Callistochiton Carpenter, in Dall, 1879 in the tropical west
American region is in need of revision. The views of Ferreira (1979) and Kaas and
Van Belle (1994) differ on several points, and there appear to be more species in
the region than are currently recognized.

Lepidochitona beanii Carpenter, 1857
(Fig. 4)

Two tiny specimens (3.0 & 4.0 mm long)(RNC 3132) of this widespread
species were taken at El Bajo (leg. Marco Medina-Lopez, SCUBA, 14 m, 30 March
1995). The specimens are typical in all characters, and are readily distinguished
from the similar Lepidochitona corteziana by the presence of large (to 350 ym long)
hyaline spicules on the girdle.

Lepidochitona corteziana new species

(Figs. 15-19)

Diagnosis: Very small chitons (to 3.5 mm); roundly oval in outline; tegmentum
microscopically lirate; intermediate valves beaked, about 1/4 as long as wide; tail
valve small, about 1/2 as long as wide, mucro prominent, at anterior 1/3, post
mucronal slope concave; girdle medium wide, fleshy, bearing no large spicules.

Description: Very small chitons (to 3.5 mm); roundly oval in outline; valves
subcarinate, beaked, slopes slightly convex (Fig. 15); tegmentum very spongy;
microscopically lirate, lirae low, broad, flattened and bearing a_ nearly
imperceptible medial sulcus; slit formula 10-1-12; color light orange to pink-
orange.

Head valve semicircular, convex, bearing about 80+ radial micro-lirae;

October 30, 2000 Clark iat

posterior margin widely v-shaped, insertion teeth fairly short, thick. Intermediate
valves (Figs. 16,17) rectangular, central areas with 120 + micro-lirae; lateral areas
barely defined, bearing about 20-25 micro-lirae; articulamentum (Fig. 17) thin,
white; sutural laminae relatively short, narrow, roundly triangular, jugal sinus

Figures 15-19. Lepidochitona corteziana Clark, spec. nov. 15. Whole animal, length
3.0 mm, Paratype, RNC 3136, Canal de San Lorenzo. 16. Intermediate valve five
(scale on figure), Holotype, LACM 2878. 17. Interior of valve five. 18. Close-up of
valve five tegmentum, showing esthete pores. Holotype. 19. Girdle (scale on
figure), Holotype.

Its NEMOURIA No. 43

wide, porous; insertion teeth short, broad, thick, bounded on both sides by
porous slit rays. Tail valve small, about 1/2 as long as wide, low; mucro
prominent, located at anterior 1/3, post-mucronal slope concave; ante-mucronal
area with about 60 longitudinal micro-lirae; post-emucronal area with about 60+
radial lirae; sutural laminae short, fairly wide, anterior. edge straight, jugal sinus
wide.

Macresthetes round, about 7 um in diameter, located in single, linear row,
atop the microlirae, each separated by two micresthetes about 3.5-4 pm in
diameter; micresthetes also located in single linear row, separate by about one
diameter, in grooves between micro-lirae (Fig. 18).

Girdle medium wide (somewhat less than 1/2 of valve 5 tegmentum), fleshy
appearing (Fig. 19); dorsally clothed with tiny, close-set, erect, pointed spicules,
about 15 pm x 5 pm; ventral surface of girdle covered with very minute, bluntly
rounded granules measuring about 5 um x 3 pm; marginal spicules about 50-60
um in length, and up to about 12.5 ym in width, relatively slender, elongated,
flattened, bluntly pointed or slightly broadened and rounded at the distal end.

Radula unknown.

Type Locality: Gulf of California, Canal de San Lorenzo, about 40 km N of La
Paz, Baja California, Mexico (24°21'N, 111°15'W); 12 m.

Type Material: Holotype (LACM 2878) & three Paratypes (1, LACM 2879); 2,
RNC 3154 (leg. , Marco Medina-Lopez, SCUBA, 2 February, 1995).

Additional Material: Two (RNC 3143), Punta Chivado, 10-12 m (leg.
Michael S. Foster, 18 March, 1998); two (RNC 3144), S side of SW tip of Isla San
Jose, 12 m (leg. Marco Medina-Lopez, SCUBA, 10 February, 1995); RNC 3136,
Punta Bajo, 12 m (leg. Marco Medina-Lopez, SCUBA, 11 November, 1994).

Distribution: East coast of Baja California Sur, from Punta Chivado (about 25
km N of Mulege) (27°05'N, 112°05'W) S to the type locality at depths of 10-12 m.

Discussion: Lepidochitona corteziana bears a close resemblance to Lepidochitona
beanii Carpenter, 1857, of the same size, with which it may occur, but may be
easily distinguished by 1) the sculpture of the valves, and 2) the lack of large
hyaline needles (up to 250 x 30 im long) on the girdle (especially bunched at the
sutures). It also shows affinities with the South African L. turtoni (Ashby, 1928),
which likewise has fine radial micro-sculpture and a secondary slit ray behind the
insertion tooth. The overall sponginess of the valves, especially the
articulamentum with its multiple slit rays, and the armature of the girdle
(although greatly reduced dorsally in L. corteziana) shows affinities with
Spongioradsia aleutica (Dall, 1878) from the Aleutian Is. Spongioradsia aleutica also
has a coralline algal habitat.

October 30, 2000 Clark 13

Etymology: Named for the Sea of Cortez (Gulf of California) where the species
lives.

Acanthochitona avicula (Carpenter, 1864)
(Fig. 5)

Two specimens of this small species were taken, one at El Pardito, 12 m
(RNC 3145)(leg. Marco Medina-Lopez, 7 April, 1995), and one at Coronados Id.,
7 m (RNC 3146)(leg. Marco Medina-Lopez, "Winter" 1994-95). The specimens

measure 7.5 mm and 7.0 mm in length respectively and appear typical.

Acanthochitona burhardtae new species

(Figs. 20-25)

Diagnosis: Very small chitons (to 3.5 mm), broadly oval in outline, intermediate
valves twice as wide as long; lateral areas barely raised; tegmentum with coarse,
close-set, flattened granules; granules of the central areas arranged in 15-17
longitudinal rows; girdle fleshy, with scattered spicules (to 350 pm in length).

Description: Very small chitons (up to 3.5 mm), broadly oval in outline (Figs.
20,21); tegmentum consisting of coarse, flattened, close-set granules; slit formula
5-1-2; color of tegmentum light orange, or white with brown markings.

Head valve semi-circular, posterior margin broadly v-shaped; insertion teeth
short, wide, fairly thick. Intermediate valves (Fig. 22) about 1/2 as long as wide,
beaked; granules arranged in longitudinal rows on central areas (often obsolete at
the jugum), with distinct grooves between; lateral areas poorly defined, slightly
raised and bearing smaller granules than central areas; granules of jugal areas
more or less obsolete; articulamentum thin, white; sutural laminae short, fairly
wide; jugal sinus wide; insertion teeth very short. Tail valve (Fig. 23) small, rather
oval shaped, mucro at posterior 1/3, post-emucronal slope straight or slightly
convex; insertion teeth fairly short, thick, with medial sinus; sutural laminae fairly
long, extending well around anterolateral edge, jugal sinus fairly wide.

Macresthete sub central, oval about 7-8 um x 5 pm; micresthetes six (three
per side) in linear rows, about the same size as macresthete (Fig. 24).

Girdle (Fig. 25) of medium width, about 1/4 the width of valve five
tegmentum, fleshy, beset dorsally with scattered long, slender, pointed spicules up
to 350 pm long and 25 pm in diameter at the base, occurring singularly or in
groups of two to four; ventral surface of girdle covered with rectangular scales
measuring 20 pm x 7.5 pm; margin of girdle armed with pointed spicules 125-150
um long and 20 pm wide at the base.

Radula about 800 pm long; the number of mature rows of teeth is unknown;
major lateral teeth relatively small, caps tridentate, about 25 um x 20 pm, central
cusp slightly longer than lateral cusps. Unfortunately, the SEM preparation of the

14 NEMOURIA No. 43

Figures 20-25. Acanthochitona burghardtae Clark, spec. nov. 20. Whole animal,
length 2.4 mm, RNC 3137. Isla San Jose. 21. Whole animal, length (curled) ca.
3.0 mm, Paratype, LACM 2880. 22. Intermediate valve five (scale on figure),
Holotype, LACM 2881. 23. Tail valve (scale on figure), Holotype. 24. Close-up of
tegmentum (valve five) showing esthete pores, Holotype. 25. Girdle (scale on
figure), Holotype.

October 30, 2000 Clark 15

radula of this species was unsuccessful, so the details of the radula are largely
unknown.

Type Locality: Gulf of California, Canal de San Lorenzo, about 40 km N of La
Paz, Baja California Sur, Mexico (24°21'N, 111°15'W); 12 m.

Type material: Holotype (LACM 2880) and two paratypes (1, LACM 2881); (1,
RNC 3147).

Additional Material: One specimen (RNC 3137) N side of SW tip of Isla San
Jose, 7 m. (leg. Marco Medina-Lopez, SCUBA, 10 February 1995).

Distribution: Gulf of California, east coast of Baja California Sur, vicinity of
Bahia de La Paz, from the N side of the SW tip of Isla San Jose (approx. 24°52'N,
110°38'W) to the type locality. At depths of 7-12 m.

Discussion: At first this species appears to be a juvenile Dendrochiton lirulatus
Berry, 1963, but close examination of the tegmental sculpture and the insertion
teeth show that it is in fact an acanthochitonid. Due to the shape of the valves
and the peculiar sculpture of the tegmentum A. burghardtae cannot be confused
with any of the known members of Acanthochitona from the eastern Pacific.

Etymology: It is with great pleasure that I name this chiton after Laura and Glenn
Burghardt of Oakdale, California whose book "A Collector’s guide to West Coast
Chitons" inspired me and many others to study these fascinating creatures.

DISCUSSION

The shallow water rhodolith beds provide good habitat for many
invertebrates. Steller (1993) reports that 75 species of invertebrates had been
identified from the Gulf of California rhodolith beds. The chiton fauna is
particularly rich and diverse, with eight species in four genera and three families.
Four of these species appear to be exclusive to the rhodolith habitat. It is likely
that the investigation of rhodolith beds in other areas of the world, as well as the
deeper water beds in the Gulf of California could yield many more species of
chitons.

The molluscan faunas of the central eastern Pacific and Caribbean regions
have many affinities. This is particularly true of many of the tropical elements of
their respective chiton faunas, and at least one species Stenoplax boogii (Haddon,
1886) is present in both faunas. Most of the chitons found in the rhodolith
habitat have affinities in the Florida-Caribbean area, however it is not known
whether any of the Caribbean species occur on rhodoliths. Ischnochiton tomhalei
shows a possible kinship with I. papillosus (C. B. Adams, 1843), and I
thodolithophilus is very similar to I. pseudovirgatus Kaas, 1972 particularly in the
small size, color of the tegmentum, and relatively large, heavy major lateral teeth

16 NEMOURIA No. 43

50 km 28° X i wy Punta
(peor Y Chivato
oO Bahia
ne Concepcion
Q
oN CALIFORNIA
PACIFIC O 8
5°)
OCEAN } Z WOK
>
YP
Ih &

d A
H Fossil Bed Ye Bi \
* Living Wave Bed

4 Living Current Bed
Isla

Coronados

Isla @
Espiritu

Canal
de
e. San Lorenzo

Isla
San
Francisco

Figure 26. Map of collecting sites (after Foster et al., 1997).

of the radula. Callistochiton sp., cf. C. elenensis is similar to C. porosus Neirstrasz,
1905 from Brazil especially in the sculpture of the radial ribs, which are topped
with small pustules. Lepidochitona beanii shows an affinity with L. liozonis (Dall &
Simpson, 1901) and L. rosea Kaas, 1972. Lepidochitona corteziana with its lower
profile, shorter, wider valves and lack of large needle-like spicules on the girdle
appears to be more distantly related to this group. Acanthochitona avicula is part of
a complex of several closely related Panamic and Caribbean species.
Acanthochitona burghardtae is particularly interesting, and is unique among
eastern Pacific acanthochitonids in the shape and form of its valves, and the
distribution and sparse number of its girdle spicules. This species appears to be
similar to Acanthochitona terezae Guerra Junior, 1983 from the Caribbean, both

October 30, 2000 Clark 17

species have similarly shaped and proportioned valves (including their sutural
laminae, which are quite short), illdefined juga covered with pustules and
apparently similar girdle spicules. However I have not seen any specimens of A.
terezae. Acanthochitona minutus Leloup, 1980 from Brazil also shows affinities with
A. burghardtae, particularly in the ill-defined, pustulose juga, however the sutural
laminae of A. minutus are much longer and rounder than either A. burghardtae, or
A. terezae. The arrangement of the girdle spicules appears similar also.

ACKNOWLEDGMENTS

Mr. Barry F. Putman, Cuesta College (California) referred this project to me.
Dr. Michael S. Foster, Moss Landing Marine Labs provided specimens data and
support; Dr. James H. McLean, LACM provided some much needed literature;
Dr. Darlene Southworth, Southern Oregon University made available to me the
SEM in her care (without which this paper would have been impossible); Mr.
Marco Medina-Lopez, Universidad Autonoma de Baja California Sur, La Paz
collected and preserved many of the specimens used in this study. Robert van
Syoc and Elizabeth Kools, CAS provided Mr. Putman's proposed type material
and Dr. Douglas J. Eernisse, California State University at Fullerton critically read
the manuscript. The review of two anonymous reviewers provided many useful
comments.

Collections were made in connection with field work supported by Mexican
Government CONABIO-UABCS Grant # FB135/B086/94 to Dr. Rafael
Riosmena, Universidad Autonoma de Baja California Sur, Inter-American
Institute Grant UCAR $97-74025 to Dr. Michael Foster and National
Geographic Society Grant # 5774-96 to K.H. Meldahl.

AGE AIR sy. Os Wel B)

Bosence, D.W.J. 1983. The occurrence and ecology of recent rhodoliths-a review,
in Peryt, T. M., ed., Coated grains: Berlin, Springer-Verlag, p. 225-242.

Ferreira, A.J. 1979. The genus Callistochiton Dall, 1879 (Mollusca:
Polyplacophora) in the eastern Pacific, with the description of a new species.
The Veliger 21(4): 444-466.

Foster, M.S., R. Riosmena Rodriguez, D.L. Steller, and W.J. Woelkerling. 1997.
Living rhodolith beds in the Gulf of California and their implications for
paleoenviromental interpretation. Geological Society of America, special
paper No. 318: 127-139.

Kaas, P. and R.A. Van Bell. 1985. Monograph of Living Chitons, Vol. II. Dr.
Backhuys Publisher: Rotterdam. 302 pp.

18 NEMOURIA No. 43

Kaas, P. and R.A. Van Belle. 1990. Monograph of Living Chitons, Vol. IV. Dr.
Backhuys Publisher: Rotterdam. 298 pp.

Kaas, P. and R.A. Van Belle. 1994. Monograph of Living Chitons, Vol. V. Dr.
Backhuys Publishing: Rotterdam. 402 pp.

Sirenko, B.I. 1993. Revision of the system of the order Chitonida (Mollusca:
Polyplacophora) on the basis of correlation between the type of gills
arrangement and shape of the chorion processes. Ruthenica 3(2): 93-117.

Sirenko, B.I. 1997. The importance of the development of articulamentum for
taxonomy of chitons (Mollusca, Polyplacophora). Ruthenica 7(1): 1-24.

Skoglund, C. 1997. New Panamic Province distribution records for thirteen
chiton (Polyplacophora) species. The Festivus XXIX(8): 69-74.

Steller, D.L. 1993. Ecological studies of rhodoliths in Bahia Concepcion, Baja
California Sur, Mexico. [MS Thesis]: San Jose, California, San Jose State
University, 89 pp.

Steller, D.L. and M.S. Foster. 1995. Environmental factors influencing
distribution and morphology of rhodoliths in Bahia Concepcion, B.C.S.,
Mexico. Journal of Experimental Marine Biology and Ecology 194 (1995):
201-212.

October 30, 2000

Clark

19

20 NEMOURIA No. 43

Founding Editor Managing Editor
JOHN E. DU PONT TIMOTHY A. PEARCE

Consulting Editors

M.G. HARASEWYCH, National Museum of Natural History, Smithsonian
Institution, Washington, DC.

KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh, PA.

RICHARD W. THORINGTON, JR., National Museum of Natural History,
Smithsonian Institution, Washington, DC.

Published by the
DELAWARE MUSEUM OF NATURAL HISTORY

P.O. Box 3937, Wilmington,
Delaware 19807-0937, U.S.A.

vy
Supe

iy ¥ Sanh

aN
iy

5
lives

ware

Me sot ie 1)

paren ee By SUN ny Aen ie y ‘ .
Museum of Natural History

Ai

Ria ats ie Me eae a: aes a

\ : 1) ire
Hh

THE DEVELOPMENT OFANISLAND

i
eh Thee

‘TERRESTRIAL MOLLUSKS OFBERMUDA

%

f

Rudiger Bieler

“ <." Jolin Slapcinsky

ke

‘DECEMBER 30, 2000

¥ t ree

ei ta : Me 1 3 iy tp \

eel]

E MUSEUM OF NATURAL HISTORY |

5

y

y,

a i ee eae aA ‘ ‘ f j oat ase
rt { Tt t 1 planta t yy > aif

3 yy 7 + ; ¥ )

Powe gos7, Wikatmstony

> 2" y
; ; 490
;

tive

a ale 4:
(tae

; Vk : USA

4 i f ; \ r

Delaware 198070

NEMOURIA

Occasional Papers of the Delaware Museum of Natural History

NUMBER 44 DECEMBER 30, 2000

A CASE STUDY FOR THE DEVELOPMENT OF AN ISLAND
FAUNA: RECENT TERRESTRIAL MOLLUSKS OF BERMUDA

Riidiger Bieler’ and John Slapcinsky””

ABSTRACT. The isolated western Atlantic island group of Bermuda provides a
model case for the development of an island snail fauna. A unique combination
of factors, such as an excellent Pleistocene fossil record, late (and precisely dated)
human colonization, and extensive historic collecting and documentation efforts
(last published in 1926), provide the background for this study. New field
collecting, a survey of museum collections, and a critical review of published and
unpublished information (resulting in several synonymies and reinterpretations of
previously assumed endemics), were used to determine the presence and range of
all terrestrial mollusks on Bermuda. At least 49 species of snails and slugs from 22
families are now known from Bermuda, including 11 endemic, 6 native non-
endemic, 28 accidentally introduced and 4 intentionally introduced species.
Starting in 1958, several exotic predatory snails including Gonaxis
quadrilateralis and Euglandina rosea were released in illadvised attempts at
biologically controlling the accidentally introduced snail species Otala lactea and
Rumina decollata. Euglandina rosea is implicated in the extinction of several land
snails native to the Pacific islands where it was also introduced. In Bermuda,
densities of the three extant species in the endemic land snail genus Poecilozonites
have been reported declining since the early 1960s and the group now appears to
be nearly extinct. Individuals (or fresh shells) of two Poecilozonites species (P.
circumfirmatus and P. bermudensis) were found although neither is common. This
faunal survey finds E. rosea spread from its original release sites and generally
distributed on the larger islands of Bermuda, with the possible exception of the

' Department of Zoology, Field Museum of Natural History, Serial Publication
1400 S Lake Shore Drive, Chicago, Illinois 60605-2496, ISSN 0085-3887
US.A.; bieler@fmnh.org

? Current address: Florida Museum of Natural History, University of Florida, Gainesville,
FL 32611, U.S.A., slapcin@flmnh.ufl.edu

2 NEMOURIA No. 44

southeastern corner of the main island. Gonaxis is beginning to spread beyond its
release area. Several endemic and native non-endemic species including P.
reinianus, Vertigo marki, V. bermudensis, and Strobilops hubbardi have only been
found in the southeastern portion of the main island near caves and remnant
native vegetation. The present study records three species new to Bermuda,
Tornatellides sp., Streptostele musaecola, and Opeas hannense. Zonitoides bermudensis,
previously recorded as an endemic species, is here synonymized with Z. arboreus
and is recognized as a species introduced through human commerce. Gastrocopta
barbadensis crassilabris is synonymized with G. pellucida, and recognized as a native
non-endemic species. Strobilops pilsbryi, described as a species endemic to Bermuda,
is synonymized with S. hubbardi. Pupisoma macneilli, Punctum bristoli, and Striatura
meridionalis, previously known from Pleistocene fossils, are here recorded as
members of Bermuda’s extant fauna for the first time.
Key words: Bermuda, endemic, exotic, fauna development, land snails.

INTRODUCTION

Bermuda, a crescentshape group of several large and numerous smaller
western Atlantic islands, lies over 900 kilometers east of the nearest landfall at
Cape Hatteras, North Carolina, and is among the most isolated island groups in
the world. This remoteness is reflected in its depauperate native fauna (Wallace,
1895), especially by the near absence of non-avian vertebrates, represented by only
one native species, the Bermuda Rock Skink, Eumeces longirostris (Cope, 1861).

Land snails are a major component of the native fauna and comprise nearly
all of the Pleistocene fossils from terrestrial deposits (Gould, 1971b). Pulmonate
snails are arguably among the most common and diverse animals on Bermuda.
Previous authors (Pilsbry, 1900; Verrill, 1902) estimated the number of native
species at 15 or 13 respectively (but, as will be shown below, included several
species now known to be introduced through human commerce). Bermuda’s land
snail fauna is best known for its endemic zonitid genus, Poecilozonites. This group
of several nominal extinct and extant species became the focus of numerous
taxonomic, morphometric, and evolutionary studies (e.g., Peile, 1924, 1925;
Kutchka, 1934; Pilsbry, 1934; Gould, 1966, 1968, 1970a,b, 1971a, b). The
postulated rapid adaptive radiation of the P. bermudensis complex was cited as one
of the two evidential examples for the Punctuated Equilibrium hypothesis
(Eldredge & Gould, 1972), although the hypothesized evolutionary scenario for
this “Darwin’s finch among mollusks” (Gould, 1991: 12), based on ice-age
climatic oscillations, has not remained undisputed (Brown, 1987; Glaubrecht,
1990).

The rich history of faunal surveys before 1926 has provided a remarkably
complete picture of Bermuda's terrestrial snail fauna to that date. The lack of

December 30, 2000 Bieler & Slapcinsky 3

recently published surveys, the rapid decline of Bermuda's cedar forests, and the
deliberate introductions of four species of predatory snails as agents of biological
control make the current survey especially timely.

Bermuda was one of the few landmasses uninhabited by humans in Pre
Columbian times and was not settled until 1612. Late colonization and good
written records have made the human-mediated changes to Bermuda's flora and
fauna among the best documented of any region. Human intervention in
Bermuda’s ecosystem began long before settlement, in 1515, with the release of
pigs by Bermudez who had discovered the islands in 1503. The first settlers, after
1612, added cats and dogs. Between 1614 and 1618, large populations of
introduced rats, Rattus rattus (Linné, 1758), led to crop failures and the starvation
of many colonists (Lefroy, 1876, cited from Verrill, 1902). Deliberate burns to
eradicate the rats eliminated most of the already heavily logged primary cedar
forest. Early last century, only a small tract of unburned native forest remained at
Walsingham Jungle (Verrill, 1902). Several land snails appear to have been
limited to this; however with only one previously published distributional study
(Vanatta, 1911) among those merely recording “Bermuda,” actual snail
distributions in the island group were poorly known. This survey is the first since
the remaining stands of the Bermuda cedar (Juniperus bermudensis) were severely
damaged by scale blight, shortly after World War II.

Slightly east of the Gulf Stream (at 32°18’N, 64°46’W), Bermuda's climate is
frost-free and mild, with about 1400 mm of evenly distributed rainfall annually.
This mild climate has allowed the introduction and naturalization of many
agricultural and ornamental plants, and with these, of numerous foreign plants
and animals, including terrestrial mollusks. It is thus no surprise that many of
today’s non-native land snails in Bermuda are species commonly found associated
with greenhouses and nurseries. Land snail introductions on Bermuda are
particularly well documented with more than a dozen qualitative faunal surveys
published between 1852 and 1926, leading Tomlin to assert "There is probably no
island in the world that has been more thoroughly combed for non-marine shells
than Bermuda" (1932: 98). Already by the time of the first faunal survey of
Bermuda, at least six introduced species were present (Prime, 1853). By 1926, the
date of the last published survey, at least 14 species of native and 22 species of
introduced land snails inhabited Bermuda (Peile, 1926). During that 73-year
period, little if any turnover in species had occurred. Each survey found nearly all
but the smallest (< 3 mm) of the species previously recorded for the islands,
despite the fact that many of these surveys were short and secondary to other
projects.

Pilsbry (1900) saw three origins for Bermuda’s land snail fauna: (a)
“Autocthonous species, peculiar to the islands,” (b) “Drift waifs from the West
Indies,” and (c) “Species imported by the agency of man.” We have categorized

4 NEMOURIA No. 44

our records (and our interpretations of earlier records) in four different groups:
(1) endemic species (species not known from elsewhere, and also present in
Bermuda’s good Pleistocene fossil record), (2) native, but not endemic, species
(species that were present before the arrival of the first settlers), (3) accidentally
introduced species, and (4) deliberately released species. Some of these might no
longer have viable populations on the islands, as will be discussed below.

Several additional introductions have been recorded since the most recent
faunal survey. Rumina decollata - an omnivorous snail now widely distributed in
tropical and subtropical areas through human commerce - was originally native to
the Mediterranean region and is considered a crop pest. This snail, introduced
accidentally with plants in the late 1870s has become a widespread agricultural
pest in Bermuda (Bennett & Hughes, 1959). In 1928, a second important
horticultural pest was introduced. The Milk Snail Otala lactea, imported in 1928
for food, escaped and became naturalized (Bennett @ Hughes, 1959). By the
1950s, the Bermuda Department of Agriculture was looking into methods to
control these crop pests. With help from the Hawaiian Department of Agriculture
several species of predatory snails were introduced in illfated attempts at
controlling Otala and Rumina (Simmonds & Hughes, 1963). One of these
predatory species, Euglandina rosea, is implicated in the extinction and
endangerment of many land snails endemic to the Pacific Islands (Hadfield &
Mountain, 1981; Clarke et al., 1984; Murray et al., 1989) and can also impact
upon freshwater snails (Kinzie, 1992). The same fate might have befallen
Bermuda's endemic genus Poecilozonites. Numbers of Poecilozonites bermudensis have
been declining since the early 1960s (Gould, 1969, 1991).

The unique situation of Bermuda’s historically well-documented land snail
fauna provides an opportunity to track each species through recorded time - in
several cases even establishing the exact event of first introduction. A complete
bibliography for Bermuda land mollusks is also provided. This paper is part of an
informal series on the terrestrial flora and fauna of Bermuda initiated by the
Bermuda Department of Agriculture and Fisheries. Previous studies have
included spiders (Sierwald, 1988), Coleoptera (Hilburn & Gordon, 1989),
Lepidoptera (Ferguson et al., 1991), and Diptera (Woodley & Hilburn, 1994).

MATERIALS AND METHODS

The core of the material for the present survey stems from intensive
collecting efforts during one week in May of 1988. That survey involved three
collectors (RB, Dr. Daniel Hilburn, Dr. Petra Sierwald), and most of the
collecting sites are identical to those reported by Sierwald (1988) in a survey of
Bermuda’s spider fauna. Additional data originate from preliminary collections
made by RB during a visit to the Bermuda Biological Station in 1983, from the

December 30, 2000 Bieler & Slapcinsky 5

Map 1. Collecting localities.

collections previously held at the Department of Agriculture (now at the Natural
History Museum, Bermuda), the Natural History Museum, Bermuda, and the
Neil Fahy (Daly City, California) private collection.

Unique or questionable literature records were verified based on available
voucher material, as indicated. In addition, Bermuda land snail holdings of the
Academy of Natural Sciences of Philadelphia, the American Museum of Natural
History, the California Academy of Sciences, the Delaware Museum of Natural
History, the Field Museum of Natural History, the Florida Museum of Natural
History, and the University of Michigan Museum of Zoology were studied. The
existing literature on Bermuda land snails and all original species descriptions
were researched, leading to new synonymies and re-interpretations of earlier
records.

Maps: The distributions of all Recent species found on Bermuda are plotted using
locality data from this survey and data from non-fossil museum specimens (see
map | and museum collections studied).

This survey:
@ Closed squares: Live specimens.
f™ Partly filled squares: Recently dead specimens (with periostracum).

& X-filled squares: Long dead specimens.
Other records:

@ Closed circles: Museum specimens (nearly all collected prior to 1926),
and in the case of the four deliberately introduced
species (Euglandina rosea, Gonaxis kibweziensis, G. quadri-
lateralis and Natalina cafra) from surveys and annual
reports of Bermuda’s Department of Agriculture.

R Release sites of deliberately introduced predatory snails.

6 NEMOURIA No. 44

Specimens collected during the 1983/1988 survey (map 1) are deposited at
Field Museum of Natural History (FMNH 224755-224920 and 227001-227056),

with duplicate voucher series deposited at BAMZ and DMNH.

1988 Survey sites:
O1 Sandy's Parish
02 Sandy's Parish
03 Sandy’s Parish
04 Southampton Parish
05 Southampton Parish
06 Warwick Parish
O7 Paget Parish
O8 Paget Parish
09 Devonshire Parish
10 Smith's Parish
11 Hamilton Parish
12 Hamilton Parish
13 Hamilton Parish
14 St. George’s Parish
15 St. George's Parish
16 St. George's Parish

Museum collections cited:

Gilbert Nature Reserve
Fort Scaur

Hog Bay Level
Seymour’s Pond
Horseshoe Bay
Warwick Pond

Que Sera Guesthouse
Botanical Garden
Devonshire Marsh
Spittal Pond
Bermuda Aquarium
Shelly Bay
Walsingham Pond
Whalebone Bay
Mullet Bay Park
Smith's Island

AMNH American Museum of Natural History, New York

ANSP Academy of Natural Sciences, Philadelphia

BAMZ Bermuda Aquarium, Natural History Museum and Zoo
BPBM Bernice P. Bishop Museum, Honolulu

CAS California Academy of Sciences, San Francisco

DMNH Delaware Museum of Natural History, Wilmington
FMNH Field Museum of Natural History, Chicago

MCZ Museum of Comparative Zoology, Cambridge

UF Florida Museum of Natural History, Gainesville
UMMZ University of Michigan Museum of Zoology, Ann Arbor

LIST OF RECENT BERMUDIAN TERRESTRIAL MOLLUSKS

The taxonomic arrangement in superfamilies and families follows Smith &
Stanisic (1998) and Turgeon et al. (1998), respectively. In addition,
Agriolimacidae Wagner, 1935 (see Wiktor, 2000), and Cochlicellidae Schileyko,
1972 (see Schileyko & Menkhorst, 1997) are used. Within superfamilies, families,

December 30, 2000 Bieler & Slapcinsky 7

genera, and species are each arranged alphabetically. The listings of synonyms and
subsequent citations are restricted to records with relevance to Bermuda and
those impacting taxonomic decisions. Species known from Bermuda only as
fossils were not included. All species listed below were found in the current survey
unless otherwise indicated.

Species were determined to be either (1) endemic to Bermuda, (2) native, but
not endemic, (3) accidentally introduced by human activities, or (4) intentionally
introduced for pest control. All native species are found in Bermuda's Pleistocene
deposits which are among the best sampled and studied of their kind. Species
missing from the fossil deposits are considered introduced. This assumption is
reasonable in this case given that (1) all endemic species are found as fossils, (2)
even uncommon or patchily distributed native non-endemic species are present in
fossil deposits, and (3) most species without a fossil record in Bermuda are well
known adventive species.

Distributions for all species on Bermuda were plotted using (1) new records
from the present survey, (2) published surveys, especially that of Vanatta (1911),
(3) museum and private collection records, including most voucher specimens of
previous surveys, and (4) unpublished records of the Bermuda Department of
Agriculture & Fisheries. The notes for each species include information about
species ranges, habitat preferences, and method of introduction.

NERITIMORPHA: NERITOIDEA
HELICINIDAE
Oligyra convexa (Pfeiffer, 1849)
(Map 2, Fig. 1)

Helicina convexa Pfeiffer, 1849: 120-121 (“Locality unknown”).

Helicina ---, - Redfield, 1853: 17;

Helicina variabilis, - Prime, 1853: 55; Jones, 1859: 107; Tristram, 1861: 405
[non Helicina variabilis Wagner in Spix, 1827: 25, pl. 16, figs. 3-4, Brazil];

Helicina subdepressa Poey, 1851: 420; - Jones, 1864: 9 [non Helicina subdepressa
Poey, 1851];

Helicina convexa, - Bland, 1861: 351; Rein, 1867: 77; Morch, 1878: 3;
Bartram, 1878: 108; Kobelt, 1880: 286; Bland, 1881: 257; E. A. Smith,
1884: 277; Heilprin, 1889: 184; Bland, 1895: 271; Gulick, 1904: 421;
Pilsbry, 1900: 506, pl. 62, figs. 15, 15a; Verrill, 1902: 728, fig. 69;
Vanatta, 1911: 665-668; Honigmann, 1914: 377; Olmsted, 1917: 225;
Vanatta, 1923: 33; Peile, 1926: 76 [“the only true land operculate in the
islands”]; Hughes, 1968: 1;

8 NEMOURIA No. 44

Figure 1. Oligyra convexa. - FANH 29795, St. George’s Parish, Bermuda; diameter

= 7.7 mm (apical view); diameter = 7.4 mm (basal and apertural views).

Map 2. Oligyra convexa.

Helecina [sic] convexa, - Jones, 1876: 139;

Helicinia varibilis [sic], - Bartram, 1878: 108;

Helicina (Pachystoma) fasciata, - Angas, 1884: 597 [placed H. convexa in
synonymy of Helicina fasciata Lamarck, 1822: 102 (type locality “Puerto
Rico”; ? Guadeloupe, see van der Schalie, 1948: 21)];

Helix Convexus, - Lane, 1891: 1, pl. 2, fig. 9 [plate not seen];

Oligyra convexa, - Baker, 1922: 44 [based on material collected in Bermuda by
Gulick in 1903};

Helicina fasciata convexa, - Wagner, 1911: 335, pl. 67, figs. 8, 9.

December 30, 2000 Bieler & Slapcinsky 9

Helicina (Helicina) fasciata convexa, — Haas, 1952: 104.

Bermuda Material Studied: BAMZ 1994-118-006, 1995-132-023, 1998-166021,
1998-166-034, 1998-166-049, 1998-166-066, 1998-166-068, 1998-166-080, 1998-
166-104; DMNH 210240, 210265, 210270, 210328, 210330, 210331, 210333;
FMNH 3624, 3625, 5473, 8835, 13485, 13499, 13510, 13521, 13528, 13541,
13548, 13552, 13560, 13563, 13567, 13571, 13574, 13576, 13594, 13683,
16465, 28193, 28615, 28769, 29795, 29796, 41737, 59081, 92869, 98243,
98251, 98286, 98317, 103289, 115079, 126379, 130912, 147968, 216085,
224764, 224778, 224795, 224796, 224820, 224833, 224880, 224897, 224911,
227019, 227025.

Note: Accidentally Introduced - One of the earliest introductions, this species was
recorded for the year 1852 by Prime (1853) and by nearly every survey since. A
very common species, it is found under rocks and on vegetation often in gardens
and plantings. Hughes' (post 1969: 1) reference to fossil material is erroneous; the
complete lack of helicinids in Bermuda's thoroughly investigated Pleistocene
deposits suggests a more recent arrival. Bermuda's small high-spired helicinids are
similar to forms found on the islands of the Lesser Antilles. Bermuda's early
governors promoted the importation of exotic vegetation from other British
colonies to provide food sources for early colonists (Verrill, 1902) and it is likely
that this species was imported with these agricultural products. Revision of the
West Indian helicinids is necessary before the source of O. convexa can be
determined. The species was transferred to Oligyra by Baker (1922) based on
characters of the radula and operculum.

BASOMMATOPHORA: ELLOBIOIDEA
CARYCHIIDAE
Carychium bermudense Gulick, 1904
(Map 3, Fig. 2)

Carychium bermudense Gulick, 1904: 415, pl. 36, figs. 11, 12 (Bermuda; fossil).

Carychium exiguum (Say, 1822) - Davis, 1904: 126 [non Pupa exigua Say, 1822:
375, which was cited by Pfeiffer, 1841b: 224. as Carychium exiguum
“naib

Carychium bermudense, - Verrill, 1906: 171, figs. 54a, b (type specimen; fossil);
Vanatta, 1924: 7; Gould, 1970b: 490, fig. 1h (fossil); Gould, 1970a: 572
(fossil);

Carychium bermudensis [sic], - Vanatta, 1911: 666-668; Vanatta, 1923: 33;
Peile, 1926: 88.

10 NEMOURIA No. 44

Figure 2. Carychium bermu Figure 3. Tornatellides sp. - FMNH
dense. - BAMZ 1995-133-003, 227039, Sandy’s Parish, Bermuda;
Bermuda; height = 2.0 mm. height = 2.9 mm.

Bermuda Material Studied: ANSP 88559; BAMZ 1995-133-003; FMNH 30340,
227028, 227029; MCZ Gould Station 1 (fossil), Gould Station 11 (fossil), Gould
Station 21 (fossil), Gould Station 21b (fossil), Gould Station 30 (fossil), Gould
Station 41 east (fossil), Gould Station 48 (fossil).

Note: Endemic - This species was described from fossil material collected during
expeditions sponsored by the then new Bermuda Biological Station (Gulick,
1904). Recent specimens were later collected by Stewerdson Brown (Vanatta,
1911). Specimens can be sifted from leaf litter, especially in the vicinity of
marshland. The heavily callused and flared lip, tapered whorls, fine spiral striae,
as well as the strongly dilated internal lamella which emerges strongly on the
parietal wall and as a low basal thickening on the columellar wall, are similar to

Map 3. Carychium bermudense.

December 30, 2000 Bieler & Slapcinsky 11

Carychium mexicanum of the coastal region of southeastern North America; the
relationship between the two nominal taxa needs further study.

STYLOMMATOPHORA: ACHATINELLOIDEA
ACHATINELLIDAE
Tornatellides sp. NEW RECORD
(Map 4, Fig. 3)

Bermuda Material Studied: BAMZ 1998-166-120, 1998-166-091, 1998-166-089,
1998-166-067, 1998-166-015; DMNH 210255, 210259, 210275, 210310; FMNH
224809, 224810, 227013, 227014, 227015, 227016, 227017, 227030, 227039.

Note: Accidentally Introduced - The genus Tornatellides is native to the Pacific Basin.
This species (here treated as Tornatellides “sp.” because of the current unresolved
taxonomic situation of the genus) probably arrived in Bermuda sometime after
the last faunal survey (Peile, 1926). It is now widespread on Bermuda and can be
collected in great numbers from the bark of trees, suggesting possible importation
through horticulture. This is the first record of this family in the Atlantic Basin.

Map 4. Tornatellides sp.

PUPILLOIDEA
PUPILLIDAE
Gastrocopta pellucida (Pfeiffer, 1841)
(Map 5, Fig. 4)

Pupa pellucida Pfeiffer, 1841a: 46 (Cuba).

? Pupa bermudensis Prime, 1853: 55 [nomen nudum];

12

NEMOURIA

Pupa jamaicensis C. B. Adams, 1849, -
Bland, 1861: table 2, p. 351; Jones,
1876: 139; Kobelt, 1880: 286;
Bland, 1881: 257; Bland, 1895:
271; Heilprin, 1889: 182 [non Pupa
jamaicensis C. B. Adams, 1849];

? Pupa Bermudensis Jones, 1859: 107
[nomen nudum; based on Prime’s list,
but indicated as “n.s.”];

Pupa pellucida, - Jones, 1876: 139;
Kobelt, 1880: 286;

Pupa barbadensis, - Bland, 1881: 257;
Bland, 1895: 271; Heilprin, 1889:
183 [non Pupa barbadensis Pfeiffer,
1853: 69 (Barbados)];

Pupa (Bifidaria) jamaicensis, - Pilsbry,
1900: 498 [non pl. 62, fig. 7 = G.
rupicola]; Verrill, 1902: 729 [non fig.
74b);

Bifidaria jamaicensis, — Gulick, 1904: 415;

Bifidaria pellucida hordeacella (Pilsbry,
1890), - Vanatta, 1911: 665-668;

No. 44

Figure 4, Gastrocopta
pellucida. - FMNH 224832,
Hamilton Parish, Bermuda;
height = 2.6 mm.

Gastrocopta barbadensis crassilabris Pilsbry, 1916: 85-86, pl. 18, figs. 69
(Bahamas and Bermuda), new synonymy;

Gastrocopta barbadensis crassilabris, — Peile, 1926: 88;

Gastrocopta barbadensis (Pfeiffer, 1853), - Vanatta, 1923: 33;

Gastrocopta pellucida hordeacella, - Vanatta, 1923: 33; Peile, 1926: 88; Gould,

1970b: 490, fig. 1b (fossil);

Gastrocopta (Gastrocopta) pellucida hordeacella, - Haas, 1952: 104.

Map 5. Gastrocopta pellucida.

December 30, 2000 Bieler G Slapcinsky i

Bermuda Material Studied: ANSP 3816, 143338; BAMZ 1995-133-008; FMNH
29802, 30201, 30203, 41009, 41010, 41011, 48647, 59074, 224832, 227040,
227048; MCZ Gould Station 4 (fossil), Gould Station 41 east (fossil), Gould
Station 63 lower (fossil), Gould Station 73 (fossil); UMMZ 106756.

Note: Native Non-Endemic - This species ranges over much of the West Indies and
North America from Florida north along the coast to New Jersey, west to
southern California and southern Mexico. This and the following species are the
most common Gastrocopta species on Bermuda and can be found on rocks in
open areas. “Pupa bermudensis” probably refers to Gastrocopta pellucida, as the only
material originally labeled “Pupa bermudensis” found in this study is a lot of G.
pellucida, collected by A. D. Brown (ANSP 3816). Gastrocopta pellucida is extremely
variable, not only in overall shell shape and size but also in the thickness of the
peristome and apertural barriers, as was pointed out by Pilsbry (1916: 79). Several
names have been applied to Bermuda's Gastrocopta pellucida depending on the
thickness of the lamellae and peristome and the length of apertural plicae. These
names include Gastrocopta pellucida hordeacella (for larger forms similar to those of
the North American mainland with longer palatal plicae and a slight crest behind
the lip), as well as forms misidentified as G. barbadensis (with thicker lamellae
including a strongly bifid angulo-parietal barrier). Those specimens also having a
thickened peristome have been described under the name G._ barbadensis
crassilabris. Pilsbry (1916: 85) stated: “In shape and size, crassilabris resembles
pellucida, but the thick lip and the teeth are more like marginalba, the peristome
being thicker and the crest stronger than in barbadensis.” However, large series
both from the North American mainland (FMNH 232835) and Bermuda (ANSP
143338) contain individuals that range from those similar to the holotype and
paratypes of pellucida hordeacella (ANSP 60460) to individuals with stronger
barriers and well developed bifid angulo-parietal teeth and thick peristomes that
could be identified as G. barbadensis crassilabris. This variation is well illustrated by
Vanatta (1912: plate 2; later reprinted by Pilsbry, 1948: fig. 495). Differences
noted by Pilsbry between these nominal taxa, including the strength of the
lamellae and plicae, the degree of bifurcation of the angulo-parietal lamellae, and
the color differences between thin translucent yellowish shells and thicker brown
shells can all potentially be caused by differences in calcium availability or by the
age of individuals within a population. This view is supported by evidence of far
greater changes in shell form due to differences in calcium availability within
populations of other Bermuda species (Gould, 1970b, 1971a). The nominal
subspecies Gastrocopta barbadensis crassilabris Pilsbry, 1916, is here synonymized
under G. pellucida.

NEMOURIA

No. 44

Gastrocopta rupicola (Say, 1821)

(Map 6, Fig. 5)

Pupa rupicola Say, 1821: 163 (East Florida).

Pupa pellucida Pfeiffer, 1841 (in part; see also G. servilis, below), - Bland,
1861: 351; Rein, 1867: 77; Bland, 1881: 257; Bland, 1895: 271; Jones,
1876: 139; Heilprin, 1889: 183 [non Pupa pellucida Pfeiffer, 1841];

Gastrocopta (Gastrocopta) barbadensis, - Haas, 1952: 104 [non Pupa barbadensis

Pfeiffer, 1853];

Pupa (Bifidaria) rupicola, - Pilsbry, 1900:
498, pl. 62, figs.7 [sic], 8; Verrill, 1902:
729, figs. 4b [sic], 74c; Verrill, 1906:
170, fig. 68c;

Bifidaria rupicola, - Gulick, 1904: 414;
Vanatta, 1911: 665-668; Vanatta,
1912: 12;

Gastrocopta rupicola duplex Pilsbry, 1916: 60,
pl. 11, figs. 4, 7, 9 (Bermuda);

Pupa marginalba Pfeiffer, 1840: 253 (Cuba);

Gastrocopta rupicola marginalba (Pfeiffer,
1840), - Pilsbry, 1916: 61, pl. 11, figs.
10,11; Vanatta, 1924: 7; Peile, 1926:
88 (fossil);

Gastrocopta rupicola, - Vanatta, 1923: 33;
Peile, 1926: 88; Gould, 1970a: 572
(fossil); Gould, 1970b: 490, fig. 1d
(fossil);

Pupa rupicola, - Hughes, post 1969: 1;

Gastrocopta (Gastrocopta) rupicola, - Pilsbry,
1948: 905, 906.

Figure By Gastrocopta
rupicola. - FMNH 224805,
Smith’s Parish, Bermuda;
height = 2.5 mm.

Map 6. Gastrocopta rupicola.

December 30, 2000 Bieler & Slapcinsky 15

Bermuda Material Studied: BAMZ 1995-
133-007; FMNH 30200, 41012, 59075,
224805, 224806, 227049; MCZ Gould
Station 1 (fossil), Gould Station 4 (fossil),
Gould Station 11 (fossil), Gould Station 21
(fossil), Gould Station 21b (fossil), Gould
Station 30 (fossil), Gould Station 41 east
(fossil), Gould Station 48 (fossil), Gould
Station 63 lower (fossil), Gould Station 73
(fossil).

Note: Native Non-Endemic - This species is
usually found in open, dry areas under rocks,
often together with Gastrocopta pellucida and
Pupoides albilabris. Gastrocopta rupicola ranges

along the extreme southeastern coast of
North America from North Carolina to Figure 6. Gastrocopta servilis. -
Texas and the West Indies. Like the previous FMNH 30302, Devonshire
species it is widespread on Bermuda. Old Parish, Bermuda; height = 2.6
individuals in calcareous areas often develop mm (lip chipped).

very thick lips, lamellae and plicae.

Gastrocopta servilis (A. A. Gould, 1843)
(Map 7, Fig. 6)
Pupa servilis Gould, 1843: 356, pl. 16, fig. 14 (Mexico).
Pupa pellucida Pfeiffer, 1841 (in part; see also G. rupicola, above), - Bland,

1861: 351; Bland, 1881: 257; Bland, 1895: 271; Heilprin, 1889: 183
[non Pupa pellucida Pfeiffer, 1841];

Map 7. Gastrocopta servilis.

16 NEMOURIA No. 44

Pupa (Bifidaria) servilis, -
Pilsbry, 1900: 497, pl.
62, fig. °6;" -Verzill,
1906: 169-170, | fig.
68a; Verrill, 1902:
729, fig. 74a;

Bifidaria servilis, - Gulick,
1904: 414; Vanatta,
1911: 665-668;

Gastrocopta servilis, - Peile,

1976.88.

Bermuda Material Studied:
FMNH 30202.

Note: Accidentally Introduced - Figure 7. Pupisoma macneilli. - FMNH

Gulick (1904: 414) reported a 227043, Hamilton Parish, Bermuda; height

single fossil specimen from = _ 1,08 min (aperture dammaeeen
Bermuda. However, no

additional records were recorded

by Gould (1970b) or located in his samples from stations 4, 41, and 48. Gulick’s
record is likely due to contamination with Recent material as this common
Neotropical species has been widely introduced through commerce (Hubricht, 1985).

Pupisoma macneilli (Clapp, 1918) NEW EXTANT RECORD
(Map 8, Fig. 7)
Thysanophora macneilli Clapp, 1918: 74, pl. 8, fig.1 (Alabama).
Pupisoma sp., —- Gould, 1970b: 490 (fossil).

Bermuda Material Studied: FMNH 227043; MCZ Gould Station 4 (fossil),

Map 8. Pupisoma macneilli.

December 30, 2000 Bieler & Slapcinsky 17

Gould Station 11 (fossil), Gould Station 41
east (fossil), Gould Station 48 (fossil), Gould
Station 73 (fossil). This present study records
the first Recent specimens of this species for
Bermuda (fresh specimens with
periostracum, collected in surface sand
samples from Walsingham Pond; FMNH
227043). Also studied was the Pupisoma
material as cited by Gould (1970b: 490).

Note: Native Non-Endemic - This widespread
Neotropical species ranges from the
southeastern United States to Central and
South America and the West Indies.
Pupisoma macneilli can be found on smooth-
barked tree trunks and _ under _ leaves
(Hubricht, 1985: 12). Pupisoma was first
recorded from Bermuda by Gould (1970b)
from Pleistocene deposits. Material examined
from Gould stations 4, 41, and 48 (S. J.
Gould research collection, specimens not
individually numbered or sorted) are here
identified as P. macneilli.

Figure 8. Pupoides albilabris. -
BAMZ 1995-132-004, Ber-
muda; height = 4.6 mm.

Pupoides albilabris (C. B. Adams, 1841)

(Map 9, Fig. 8)

Pupa albilabris C. B. Adams, 1841 [new name for Cyclostoma marginata Say,
1821: 172-173, non Cyclostoma marginatum G. Fischer, 1807: 219] (Cuba).

Map 9. Pupoides albilabris.

18 NEMOURIA No. 44

Bulimus nitidulus Pfeiffer, 1839: 352 (Cuba) [not to be confused with
Bulimulus nitidulus Beck, 1837: 67];

? Bulimus Sandysii Prime, 1853: 55 [nomen nudum];

? Bulimus Sandysii Jones, 1859: 107 [nomen nudum; based on Prime’s list, but
indicated as “n.s.”];

Bulimus nitidulus, - Bland, 1861: table 2, p. 351; Rein, 1867: 77; Bland, 1881:
256; Bland, 1895: 271;

Pupa fallax Say, 1825, - Jones, 1876: 139; Bartram, 1878: 108;

Bulimulus fallax Say (nitidulus Pfr.), - Kobelt, 1880: 286;

Pupoides fallax, - Heilprin, 1889: 183;

Pupoides marginatus (Say, 1821), - Pilsbry, 1900: 498, pl. 62, fig. 16; Verrill,
1902: 729, fig. 73; Gulick, 1904: 415; Vanatta, 1911: 666-668; Peile,
1926: 89 [non Pupa marginata Draparnaud, 1801: 58; nec Cyclostoma
marginata G. Fischer, 1807; see Pilsbry, 1948: 923];

Pupoides (Pupoides) albilabris, - Pilsbry, 1948: 921,

Pupoides albilabris, - Gould, 1970b: 490, fig. 1c (fossil);

Pupoides nitidulus, - Abbott, 1989: 64, fig.

Bermuda Material Studied: BAMZ 1995-132-004; Gould Station 63 lower
(fossil), Gould Station 73 (fossil).

Note: Native Non-Endemic - Pupoides albilabris ranges over much of eastern and
southern North America from southeastern Canada to Mexico (Pilsbry, 1948) and
has been introduced in California (Hanna, 1966). It is a calciphile often found in
glades and other open areas (Hubricht, 1985). This species appears to be patchily
distributed on Bermuda (Jones, 1876, recorded it as rare) and was not found in
this survey.

Vertigo bermudensis Pilsbry, 1919
(Map 10, Fig. 9)
Vertigo bermudensis Pilsbry, 1919: 149, pl. 13, figs. 8-10, 12 (Bermuda).
Vertigo eyriesii (Drouét, 1859), - Vanatta, 1911: 666 [non Pupa eyriesii Drouét,
1859];
Vertigo bermudensis, - Peile, 1926: 89.

Bermuda Material Studied: ANSP 91159, 105610; BAMZ 1995-132-001.

December 30, 2000 Bieler @ Slapcinsky 19

Note: Endemic - To date, the only
specimens of this species with verifiable
data were collected in the early 1900s
from the vicinity of Church Cave near
Tuckerstown. Pilsbry (1919: 149, 150)
compared this species to the similar
Vertigo milium (Gould, 1840) of eastern
North America.

Vertigo marki Gulick, 1904
(Map 11, Fig. 10)

Vertigo marki Gulick, 1904: 414, pl.
36, fig. 7 (Bermuda, fossil).

Vertigo marki, - Verrill, 1906: 170, Figure 9. Vertigo bermudensis. -
fig. 54d (type; fossil); Vanatta, BAMZ 1995-132-001, Bermuda;
1911: 666; Pilsbry, 1919: 107, height = 1.3 mm.
pl. 13, fig. 17; Peile, 1926: 89
(fossil); Gould, 1970b: 490
(fossil).

Bermuda Material Studied: ANSP 85574, 91156; BAMZ 1995-132-002; MCZ
Gould Station 4 (fossil).

Note: Endemic - This species has never been documented alive, although empty
Recent shells are recorded from leaf litter around Church Cave (Pilsbry, 1919).
This species is believed to be most closely related to Vertigo tridentata Wolf, 1870,
of eastern North America (Pilsbry, 1919).

Map 10. Vertigo bermudensis.

20 NEMOURIA No. 44

Figure 10. Vertigo marki. - Figure 11. Vertigo numellata. -
BAMZ 1995-132-002, Bermuda; ANSP 105611, Hamilton Parish,
height = 2.2 mm. Bermuda; height = 1.7 mm.

Vertigo numellata Gulick, 1904
(Map 12, Fig. 11)

Vertigo numellata Gulick, 1904: 413-414, pl. 36, fig. 6 (Bermuda, fossil).

Vertigo numellata, - Verrill, 1906: 170, fig. 54c (type); Vanatta, 1911: 666;
Pilsbry, 1919: 91, pl.13, figs. 14, 15; Vanatta, 1924: 7; Peile, 1926: 88;
Hughes, post 1969: 1; Gould, 1970b: 490, fig. Le.

Bermuda Material Studied: ANSP 85583, 91158, 105611, 85583; BAMZ 1995-
132-003; FMNH 224903; MCZ Gould Station 1 (fossil), Gould Station 4 (fossil),

Map 11. Vertigo marki.

December 30, 2000 Bieler G Slapcinsky DI

Map 12. Vertigo numellata.

Gould Station 11 (fossil), Gould Station 21 (fossil), Gould Station 21b (fossil),
Gould Station 30 (fossil), Gould Station 41 east (fossil), Gould Station 48 (fossil),
Gould Station 63 lower (fossil), Gould Station 73 (fossil).

Note: Endemic - Originally described from fossil material, this is the most common
living Vertigo species on Bermuda. Recent specimens were first reported by
Vanatta (1911) from collections by Stewerdson Brown (ANSP 91158). Although
previously considered similar to Vertigo oralis Sterki, 1898 (Pilsbry, 1919), and V.
ovata Say, 1822 (Gulick, 1904), the strong contraction and large oblique crest
behind the shell aperture as well as the long lower palatal fold indicate this
species’ close affinity with V. bermudensis. Both V. numellata and V. bermudensis
have been collected around caves and not along the margins of wetlands as would
be expected of V. oralis or V. ovata. Shells of V. numellata differ only slightly from
those of V. bermudensis, having less deeply impressed sutures, larger size and a
stronger basal fold.

STROBILOPSIDAE
Strobilops hubbardi (A. D. Brown, 1861)
(Map 13, Fig. 12)

Helix Hubbardi A. D. Brown, 1861: 333, figs. 1, 2 (Texas).

Strobilops hubbardi, - Gulick, 1904: 413 (fossil); Verrill, 1906: 169 (“not
known as a recent shell in Bermuda”); Vanatta, 1911: 666; Vanatta,
1912: 12; Vanatta, 1924: 7; Peile, 1926: 89 (fossil); Pilsbry, 1948: 865-
868; Gould, 1970b: 490, fig. 1g (fossil).

Strobilops pilsbryi Morrison, 1953: 53-54, pl. 6, figs. 1-3 (Bermuda), new
synonymy.

Bermuda Material Studied: BAMZ 1995-133-009; FMNH 227033; MCZ Gould
Station 4 (fossil), Gould Station 21b (fossil), Gould Station 30 (fossil), Gould

LL NEMOURIA No. 44

Figure 12. Strobilops hubbardi. - BAMZ 1995-133-009, Bermuda; diameter =

2.7 mm.

Station 41 east (fossil), Gould Station 63 (fossil); USNM 251195 (paratypes of
Strobilops pilsbryi). Only one empty shell was located in this latest survey (FMNH
226908, Walsingham Pond).

Note: Native Non-Endemic - This species ranges over southeastern North America
from coastal Georgia south to Central America and east to Jamaica, Cuba and the
Bahamas (Pilsbry, 1948).

Adult Strobilops hubbardi specimens from Bermuda usually have four basal

Map 13. Strobilops hubbardi.

December 30, 2000 Bieler & Slapcinsky US

lamellae and a weak columellar lamella. Morrison (1953), citing the presence of
this columellar lamella, larger shell size and umbilical width, described Strobilops
pilsbryi as endemic to Bermuda. However, mainland populations of Strobilops
hubbardi also contain individuals with such lamellae (UF 110589). Furthermore,
there is ontogenetic variation in the number and strength of lamellae in Strobilops
hubbardi (UF 110593). Only the second and fourth basal lamellae are found in
many immature individuals (see Pilsbry, 1948, p. 849, for numbering and
terminology of Strobilops lamellae). Four lamellae are found in most adults,
however some adults do not develop all lamellae (see Pilsbry, 1948, fig. 468). The
columellar lamella is the smallest and latest lamella to develop. The prevalence of
this structure in the Bermuda populations is likely due to the high calcium
content of ‘many of Bermudas soils allowing complete and pronounced
development of apertural barriers.
This species is usually found on rotting logs and branches in open woods.

VALLONIIDAE
Vallonia pulchella (Miller, 1774)
(Map 14, Fig. 13)

Helix pulchella Miller, 1774: 30-31 (Europe).

? Helix paludosa da Costa, 1778: 59 [see Gerber, 1996: 68; Gerber erroneously
stated “Locus typicus’”; however, da Costa (1778: 59) clearly indicated
several localities in Britain, “in Oxendon and Arthingworth, in
Northamptonshire” ];

? Helix palludosa [sic; no author given], - Prime, 1853: 55; Jones, 1859: 106
[citing Prime];

? Helix paludosus Say [sic|—Tristram, 1861: 405;

Helix putchella [sic] Miller, - Rein, 1867: 77;

Helix pulchella, - Jones, 1876: 138; Bland, 1881: 256; Bland, 1895: 271;

Helix (Vallonia) pulchella, - Heilprin 1889: 182;

Vallonia pulchella, - Sterki, 1893: 278; Pilsbry, 1900: 495; Verrill, 1902: 732,
733, fig.; Bryant, 1905: 129-130; Vanatta, 1911: 665-666; Peile, 1926:
89; Pilsbry, 1948: 1023-1025; Glaubrecht, 1990: 218; Gerber, 1996: 56
[based on Pilsbry, 1948).

Bermuda Material Studied: BAMZ 1995-132-005.

Note: Accidentally Introduced - “Helix paludosa” of Prime's (1853) list and “Helix
paludosus” Say of Tristram's list (1861) probably refer to this species rather than to

24 NEMOURIA No. 44

Figure 13. Vallonia pulchella. - BAMZ 1995-132-005, Bermuda; diameter = 2.3

mim.

Polygyra plana (Dunker, 1843) as surmised by Pilsbry (1900). The source of this
confusion stems from Prime's omission of the species authority for his record of Helix
paludosa. Tristram later amended Prime’s record to paludosus and erroneously
attributed it to Say. Unfortunately, H. paludosa has two homonyms: Helix paludosa
Pfeiffer, 1839, was associated with Polygyra plana (Dunker, 1843) by Pilsbry (1900:
506), while Helix paludosa da Costa, 1778, is a synonym of Vallonia excentrica Sterki,
1893, a species very similar to and often confused with V. pulchella. Both Vallonia
pulchella and Polygyra plana are found on Bermuda. Pilsbry may have overlooked H.

Map 14. Vallonia pulchella.

December 30, 2000 Bieler & Slapcinsky 25

paludosa da Costa when he concluded that the early authors were looking at Polygyra
plana. This view is supported by the fact that Tristram recorded both paludosa and
microdonta (see Polygyra plana synonymy).

Vallonia excentrica ranges over much of the Northern Hemisphere. In North
America it is native to the Northeast from southern Ontario (Oughton, 1948) to
Missouri (Hubricht, 1985), and has been introduced to Texas, New Mexico, the
Rocky Mountain States, California, and Oregon (Bequaert & Miller, 1973). It is
found in open areas including meadows, marshes and sand dunes (Kerney &
Cameron, 1979), as well as on roadsides and lawns (Hubricht, 1985), usually
under cover of boards, wood or on_ house
foundations and other cement structures.

ACHATINOIDEA
FERUSSACIIDAE
Cecilioides acicula (Miller, 1774)
(Map 15, Fig. 14)

Buccinum acicula Miller, 1774: 150-151
(Europe).

Bulimus acicula, - Bland, 1861: table 2;

Achatina acicula, - Bland, 1861: 351;

Cionella acicula, - Jones, 1876: 139; Bartram,
1878: 108; Bland, 1881: 256;

Caecilianella acicula, - Kobelt, 1880: 257, 286;
Glaubrecht, 1990: 218.

Caecilianella (Cionella) acicula, - Heilprin, 1889:
183;

Coecilianella acicula, - Bland, 1895: 271,

Figure 14. Cecilioides
acicula. - BAMZ 1995-
133-002) Bermuda;
height = 4.3 mm.

Map 15. Cecilioides acicula.

26 NEMOURIA No. 44

Cecilioides acicula, - Bryant, 1905: 130; Pilsbry, 1946: 185, 186;
Caecilioides acicula, - Pilsbry, 1900: 497; Verrill, 1902: 731, 732, fig. 78a;
Vanatta, 1911: 665-667; Peile, 1926: 90; Abbott, 1989: 84, fig.

Bermuda Material Studied: BAMZ 1995-133-002; FMNH 227024.

Note: Accidentally Introduced - First recorded in Bermuda by Bland in 1861, this
small snail is native to Europe. Its burrowing habits might explain the small
number of collection records on Bermuda and its easy transport between
greenhouses and plant nurseries across the world.

SPIRAXIDAE
Euglandina rosea (Férussac, 1821)
(Map 16, Fig. 15)

Helix (Cochlicopa) rosea Férussac, 1821: 50 [or p. 54, depending on issue]
(Florida).

Euglandina rosea, - Bennett & Hughes, 1959: 432; Hughes, 1962; Hughes,
1963; Simmonds & Hughes, 1963: 219 ff.; Hughes, post 1968: 1;
Monkman, 1982a: 1; Monkman, 1982b: 71; Cock, 1985: 138;
Glaubrecht, 1990: 221, fig. 11.

Figure 15. Euglandina rosea. - left: BAMZ 1994-118-010, Pembroke Parish, height
= 48.6 mm; right: FANH 224776, Paget Parish, Bermuda, height = 50.0 mm.

December 30, 2000 Bieler & Slapcinsky 24

Map 16. Euglandina rosea.

Bermuda Material Studied: BAMZ 1998-166-002, 1998-166-005, 1998-166026,
1998-166-039, 1998-166046, 1998-166-051; DMNH 210302, 210308, 210316,
210323, 210338; FMNH 224761, 224792, 224814, 224831, 224844, 224853,
224855, 224861, 224866, 224869, 224879, 224896, 224905, 224913, 224920,
227001.

Note: Intentionally Introduced - This native of the southeastern United States was
introduced on many islands of the Pacific as a biological control against the
agricultural pest snail Achatina fulica. Euglandina rosea specimens - mostly from
Florida (Davis & Butler, 1964), not from Cuba as stated by Cock (1985) - were
released in the Hawaiian Islands in 1955 and were established there by 1957
(Krauss, 1964). In 1958, the Bermuda Department of Agriculture received a
shipment of about 50 adult Euglandina rosea from Hawaii via the West Indian
Station CIBC (Commonwealth Institute of Biological Control) in Trinidad to
breed and release in efforts to control Otala lactea and, secondarily, Rumina
decollata, two accidentally introduced horticultural pests. Later that year, after
brief tests to ascertain that Euglandina would consume Rumina, Otala and other
local snails when captive with them, Euglandina rosea was released on Bermuda.
Between January and April of 1958, 329 snails were released, and by 1960, a total
of 841 animals had been set free (Hughes, 1962 unpubl.). Releases of specimens
received via Hawaii and Trinidad continued for a few more years. By 1962, the
Department of Agriculture reported “excellent control of Otala lactea [had] been
achieved.” At that time, E. rosea had become so abundant that the Department
exported nearly 1,000 specimens to various countries, ranging from the Bahamas
to India (Hughes, 1962 unpubl.). Although ultimately failing to control either
Otala or Rumina, Euglandina has become widespread in Bermuda and might be
responsible for the decline of several of Bermuda's native snails, including the
members of the now nearly extinct endemic genus Poecilozonites.

28 NEMOURIA No. 44

Figure 16. Allopeas gracile. - Figure 17. Allopeas micrum. - FMNH

FMNH 227044, Sandy’s Parish, 30195, St. George’s Parish,

Bermuda; height = 10.8 mm. Bermuda; height = 6.3 mm.
SUBULINIDAE

Allopeas gracile (Hutton, 1834)
(Map 17, Fig. 16)

Bulimus? gracilis Hutton, 1834: 84, 93 (India).

Opeas gracile, - Vanatta, 1911: 666-667; Peile, 1926: 90;

Opeas (Opeas) gracile, - Haas, 1952: 104;

Allopeas gracile, - Naggs, 1992: 258-259; ICZN, 1994: 162-163; Naggs, 1994:
187.

Map 17. Allopeas gracile.

December 30, 2000 Bieler & Slapcinsky 29

Bermuda Material Studied: BAMZ 1994-118-016; FMNH 227044.

Note: Accidentally Introduced - Probably originally from the Neotropical region
(Solem, 1964: 134), Allopeas gracile has been introduced worldwide in cultivated
areas of the tropics and in greenhouses and was first described from India. Deisler
& Abbott (1984: 13) stated that this “species probably is the most widely
distributed land snail.” In North America, this species is found from South
Carolina to Texas (Karlin & Naegele, 1966; Dundee, 1970).

Allopeas micrum (Orbigny, 1835)
(Map 18, Fig. 17)

Helix micra Orbigny, 1835: 9 (Central America).

Opeas octonoides (C. B. Adams, 1845), - Pilsbry, 1900: 497; Verrill, 1902: 732
[Bulimus octonoides C. B. Adams, 1845, non Bulimus octonoides Orbigny in
Sagra, 1841];

Opeas micra, - Vanatta, 1911: 666-667; Peile, 1926: 90;

Opeas (Opeas) micrum, - Haas, 1952: 104.

Bermuda Material Studied: FMNH 30194, 30195.

Note: Accidentally Introduced - Probably native to Central America, the West
Indies, and northern South America, this species has been introduced into the
southeastern United States and the Bahamas (Deisler & Abbott, 1984). On
Bermuda, Allopeas micrum was first collected by the Heilprin party in 1888, but
not included in their publication of 1889 (Pilsbry, 1900). This species is often
found near greenhouses and in urban areas.

Map 18. Allopeas micrum.

30 NEMOURIA No. 44

Figure 18. Opeas hannense. - FMNH Figure 19. Obeliscus swiftianus. -
278023, St. George’s Parish, Bere FMNH 30197, St. George’s Parish,
muda; height = 9.2 mm. Bermuda; height = 7.3 mm.

Opeas hannense (Rang, 1831) NEW RECORD
(Map 19, Fig. 18)

Helix hannensis Rang, 1831: 41-42, pl. 3, fig. 8 (Cape Vert peninsula, Senegal).

Opeas hannensis [sic] (Rang, 1831), - Proschwitz, 1983: 283; Proschwitz, 1994:
184.

Bermuda Material Studied: FMNH 278023 (Mullet Bay Park), 227034 (city of

Hamilton).

Map 19. Opeas hannense.

December 30, 2000 Bieler & Slapcinsky on

Note: Accidentally Introduced - A native of the New World tropics, this species is
now widely distributed. In North America it has been introduced to southern
Florida and greenhouses in New York, Pittsburgh and Chicago (Pilsbry, 1946).
Bulimus pumilus Pfeiffer, 1840, first described from Cuba, is a synonym
(Proschwitz, 1994: 184). This species is here recorded from Bermuda for the first
time.

Obeliscus swiftianus (Pfeiffer, 1852)
(Map 20, Fig. 19)

Bulimus Swiftianus Pfeiffer, 1852: 150 (St. Thomas).

Opeas swiftianum, - Pilsbry, 1900: 497, pl. 62, fig.5; Verrill, 1902: 731, 732,
fig. 78;
Obeliscus swiftianus, - Vanatta, 1911: 665; Vanatta, 1923: 33; Peile, 1926: 90.

Bermuda Material Studied: BAMZ 1998-166-004, 1998-166-018, 1998-166-024,
1998-166-031, 1998-166045, 1998-166-077, 1998-166-082, 1998-166-085, 1998-
166-086, 1998-166-087; DMNH 210251, 210252, 210264, 210278, 210282;
FMNH 3632, 13493, 29794, 30197, 224773, 224811, 224812, 224822, 224835,
224846, 224857, 224863, 227023, 295082.

Note: Accidentally Introduced - Like the previous two species, Obeliscus swiftianus is
native to the American tropics and has been widely distributed through
commerce. This species was collected by the Heilprin expedition but not included
in their 1889 publication (Pilsbry, 1900). Obeliscus swiftianus is usually found
under cover in urban areas and near horticulture.

Map 20. Obeliscus swiftianus.

SYP NEMOURIA No. 44

Figure 20. Rumina decollata. - FMNH 224777; Paget Parish, Bermuda; left: height
= 28.5 mm; right (subadult): diameter = 18.4 mm.

Rumina decollata (Linné, 1758)
(Map 21, Fig. 20)

Helix decollata Linné, 1758: 773 (Europe).

Subulina decollata, - Bartram, 1878: 108;

Bulimulus decollatus, - Heilprin, 1889: 183; Frith, 1891: 14; Peniston, 1891:
21;

Stenogyra decollata, - Aldrich, 1889: 9; Lane, 1891: 3; Bland, 1895: 271;

Map 21. Rumina decollata.

December 30, 2000 Bieler @ Slapcinsky 33

Rumina decollata, - Riley & Howard in Hastings, 1892: 334; Johnson, 1900:
117; Pilsbry, 1900: 496; Verrill, 1902: 730, 731, figs. 75a-c, 76a, b, 79e;
Vanatta, 1911: 665-668; H. H. Smith, 1912: 4; Honigmann, 1914: 376;
Vanatta, 1923: 33; Peile, 1926: 90; Pilsbry, 1946: 170-172; Bennett &
Hughes, 1959: 432; Simmonds G& Hughes, 1963: 219-220; Monkman,
1982a: 1; Monkman, 1982b: 71; Bennett et al., 1985: 138; Cock, 1985:
138.

Bermuda Material Studied: BAMZ 1998-166-007, 1998-166-013, 1998-166-040,
1998-166058, 1998-166-062, 1998-166-065, 1998-166-076, 1998-166-099, 1998-
166-105, 1998-166-108, 1998-166-110; DMNH 210271, 210294, 210300,
210313, 210324, 210337; FMNH 3630, 8670, 8836, 13501, 13527, 13531,
13537, 13556, 13582, 13591, 13679, 13686, 16473, 16474, 29793, 41014,
112230, 135326, 216093, 224755, 224762, 224793, 224794, 224826, 224839,
224870, 224889, 224900, 224907, 227057, 227008, 227009, 227027.

Note: Accidentally Introduced - Rumina decollata, a ground dweller, is often found in
gardens and orchards and is among the most common of Bermuda's snails.
Rumina decollata is a common pest species introduced worldwide to subtropical
areas, and is well established in the southern U.S. (Dundee, 1970). This native of
the Mediterranean region is believed to have been introduced on plants from
Madeira or the Cape Verde Islands by Bermuda Governor John H. Lefroy in the
late 1870s (Frith, 1891: 16). By 1879, Rumina was abundant in Hamilton
(Aldrich, 1889: 9, cited it as “so common it threatens to become injurious to the
crops there”), and a few years later Hastings (1892: 334) described it as “literally
eating up the island.” By the 1950s it was common throughout the larger islands
of Bermuda and considered a minor crop pest. Efforts were made by the Bermuda
Department of Agriculture to biologically control crop pests including Rumina
decollata using several species of predatory snails (Bennett G Hughes, 1959).
Although original reports were promising (Simmonds & Hughes, 1963), later
Bermuda Department of Agriculture surveys found little or no evidence of control
(Monkman, 1982a). Fisher et al. (1980) showed that this omnivorous species itself
is a predator of half-grown Brown Garden Snails in California (“Helix” aspersa
Miller, 1774; recently placed in genus Cantareus by Giusti et al., 1995: 491).
Rumina decollata is still advocated as a control agent in California’s citrus industry
(UCIPM, 1999). Its impact on Bermuda’s native snail population is unknown.

34 NEMOURIA No. 44

Subulina octona (Bruguiére, 1789)
(Map 22, Fig. 21)

Bulimus octonus Bruguiére, 1789: 325
(Antilles).

? “Achatina aetona [sic] “Chemnitz”, - Prime,
1853: 76;

Stenogyra octona, - Bland, 1881: 256; Aldrich,
1889: 9; Heilprin, 1889: 183; Bland,
1895: 271;

Subulina octona, - Pilsbry, 1900: 497, pl. 62,
fig. 4; Verrill, 1902: 731, 732, fig. 77;
Vanatta, 1911: 665-667; Peile, 1926: 90.

Bermuda Material Studied: BAMZ 1998 166-
006, 1998- 166- 011, 1998- 166- 019, 1998- 166- Figure’ 21-{Snbulincene conc.
020, 1998- 166- 023, 1998- 166 028, 1998-166 = _ pyNH PASCO South:
037, 1998- 166- 061, 1998- 166- 071, 1998- 166- ampton Parish, Bermuda;
072, 1998- 166- 073, 1998- 166- 078, 1998- 166- height = 13.3 mm.

090, 1998- 166- 103, 1998 166 112, 1998 166-

125; DMNH 210242, 210247, 210257, 210267, 210283, 210285, 210286,
210297, 210298, 210305, 210309, 210311, 210326; FMNH 104119, 224770,
224771, 224783, 224803, 224804, 224827, 224843, 224852, 224854, 224860,
224860, 224862, 224865, 224874, 224890, 224901, 224908, 227010, 227011,
227020, 227026.

Note: Accidentally Introduced - Subulina octona often is listed as a native of Latin
America (Robinson, 1999), but other species of this genus are native to Africa.
Subulina octona is now widely distributed in tropical areas, including southern
Florida and the Bahamas (e.g., Deisler & Abbott, 1984). It was first recorded in
Bermuda by Bland in 1881 and is now found throughout the larger islands of
Bermuda under boards, garbage and stones.

Map 22. Subulina octona.

December 30, 2000 Bieler @ Slapcinsky 35

STREPTAXOIDEA
STREPTAXIDAE
Gonaxis kibweziensis (E. A. Smith, 1894)
(Fig. 22)

Streptaxis Kibweziensis E. A. Smith,
1894: 165, 166, fig. 1 (Kenya).

Gonaxis (Streptaxis) kibweviensis, -

Simmonds & Hughes, 1963:

DDD:
Gonaxis kibweziensis, - Monkman,
1982a: 1; Cock, 1985: 138;
Eustreptaxis kibweziensis, - Richard-

son, 1988: 198.
Bermuda Material Studied: None.

Note: Intentionally Introduced - This
predatory snail was brought to Hawaii in
1952 from Diana Beach, Kenya by
Hawaii's Department of Agriculture
(Krauss, 1964) in attempts at controlling
populations of the introduced Giant

Figure 22. Gonaxis kibweziensis. -
BPBM 217478, Marianas Islands;
height = 19.8 mm.

African Snail, Achatina fulica Bowdich, 1822. In 1959, about one hundred
Gonaxis kibweziensis specimens were sent from Hawaii via the West Indian Station
CIBC (Commonwealth Institute of Biological Control) to Bermuda's Department
of Agriculture to be used to control Otala lactea and Rumina decollata. In 1960,
these snails were released at four sites (Simmonds & Hughes, 1963). Subsequent
monitoring of these sites by Bermuda's Department of Agriculture has failed to
recover individuals of this species (Monkman, 1982a), as has this survey.

Gonaxis quadrilateralis (Preston, 1910)
(Map 23, Fig. 23)

Ennea quadrilateralis Preston, 1910: 527, pl. 7, fig. 2 (British East Africa

[Kenya]).

Gonaxis quadrilateralis, - Hughes, post 1968: 1; Monkman, 1982a: 1;
Monkman 1982b: 71; Cock 1985: 138;

36 NEMOURIA No. 44

Figure 23. Gonaxis quadrilateralis. Figure 24. Gulella bicolor. -
- Bermuda Department of ANSP 48654, ex Heilprin
Agriculture, now BAMZ; Expedition 1888, Ber-
Bermuda; height = 25.0 mm. muda; height = 6.7 mm.

Eustreptaxis quadrilateralis, - Richardson, 1988: 202.

Bermuda Material Studied: BAMZ 1995-127-012, 1998-166-014, 1998-166-088;
DMNH 210276; FMNH 224785, 227036, 227037, 227038.

Note: Intentionally Introduced - Like the previous species, Gonaxis quadrilateralis was
introduced to Bermuda from Hawaii by the Bermuda Department of Agriculture.
G. quadrilateralis was collected from Kwale, Kenya, and brought to Hawaii in 1957
in attempts to control Achatina fulica (see Krauss, 1964). In 1968 and again in
1972, G. quadrilateralis specimens sent from Hawaii were released at eight sites on

Map 23. Gonaxis quadrilateralis.

December 30, 2000 Bieler & Slapcinsky 37

Bermuda in continuing efforts at controlling Rumina decollata and Otala lactea.
[Cock (1985: 138) cited 260 specimens obtained from Hawaii in 1968 and 1972].
Since that time, G. quadrilateralis individuals have been collected at several of the
original release sites (Monkman, 1982a). This study records the first spread of G.
quadrilateralis beyond the original release sites.

Gulella bicolor (Hutton, 1834)
(Fig. 24)

Pupa bicolor Hutton, 1834: 86, 93 (India).

Ennea bicolor, - Pilsbry, 1900: 499; Verrill, 1902: 732;
Huttonella bicolor, - Peile, 1926: 90;
Gulella bicolor, - Naggs, 1989: 165-168 (discussion of generic placement).

Bermuda Material Studied: ANSP 48654.

Note: Accidentally Introduced - Native to the Indian subcontinent (Naggs, 1989),
this species has been introduced around the world in the tropics and subtropics
including the southeastern United States (Dundee,
1970; Auffenberg & Stange, 1986) and Caribbean
(van der Schalie, 1948). Records of this species on
Bermuda are based on a single shell (ANSP 48654),
collected by the Heilprin party in 1888 and first
mentioned in publication by Pilsbry in 1900. The
species was not found in the present survey and may
not be established on Bermuda (note, however, that
its burrowing habits make it difficult to find).

Streptostele musaecola (Morelet, 1860) NEW
RECORD
(Map 24, Fig. 25)

Achatina musaecola Morelet, 1860: 190 (Gui-

nea). Figure 25. Streptostele mus-
aecola. - FANH 227045,
Bermuda Material Studied: FANH 227045. Paget Parish, Bermuda;

height = 6.0 mm.

38 NEMOURIA No. 44

Map 24. Streptostele musaecola.

Note: Accidentally introduced - Originally described from West Africa, this species
was accidentally introduced to several islands in the Pacific (Solem, 1989). A
specimen from the city of Hamilton (FMNH 227045), collected in 1992, is the

first record of this species from the western Atlantic region.

RHYTIDOIDEA
RHYTIDIDAE
Natalina cafra (Férussac, 1821)
(Map 25, Fig. 26)

Helix (Helicophanta) cafra Férussac, 1821: 25 [or p. 29, depending on issue]
(South Africa).

Natalina cafra, - Simmonds & Hughes, 1963: 221;

Natalina caffra [sic] - Hughes, 1962; Hughes, 1963; Hughes, post 1968: 1;
Monkman, 1982a: 1; Cock, 1985: 138.

Figure 26. Natalina cafra. - left: FMNH 199492, Natal, South Africa, diameter =
65.4 mm; right: FMNH 76882, South Africa, diameter = 70.1 mm.

December 30, 2000 Bieler & Slapcinsky 39

Map 25. Natalina cafra.

Bermuda Material Studied: None.

Note: Intentionally Introduced - About sixty individuals of this South African species
were released in Smith’s Parish on Bermuda beginning in 1960, as part of
continuing efforts to control populations of Otala and Rumina (Simmonds &
Hughes, 1963; Hughes, post 1968). Shipments were discontinued in 1962 when
surveys recorded only one live individual at Bermuda's Arboretum, one of the
original release sites (Hughes, unpubl. reports for 1962, 1963). Since that time no
additional individuals have been recorded (Monkman, 1982a).

ARIONOIDEA
PUNCTIDAE
Punctum bristoli (Gulick, 1904) NEW EXTANT RECORD
(Map 26, Fig. 27)

Zonitoides bristoli Gulick, 1904: 421, pl. 36, fig. 13 (Bermuda, fossil).

Zonitoides bristoli, — Verrill, 1906: 168, fig. 51 (after Gulick; fossil); Vanatta,
1912: 12 (fossil); Hughes, post 1969: 1 (fossil); Gould, 1970b: 491, fig. 1f
(fossil);

Punctum bristoli, - Vanatta, 1924: 7 (fossil);

Map 26. Punctum bristoli.

40 NEMOURIA No. 44

Figure 27. Punctum bristoli. - BAMZ 1995-132-013, Bermuda, diameter = 1.28

mm.

Punctum (?) bristoli, - Peile, 1926: 89 (fossil).

Bermuda Material Studied: ANSP 85572 (fossil); BAMZ 1995-132-013 (fossil);
FMNH 227042; MCZ Gould Station 1 (fossil), Gould Station 4 (fossil), Gould
Station 21 (fossil), Gould Station 21b (fossil), Gould Station 41 east (fossil),
Gould Station 48 (fossil), Gould Station 63 (fossil).

Note: Endemic - This study records the first Recent specimens of this small-shelled
species (FMNH 227042). Pleistocene specimens are commonly found in siftings
from Bermuda's fossil deposits. The lack of Recent records of this species is
probably due to insufficient efforts to sort such micromollusks from soil and leaf
litter samples.

HELICODISCIDAE
Helicodiscus singleyanus (Pilsbry, 1889)
(Map 27, Fig. 28)

December 30, 2000 Bieler & Slapcinsky 4]

---, Pilsbry, 1888: pl. 17, figs., M, M, M;
Zonites singleyanus Pilsbry, 1889: 84 (Texas).

Zonitoides singleyanus, - Peile, 1926: 89.

Bermuda Material Studied: FMNH 227047.

Figure 28. Helicodiscus singleyanus. - FMNH 227047, Paget Parish, Bermuda;

diameter = 2.0 mm.

Map 27. Helicodiscus singleyanus.

42 NEMOURIA No. 44

Note: Accidentally Introduced - Native to eastern and central North America
(Hubricht, 1985), this species has been introduced throughout Europe and into
North Africa (Seddon & Holyoak, 1993). It is subterranean and can be sifted
from loose soil around the roots of grasses and herbaceous plants along roadsides
and other open areas. This elusive species was previously recorded from Bermuda
only by Peile (1926). The present survey found one specimen in the city of
Hamilton (FMNH 227047). Like other soilinhabiting species, Helicodiscus

singleyanus is probably much more common than the sparse records indicate.

SUCCINEOIDEA
SUCCINEIDAE
Succinea bermudensis Pfeiffer, 1857
(Map 28, Fig. 29)

Succinea bermudensis Pfeiffer, 1857: 110 (Bermuda).

Succinea ---, Redfield, 1853: 17; Tristram, 1861: 405;

Succinea Bermudensis Prime, 1853: 55 [nomen nudum];
Succinea Bermudensis Jones, 1859: 107 [nomen nudum; based on Prime’s list,

but erroneously indicated as “n.s.”];

Figure 29. Succinea bermudensis. - BAMZ 1995-132-006, Bermuda; left, height =
12.6, right, height = 12.0 mm.

December 30, 2000 Bieler & Slapcinsky 43

Succinea bermudensis, - Pfeiffer, 1859: 817; Bland, 1861: table 2, p. 351; Rein,
1867: 76; Jones, 1876: 139; Médrch, 1878: 3; Kobelt, 1880: 286; Bland,
1881: 256 E. A. Smith, 1884: 277; Heilprin, 1889: 182; Bland, 1895:
270; Gulick, 1904: 421; Davis, 1904: 125; Verrill, 1906: 171; Vanatta,
1911: 665-668; Peile, 1926: 91; Tomlin, 1932: 98-100, figs. C, D;
Hughes, post 1968: 1; Gould, 1970b: 500 (fossil);

Succinea fulgens Lea, - Bland, 1861: table 2, p. 351; Jones, 1876: 139; Kobelt,
1880: 286; Bland, 1881: 256; Heilprin, 1889: 182; Bland, 1895: 270
[non Succinea fulgens Lea, 1841);

Succinea margarita Pfeiffer, - Bland, 1861: 351; Jones, 1876: 139; Kobelt,
1880: 286; Bland, 1881: 256; Heilprin, 1889: 182; Bland, 1895: 270,
table 2 [non Succinea margarita Pfeiffer, 1853];

Succinea texasiana Pfeiffer, 1848, - Jones, 1864: 9 [non Succinea texasiana
Pfeiffer, 1848];

Sucinea [sic] putris, - Bartram, 1878: 108 [non Helix putris Linné, 1758: 774];

Succinea barbadensis Guilding, 1828, - Pilsbry, 1900: 502; Verrill, 1902: 729,
732, 733, figs. 80a, b; Vanatta, 1923:33; [? non Succinea barbadensis
Guilding, 1828];

Succinea (Tapada) bermudensis, - Clessin in Pfeiffer, 1879: 416;

Succinea gabrieli Tomlin, 1932: 99-100, figs, A, B, pl. 9A (Bermuda);

Succinea barbadensis “Gould”, - Kutchka, 1934: 295;

Succinia [sic] bermudensis, - Hughes, post 1969: 1.

Bermuda Material Studied: BAMZ 1998-166-052, 1998-166-057, 1998-166-113;
DMNH 210268, 210290, 210319; FMNH 3628, 13483, 13496, 13509, 13511,
13516, 13519, 13526, 13549, 13569, 13572, 13575, 13578, 13681, 16462
(fossil), 16463, 29800, 30191, 30216, 41016, 49546, 104491, 104612, 141807,
224780, 224798, 224828, 224841, 224891, 224902, 224909, 227012.

Note: Endemic - Pilsbry (1900: 492) considered this species to be conspecific with
Succinea barbadensis of Barbados. However, identifications based on shell

Map 28. Succinea bermudensis.

44 NEMOURIA No. 44

characteristics in species with
reduced shells, such as those of the
succineids, are notoriously unreli-
able. Comparisons of anatomical
features are necessary to determine
if S. barbadensis and S. bermudensis
are indeed conspecific. Pending
this, we have considered the two
nominal species distinct and
Succinea bermudensis endemic to
Bermuda. Succinea gabrieli Tomlin,
1932, introduced based on slight
differences in shell morphology and
radular tooth counts, is most likely
conspecific with S. bermudensis.
Succinea bermudensis is common on
grasses and on the bark of trees,
often in open areas, and is also
commonly found as a Pleistocene
fossil.

Figure 30. "Euconulus" turbinatus. -
ANSP 91153, Hamilton Parish,
Bermuda; height = 3.65 mm.

LIMACOIDEA
HELICARIONIDAE
"Euconulus" turbinatus Gulick, 1904
(Map 29, Fig. 30)

Euconulus turbinatus Gulick, 1904: 420, pl. 36, figs. 8-10 (Bermuda, fossil).

Euconulus turbinatus, - Verrill, 1906: 168, figs. 55a, b (types; “not known to be

living”); Gould, 1970b: 491;

Map 29. "Euconulus" turbinatus.

December 30, 2000 Bieler & Slapcinsky 45

Kaliella turbinatus, - Vanatta, 1911: 665;
Kaliella turbinata, - Pilsbry G@ Vanatta, 1909: 64; Vanatta, 1911: 666-667;
Peile, 1926: 89.

Bermuda Material Studied: ANSP 91153; FMNH 43379; MCZ Gould Station 1
(fossil), Gould Station 41 east (fossil).

Note: Endemic - Described from fossils on Bermuda and tentatively placed in the
genus Euconulus (Gulick, 1904), this species was later found living around Church
Cave (Pilsbry & Vanatta, 1909). Using shell and radular characteristics, Godwin
Austen (in Pilsbry & Vanatta, 1909) determined this species to belong to the
Asian genus Kaliella, leading Pilsbry @ Vanatta to consider Gulick’s specimens
subfossil and the species a very early introduction. Studies of the Pleistocene
fossils of Bermuda confirm this species’ presence in Pleistocene deposits and its
endemic status (Gould, 1971b). Anatomical material is needed to establish its
generic position and relationships with other helicarionids.

AGRIOLIMACIDAE
Deroceras laeve (Miller, 1774)
(Map 30, Fig. 31)

Limax laevis Miller, 1774: 1-2 (Europe).

Agriolimax campestris (Binney, 1842), - Cockerell in Bland, 1895: 271
[described as Limax campestris Binney, 1842];

Figure 31. Deroceras laeve. - FANH 224864, St. Georges Parish; lateral view; body
length = 14 mm.

Agriolimax laevis, - Pilsbry, 1900: 501; Verrill, 1902: 734; Vanatta, 1911: 665-
668;
Malino laevis, - Peile, 1926: 90.

46 NEMOURIA No. 44

Map 30. Deroceras laeve.

Bermuda Material Studied: BAMZ 1998-166-092, 1998-166-094, 1998-166-098,
1998-166-102, 1998-166-107, 1998-166-109, 1998-166-123; DMNH 210249,
210254, 210256, 210263, 210273, 210335, 210336; FMNH 224774, 224816,
224864, 224872, 224875, 224893, 227055, 227056.

Note: Accidentally Introduced - A very widely distributed slug, Deroceras laeve is
found throughout much of the Northern Hemisphere. Found in gardens or along
foundations it is most commonly seen in rainy weather or at night. Deroceras laeve
was first collected in Bermuda by the British ‘Challenger’ expedition in 1873.

LIMACIDAE
Limacus flavus (Linné, 1758)
(Map 31, Fig. 32)

Ope cca RO

Figure 32. Limacus flavus. - after Moquin-Tandon, 1855: pl. 3, fig. 3; body length
= 75 to 100 mm.

Map 31. Limacus flavus.

December 30, 2000 Bieler & Slapcinsky 47

Limax flavus Linné, 1758: 652 (Europe).

Limax flavus, - Pilsbry, 1900: 501; Verrill, 1902: 734, fig. 83; Peile, 1926: 90;
Hughes 1968: 1;

Limacus flavus (Linné, 1758), - Falkner in Fechter & Falkner, 1990: 190
[European distribution and placement in Limacus].

Bermuda Material Studied: FMANH 198685.

Note: Accidentally Introduced - Found in urban areas around the world, this native
of Europe was first collected on Bermuda by Verrill in 1898. Incomplete locality
data on most Bermudan specimens obscures the distribution of this species on
Bermuda. This very large slug can be found at night, especially on stone fences
and in gardens. Dissection of FMNH 198685 confirmed the specific identity of
this slug, which is often confused with Limacus maculatus (Kaleniczenko, 1851).

MILACIDAE
Milax gagates (Draparnaud, 1801)
(Map 32, Fig. 33)

Limax gagates Draparnaud, 1801: 100 (France).

Figure 33. Milax gagates. - after Moquin-Tandon, 1855: pl. 2, fig. 1; body length =
50 to 60 mm.

Map 32. Milax gagates.

48 NEMOURIA No. 44

Limax gagates, - E. A. Smith, 1884: 276;

Amalia gagates, - Bland, 1895: 271; Pilsbry, 1900: 501; Verrill, 1902: 734;

Milax sowerbyi (Férussac, 1823), var., - Vanatta, 1911: 665-668 [non Limax
sowerbii [sic] Férussac, 1823];

Milax sowerbyi, - Peile, 1926: 90;

Milax gagates, - Vanatta, 1911: 665-667; Peile, 1926: 90; Hughes, 1968: 1.

Bermuda Material Studied: FMNH 224759.

Note: Accidentally Introduced - Milax gagates was first collected on Bermuda by the
British ‘Challenger’ expedition in 1873 (E. A. Smith, 1884). A native of Europe,
it burrows in soil and feeds on plant roots. Introduced around the world in
potted plants (Karlin G@ Naegele, 1966), it has become so ubiquitous in
greenhouses it has earned the common name “The Greenhouse Slug” (Pilsbry,
1948). It is placed in genus Tandonia by some authors.

ZONITIDAE
Hawaiia minuscula (Binney, 1841)
(Map 33, Fig. 34)

Helix minuscula Binney, 1841: 435-436 pl. 22, fig. 4 (Ohio).

Zonitoides minusculus, - Pilsbry, 1900: 501, pl.62, fig.17; Verrill, 1902: 729,
fig. 71; Gulick, 1904: 421 (Recent and “fossil”); Verrill, 1906: 168, figs.
52a, b (after Binney); Peile, 1926: 89; Hughes, post 1969: 1; Gould,
1970b: 491;

Zonitoides minuscula, - Vanatta, 1911: 665-668; Vanatta, 1923: 33; Vanatta,
1924: 7;

Hawaiia minuscula, - Riedel, 1980: 28-29.

Map 33. Hawaiia minuscula.

December 30, 2000 Bieler & Slapcinsky 49

Figure 34. Hawaiia minuscula. - FMNH 227046, Hamilton Parish, Bermuda;

diameter = 2.1 mm.

Bermuda Material Studied: BAMZ 1995-132-014; FMNH 30204, 30205, 41019,
227035, 227046.

Note: Accidentally Introduced - Commerce has transported this native of North
America around the world, as attested by the generic name, which is based on
specimens mistakenly thought native to Hawaii. Hawaiia minuscula is usually
found on bare ground in yards, fields and other open areas. Gould (1970b)
questioned the fossil record by Gulick (1904); Gulick (1904: 421) himself
expressed surprise at this singular occurrence and absence at all other sites studied
by him. The later references to Bermuda fossils of this species by Verrill (1906)
and Hughes (post 1969) are based on Gulick’s single record. Gulick’s collecting
site was probably contaminated with Recent material.

50 NEMOURIA No. 44

Figure 35. Oxychilus draparnaudi. - FMNH 13547, Paget’s Parish, Bermuda;

diameter = 10.9 mm.

Oxychilus draparnaudi (Beck, 1837)
(Map 34, Fig. 35)

Helix (Helicella) Draparnaldi [sic] Beck, 1837: 6 [new name for Helix nitida
Draparnaud, 1805, non Miller, 1774]; non Helix draparnaldi Cuvier,
1816, nec Helix draparnaudi Sheppard, 1823 [see Manganelli & Giusti,
1997: 149-150].

Helix cellaria Miller, 1774 - Prime, 1853: 77 [non Helix cellaria Miller, 1774];

Hyalina lucida (Draparnaud, 1801), - Verrill, 1902: 733, fig. 8la, b; Davis,
1904: 126; [described as Helix lucida Draparnaud, 1801; non Helix lucida
Pulteney, 1799];

Vitrea cellaria Miller, 1774, - Davis, 1904: 126 [non Vitrea cellaria Miller,
1774);

December 30, 2000 Bieler & Slapcinsky 51

Map 34. Oxychilus draparnaudi.

Vitrea lucida (Draparnaud, 1801), - Bryant, 1905: 131; Vanatta, 1911: 666-
667;

Helicella draparnaldi [sic], - Peile, 1926: 89;

Helicella draparnaldi, - Ellis, 1951: 126 [petition to place on Official List of
Names];

Helicella draparnaudi, - ICZN, 1955: 81, 87, 99 [placement of the emended
specific name on the Official List of Specific Names in Zoology; with
erroneous “original” generic combination];

Oxychilus draparnaudi, - Manganelli & Giusti, 1997: 148 ff. [proposed
conservation of specific name Oxychilus draparnaudi (Beck, 1837), based
on Helix draparnaldi).

Bermuda Material Studied: BAMZ 1998-166-038, 1998-166-041, 1998-166-117,
1998-166-118; DMNH 210260, 210261, 210288; FMNH 13503, 13558, 13577,
13680, 13684, 224757, 224767, 224799, 224823, 224848, 224886, 224898,
224906, 227006, 227007.

Note: Accidentally Introduced - This species was first collected on Bermuda in 1852
and reported as the similar species “Helix” cellaria by Prime (1853). Museum
records (all prior to 1950) and localized reports (Verrill, 1902; Vanatta, 1911)
show this species limited to the vicinity of the city of Hamilton, the likely point of
its introduction. The current survey finds O. draparnaudi common throughout the
western two thirds of Bermuda. A native of Europe, western Asia and northern
Africa, this species has been introduced to cities throughout North America.
Oxychilus draparnaudi is often found in gardens and vacant lots.

Poecilozonites bermudensis (Pfeiffer, 1845)
(Map 35, Fig. 36)

Helix Bermudensis Pfeiffer, 1845: 67 (Bermuda).

yl NEMOURIA No. 44

i
Yy
Y

Figure 36. Poecilozonites bermudensis. - FMNH 224886, Southampton Parish,
Bermuda; diameter = 17.1 mm (apical view); diameter = 15.4 mm (basal and
apertural views).

? Helix ochroleuca, - Pfeiffer, 1847: 80-81; Reeve, 1854: Helix pl. 177, no.
1212; Bland, 1861: 351;

Helix (Trochomorpha) bermudensis, - Albers, 1850: 116;

? Helix (Trochomorpha) ochroleuca Férussac, - Albers, 1850: 116 [non Helix
(Helicostyla) ochroleuca Férussac, 1821: 46 (“Les Grandes Indes’?,”
subsequently changed to “Habite l’ile Bermude (Menke)” by Férussac [in
Férussac & Deshayes], 1819-1851: 183); H. ochroleuca Férussac was later
interpreted as a senior synonym of Pachystyla rufozonata H. Adams, from
Mauritius, by Pilsbry 1900: 499);

Helix bermudensis, - Prime, 1853: 55; Reeve, 1851: Helix pl. 57, no. 275;
Jones, 1859: 106; Bland, 1861: 351; Tristram, 1861: 405; Rein, 1867:
77; Bland & Binney, 1873: 221; Mérch, 1878: 3; Bartram, 1878: 108;
Kobelt, 1880: 258; E. A. Smith, 1884: 277; Martens, 1889: 201; Lane,
1891: 1;

December 30, 2000 Bieler & Slapcinsky 58

Map 35. Poecilozonites bermudensis.

2? Nanina (Trochomorpha) ochroleuca, - Martens in Albers, 1860: 60-61
(expressing doubt about the presence of this species on Bermuda because
it was not listed by Jones in 1859),

Helix (Caracolus) bermudensis, - Martens in Albers, 1860: 156;

Hyalina bermudensis, - Bland, 1875: 77; Jones, 1876: 138; Bland, 1881: 256;

Hyalina ochroleuca, - Jones, 1876: 138;

Zonites (’) bermudensis, - Binney, 1876: 183; Binney, 1884: 86, 87;

? Hyalina ochrolenca [sic], - Jones, 1876: 138;

Hyalina [sic] bermudensis, - Jones, 1876: 138; Bartram, 1878: 108;

Hyalinia ? Bermudensis Clessin in Pfeiffer, 1879: 69;

Hyalinia bermudensis, - Kobelt, 1880: 286;

? Carocolla ochroleuca, - Kobelt, 1880: 286;

Zonites bermudensis, - E. A. Smith, 1884: 31;

Poecilozonites bermudensis, - O. Boettger, 1884: 139 [introduction of genus-
group name Poecilozonites, 2 species originally included]; Pilsbry, 1888:
289-290, pl. 17, fig. E, C; Heilprin, 1889: 182; Pilsbry, 1889: pl. 17, figs.
N, O; Pilsbry, 1889: 85; Bland, 1895: 270; Pilsbry, 1900: 499; Verrill,
1902: 782; Gulick, 1904: 418; Verrill, 1906: 159 ff.; Vanatta, 1911: 665-
668; Vanatta, 1923: 33; Vanatta, 1924: 7; Peile, 1926: 89; Kutchka,
1934: 296; Gould, 1969: 507, pl. 3, figs. 6-7 (Recent), pl. 4, fig. 6 (fossil),
fig. 7 (Recent); Gould, 1970b: 502 ff. (fossil); Gould, 1971a: 88 (fossil);
Abbott, 1989: 126, fig; Gould, 1991: 12;

Helix (group Patula, section Trochomorpha, subsection Poecilozonites)
bermudensis, - Tryon, 1887: 95, 96, pl. 19 figs. 7-10 [H. bermudensis
indicated as type species of Poecilozonites Sandberger [sic, error for O.
Boettger], 1884);

Bermudia bermudensis, - Ancey, 1887: 53 [introduction of genus-group name
Bermudia, with H. bermudensis as type species by monotypy];

Juno bermudensis, - Mazyck, 1889: 210-211 [introduction of genus-group name
Juno, with H. bermudensis as type species by original designation];

54 NEMOURIA No. 44

Poecilozonites bermudensis bermudensis, - Gould, 1966: 1135; Gould, 1968: 81
ff., pl. 10, fig. 6 (fossil), fig. 7 (Recent); Hughes, post 1968: 1; Hughes,
post 1969: 1; Eldredge & Gould, 1972: 99 ff.;

Poecilozonites (Poecilozonites) bermudensis, - Thiele, 1931: 595; Zilch, 1959: 263,
fig. 933; Riedel, 1980: 23.

Bermuda Material Studied: BAMZ 1994-118-009, 1998-166-025, 1998-166-053
(fossil); 1998-166-055, 1998-166-060; DMNH 210287, 210304, 210327 (fossil);
FMNH 3633, 8833, 8858, 13482, 13492, 13502, 13506, 13507, 13524, 13532,
13580, 16455, 16456 (fossil), 29787 (fossil), 29788 (fossil), 29799, 45243 (fossil),
45244, 45245, 45246, 71536, 82048, 93871, 100125, 100184, 103063, 106932,
115118, 127348, 127430 (fossil), 147304, 216369, 224859, 224886, 224910
(fossil), 227059.

Note: Endemic - The closest relatives of the genus Poecilozonites are the members of
the southeastern United States genera Gastrodonta and Ventridens (Pilsbry, 1900).
Once extremely common on Bermuda, Poecilozonites bermudensis specimens were
collected by the bucket-full and destroyed by burning in the middle part of this
century (Gould, 1969). Jones (1864: 8) referred to this species as “Quite as
common as the Garden Snail (H. aspersa) is in England.” After the introduction of
several predatory snails, populations of Poecilozonites bermudensis rapidly declined
(Gould, 1969, 1991) and recently this species was feared extinct (Gould, 1991). In
1988, this survey located fresh-dead specimens of this species (FMNH 224886) at
one locale, giving hope to future efforts to protect this species.

This is the type species of Poecilozonites O. Boettger (1884: 139) and its two
objective synonyms Bermudia Ancey (1887: 53), and Juno Mazyck (1889: 211).

Poecilozonites circumfirmatus (Redfield, 1853)
(Map 36, Fig. 37)

Helix circumfirmata Redfield, 1853: 16, 17 (Bermuda).

Helix ptychoides Prime, 1852: 55 [nomen nudum]; Jones, 1859: 106 [nomen
nudum; based on Prime’s list]; Bartram, 1878: 108 [nomen nudum;
erroneously referring to “Rein”;

Helix circumfirmata, - Shuttleworth, 1853: 123; Bland, 1861: 351; Pfeiffer,
1863: 2; Rein, 1867: 77; Bland & Binney, 1873: 221; E. A. Smith, 1884:
277; Morch, 1878: 3; Bartram, 1878: 108; E. A. Smith, 1884: 31;

Helix circumfirmatus, - Tristram, 1861: 405;

Helix discrepans Pfeiffer, 1863: 1-2 (Bermuda); Rein, 1867: 77;

December 30, 2000 Bieler & Slapcinsky 55

Figure 37. Poecilozonites circumfirmatus. - FMNH 29786, St. George’s Parish,
Bermuda; diameter = 8.6 mm (apical and basal views); diameter = 7.8 mm
(apertural view).

Hyalina discrepans, - Pfeiffer, 1868: 216; Jones, 1876: 138; Bland, 1881: 256;
Hyalina circumfirmata, - Jones, 1876: 138; Bland, 1881: 256;

Hyalosagda discrepans, - Kobelt, 1880: 286;

Hyalosagda circumfirmata, - Kobelt, 1880: 286;

Helix (group Sagda, section Hyalosagda) circumfirmata, - Tryon, 1887: 9, pl. 1,

Map 36. Poecilozonites circumfirmatus.

56 NEMOURIA No. 44

fig. 30;

Helix (group Sagda, section Hyalosagda) discrepans, - Tryon, 1887: 9;

Poecilozonites circumfirmatus variety Pilsbry, 1888: 291, pl. 17, fig. 11;

Poecilozonites circumfirmatus, - Pilsbry, 1888: 291, pl. 17, figs. A, B, F;
Heilprin, 1889: 182; Pilsbry, 1900: 500, pl. 62, fig. 9, 9a; Verrill, 1902:
728, figs. 67a, b; Gulick, 1904: 420; Verrill, 1906: 166, figs. 48a, b;
Vanatta, 1911: 665-668; Peile, 1924: 18-19; Vanatta, 1924: 7; Gould,
1966: 1135; Hughes, post 1968: 1; Hughes, post 1969: 1; Gould, 1966:
1135; Gould, 197 1a: 88 (fossil);

Helix (Hyalosagda) discrepans, - Heilprin, 1889: 182;

Poecilozonites circumfirmatus var. corneus Pilsbry, 1890a: 95;

? Hyalnia [sic] ---, - Lane, 1891: 3;

Poecilozonites circumfirmatas [sic], - Bland, 1895: 270;

Poecilozonites discrepans, - Bland, 1895: 270; Peile, 1925: 257; Peile, 1926: 89;

Poecilozonites (Gastrelasmus) circumfirmatus, - Pilsbry, 1924: 2 [introduction of
genus-group name Gastrelasmus with H. circumfirmata as type species by
original designation];

Poecilozonites (Gastrelasmus) circumfirmatus discrepans, — Pilsbry, 1924: 3, fig. 2,
a, b;

Poecilozonites circumfirmatus var. caliban Peile, 1924: 19;

Poecilozonites circumfirmatus mut. corneus, - Pilsbry, 1924: 2, fig. 1;

Poecilozonites peilei Vanatta, 1925: 423, figs. 1, 2;

Poecilozonites circumfirmatus var. corneus, - Peile, 1925: 257; Peile, 1926:89;

Poecilozonites circumfirmatus var. caliban, - Peile, 1926: 89;

Poecilozonites circumfirmatus peilei, - Peile, 1926: 89;

Poecilozonites (Gastrelasmus) circumfirmatus, - Peile, 1926: 89; Thiele, 1931:
595; Zilch, 1959: 263, fig. 934;

Poecilozonites (Poecilozonites?) peilei, - Riedel, 1980: 23;

Poecilozonites (Gastrelasmus) circumfirmatus, - Riedel, 1980: 23.

Bermuda Material Studied: BAMZ 1994-118-008, 1998-166-012, 1998-166-032,
1998-166-056; CAS 54685; DMNH 210281, 210303, 210318; FMNH 11429,
13489, 13494, 13512, 13553, 13565, 13581, 16457, 16458, 29803 (fossil),
41763, 45247 (fossil), 45248, 45249 (fossil), 70863, 71537, 92868, 93813,
100080, 100186, 102848, 114991, 129496, 147327, 217552, 224786, 224813,
224825, 224838, 224850, 224887.

Note: Endemic - Once very common and widespread on the larger islands of
Bermuda, this is the only member of the genus to be found alive in this latest
survey. Most records appear to be associated with leaf litter under herbaceous
cover. Some individuals have weak or no lamellae, leading to the varietal name

December 30, 2000 Bieler & Slapcinsky 57

corneus. The nominal species Helix discrepans was introduced by Pfeiffer (1863) for
a depressed specimen collected by Rein. Gould (1991: 12) was probably referring
to this species when he stated he found the “smallest and most cryptic” species of
Poecilozonites alive in 1990.

This is the type species of Gastrelasmus Pilsbry (1924: 2).

Poecilozonites reinianus (Pfeiffer, 1863)
(Map 37, Fig. 38)

Helix reiniana Pfeiffer, 1863: 1 (Bermuda).

Helix Reinii [sic] Pfeiffer, - Rein, 1867: 77;

Helix Reiniana, - Pfeiffer, 1868: 172;

Hyalina Reiniana, - Jones, 1876: 138;

Patula reiniana, - Kobelt, 1880: 258, 286; Bland, 1881: 256;

Helix (group Patula, section Discus) Reiniana, - Tryon, 1887: 27;

Patula (Discus) reiniana, - Clessin in Pfeiffer, 1879: 91;

Poecilozonites reinianus var. goodei Pilsbry, 1889: 85, pl. 3, figs. 12, 13;

7 Rotundatus, - Lane, 1891: 3 [non Helix rotundatus Miller, 1774);

Poecilozonites goodei, - Vanatta, 1911: 665; Peile, 1924: 18;

Poecilozonites reinianus, - Pilsbry, 1888: 290-291, pl. 17, fig I; Heilprin, 1889:
182; Bland, 1895: 270; Pilsbry, 1900: 500; Verrill, 1902: 728; Verrill,
1906: 164-165; Vanatta, 1911: 666; Vanatta, 1924: 7; Kutchka, 1934:
296; Hughes, post 1968: 1; Hughes, post 1969: 1;

Poecilozonites (Discozonites) reinianus, - Peile, 1924: 4, fig. 3 a-d; Peile, 1925:
258; Peile, 1926: 90; Riedel, 1980: 23;

Poecilozonites blandi Pilsbry, 1924: 2, 7-8, fig. 5 [introduction of genus-group
name Discozonites with P. blandi as type species by original designation]
(Bermuda), new synonymy;

Map 37. Poecilozonites reinianus.

58 NEMOURIA No. 44

I

Figure 38. Poecilozonites reinianus. - 38a: FMNH 115598, Bermuda; diameter =
10.0 mm (apical view); diameter = 10.2 mm (basal view); diameter = 9.4 mm
(apertural view). Poecilozonites reinianus “form blandi” - 38b: FMNH 45251,

Bermuda; diameter = 8.9 mm.

December 30, 2000 Bieler & Slapcinsky 59

Poecilozonites (Discozonites) blandi, - Peile, 1925: 258; Peile, 1926: 90; Thiele,
1931: 595; Zilch, 1959: 263, fig. 935 [after Pilsbry, 1924]; Riedel, 1980:
233

Bermuda Material Studied: FMNH 16454, 45238, 45250 (fossil), 45251, 45252
(fossil), 45253 (fossil), 45254 (fossil), 71538, 115598.

Note: Endemic - Historically much less common and more restricted in range than
Poecilozonites bermudensis and P. circumfirmatus, P. reinianus has only been collected
on the eastern side of the main island of Bermuda. The varietal name goodei was
introduced by Pilsbry (1889) for widely umbilicate and flattened shells that he
wished to distinguish from narrowly umbilicate forms identified as reinianus in
ANSP’s collections. He mistakenly believed his narrowly umbilicate forms to be
typical reinianus (Pilsbry, 1924). Pilsbry (1924) then synonymized goodei with
reinianus and described Poecilozonites blandi for the narrowly umbilicate forms
previously identified as reinianus. Umbilical width and the spire height of P.
reinianus are surprisingly variable, as variable as in species of Polygyra. Separation
of these two forms based on spire height and umbilical width alone is not possible
as the full range of morphologies from tall narrowly umbilicate “P. blandi” to
widely umbilicate “P. reinianus” and “P. goodei” exist in museum collections
(FMNH 16454, 45238, 45251, 71538 and 115598). Poecilozonites blandi, the type
species of Discozonites Pilsbry, 1924, is here synonymized with Poecilozonites
reinianus.

This species was last reported alive by Peile (1926). Gould was unable to
locate P. reinianus in 1973 (Gould, 1991: 12), as was this survey in 1988.

Striatura meridionalis (Pilsbry & Ferriss, 1906)
(Map 38, Fig. 39)

Vitrea (Striatura) milium meridionalis Pilsbry & Ferriss, 1906: 152-153 (Texas,
New Mexico, Arizona).

Striatura milium meridionalis, - Vanatta, 1912: 12 (fossil);
Striatura milium viar]. meridionalis, - Peile, 1926: 89 (fossil);
Striatura meridionalis, - Pilsbry, 1946: 493-495 (fossil);
species A, - Gould, 1970b: 491,

Striatura (Pseudohyalina?) meridionalis, - Riedel, 1980: 23.

Bermuda Material Studied: ANSP 105793; FMNH 227041; MCZ Gould Station
4 (fossil), Gould Station 48 (fossil), Gould Station 63 (fossil).

60 NEMOURIA No. 44

Figure 39. Striatura meridionalis. - ANSP 105793, Hamilton Parish, Bermuda;

diameter = 1.68 mm.

Note: Native Non-Endemic - The paucity of live records of this species on Bermuda
reflects its small size. Vanatta (1912) was the first to recognize this species from
Bermuda. Striatura meridionalis is found in the southeastern United States
(Maryland to Texas), in Mexico, and the Greater Antilles (van der Schalie, 1948).

It lives in moist leaf litter.

Map 38. Striatura meridionalis.

December 30, 2000 Bieler & Slapcinsky 61

Zonitoides arboreus (Say, 1816)
(Map 39, Fig. 40)

Helix arboreus [Say, 1816]: species no. 2, pl. 4, fig. 4 (North America).

Zonitoides arboreus, - Bryant, 1905: 130;

Zonitoides bermudensis Pilsbry @& Vanatta, 1909: 63-64, fig. 1 ad, new
synonymy;

Zonitoides bermudensis, - Vanatta, 1924: 7; Vanatta, 1911: 665-667; Peile,
1926: 89.

Bermuda Material Studied: ANSP 91152, 100760, 99100, 131601, 85500,
110690, 113129, 85562; BAMZ 1998-166-081, 1998-166-114; DMNH 210277,
210258; FMNH 45024, 227050, 227051, 227052, 227053, 227054.

Note: Accidentally Introduced - The first record of Zonitoides on Bermuda came from
a single lot collected by Stewerdson Brown in 1905. These specimens were used
by Pilsbry & Vanatta (1909) to describe Zonitoides bermudensis, which is here
synonymized with Zonitoides arboreus (Say, 1816).

In their description, Pilsbry & Vanatta cite the wide umbilicus of Zonitoides
bermudensis as the main feature distinguishing it from Z. arboreus. However, Pilsbry
(1946: 481) later demonstrated the variability of this character in Zonitoides
arboreus, stating the “umbilicus [is] contained about 4 ”% to 5 times in the
diameter.” On the same page, measurements are given of a specimen from
Williams County, Colorado, which is even more widely umbilicate (the umbilicus
width fitting only four times in the shell diameter). Examination of the holotype
and paratypes of Zonitoides bermudensis (ANSP 91152) collected by Brown reveals
the holotype to be quite widely umbilicate. In fact, the holotype is among the
most widely umbilicate of the hundreds of specimens collected by Brown, having
a ratio of umbilical width to shell diameter of 0.338. A random sample of 50 of

Map 39. Zonitoides arboreus.

62 NEMOURIA No. 44

Figure 40. Zonitoides arboreus. - 40a: ANSP 91152, Holotype of Zonitoides
bermudensis Pilsbry & Vanatta; diameter = 5.7 mm (body whorl damaged). - 40b:
ANSP 91152, one of numerous paratypes of Zonitoides bermudensis Pilsbry &

Vanatta; diameter = 4.6 mm.

December 30, 2000 Bieler @ Slapcinsky 63

the other specimens collected by Brown average only 0.265 umbilical
width/diameter and can not be differentiated from widely umbilicate populations
of Zonitoides arboreus from Missouri (FMNH 250319), New York (FMNH 250356),
Tennessee (FMNH 250607, 250621), and Texas (FMNH 250638, 250640). Most
of these widely umbilicate populations appear to occur in highly calcareous areas.
The most widely umbilicate lot (FMNH 250621), like the lot collected by Brown
from Bermuda, came from the vicinity of a cave. Other specimens collected from
Bermuda (ANSP 85500, 110690, 113129; FMNH 227050, 227051, 227052,
227053, 227054) have the more “typical” morphology of Zonitoides arboreus.

Zonitoides arboreus is a native of North America and is found over most of the
continent. It is also well known as an adventive species and has been recorded
from South Africa, Hawaii, Japan and Australia (Pilsbry, 1946) and can often be
found in potted plants. Zonitoides species are not found in Bermuda’s abundant
Pleistocene deposits. Z. arboreus is found under almost any type of cover and often
is associated with rotting wood.

POLYGYROIDEA
POLYGYRIDAE
Patera appressa (Say, 1821)
(Map 40, Fig. 41)

Helix appressa Say, 1821: 151 (“banks of the Ohio and Missouri”; restricted to
Ohio, Pilsbry, 1940: 751).

Helix Sancta Georgiensis Prime, 1853: 55 [nomen nudum];

Helix Sancta Georgiensis Jones, 1859: 107 [nomen nudum; based on Prime’s list,
but erroneously indicated as “n.s.”);

Helix appressa, - Jones, 1876: 138; Bland, 1881: 256; Heilprin, 1889: 182 ;
Lane, 1891: 2; Bland, 1895: 270;

Hex [sic] appressa, - Bartram, 1878: 108;

Triodopsis appressa, - Aldrich, 1889: 9;

Helix Sancta Georgiensis ‘Bartram’ - Lane, 1891: 2 [introduced in synonymy of
Helix appressa];

Polygyra appressa, - Pilsbry, 1900: 496; Verrill, 1902: 730, 732, fig. 75d-f [with
H. sancta-georgiensis in synonymy]; Pilsbry & Vanatta, 1924: 7;

Polygyra appressa sculptior Chadwick, 1899 [originally described as variety from
Virginia], - Pilsbry, 1900: 496; Vanatta, 1911: 665; Peile, 1926;

Polygyra appressa var. sanctaegeorgiensis “(Prime), P. & V.”, - Peile, 1926: 90;

Polygyra appressa var. sculptior, - Peile, 1926: 90;

Mesodon (Patera) appressus f. sanctigeorgiensis [sic] - Pilsbry, 1940: 751, fig. 452c;

64 NEMOURIA No. 44

Figure 41. Patera appressa. - FMNH 29789, St. George’s Parish, Bermuda;

diameter = 13.3 (apical view); diameter = 13.0 mm (basal and apertural views).

Mesodon (Patera) appressus form sanctageorgiensis ”’Prime’ Verrill”, - Pilsbry,

1940:.752.

Bermuda Material Studied: BAMZ 1998-166-001, 1998-166-008, 1998-166-016,
1998-166029, 1998-166042, 1998-166-047, 1998-166-115; DMNH 210241,
210250, 210280, 210312, 210320; FMNH 3639, 13484, 13525, 13533, 13539,
13583, 13592, 16460, 16461, 45850, 100191, 124505, 157504, 216047, 224768,

Map 40. Patera appressa.

December 30, 2000 Bieler & Slapcinsky 65

224788, 224824, 224837, 224849, 224858, 224885, 224899.

Note: Accidentally Introduced - Native to the southeastern United States, from
eastern Tennessee to eastern Virginia, this species was introduced to Bermuda
before the first faunal survey of the islands. For many years it was restricted to the
area around St. George, but it is now widespread on the larger islands of
Bermuda. Patera appressa is often found in leaf litter in rocky areas and under
cover in urban areas.

Polygyra plana (Dunker, 1843)
(Map 41, Fig. 42)

Helix plana Dunker in Philippi, 1843: 51-52, pl. 3, fig. 11 (“Indiae
occidentales?”).

Helix microdonta Deshayes [in Férussac G Deshayes], 1839, - Redfield, 1853:
17; Jones, 1859: 107; Bland, 1860: 93, 94; Tristram, 1861: 405; Bland,
1861: table 2, p. 351; Jones, 1864: 9; Bland, 1866: 360; Rein, 1867: 77;
Jones, 1876: 138; Morch, 1878: 3; Bland, 1881: 256; E. A. Smith, 1884:
277; Bland, 1895: 270 [non Helix microdonta Deshayes, 1832, teste
Vanatta, 1911: 664};

Helix cheilodon “Say MS.” Bland, 1860: 94 [introduced as synonym of H.
microdonta sensu Bland, from Bermuda);

Polygyra microdonta, - Kobelt, 1880: 258, 286; Pilsbry, 1900: 496, pl. 62, fig. 3;
Verrill, 1902: 729, fig. 72; Gulick, 1904: 413; Hughes, post 1968: 1;
Hughes, post 1969: 1;

Helix Ringens Lane 1891: 2, pl. 1, fig. 3 [plate not seen] (Bermuda) [non Helix
ringens Linné, 1758: 769);

Helix (Polygyra) microdonta, - Heilprin, 1889: 182;

Polygyra plana, - Vanatta, 1911: 665-668 [citing Thiele in litt.; with H.
microdonta Deshayes, 1839 in synonymy]; Honigmann, 1914: 375;

Map 41. Polygyra plana.

66 NEMOURIA No. 44

Figure 42. Polygyra plana. - FMNH 224888, Southampton Parish, Bermuda;
diameter = 8.2 mm (apical and apertural views); diameter = 8.2 mm (basal view;

different specimen).

Vanatta, 1923: 33; Peile, 1926: 90; Abbott, 1989: 134, fig.

Bermuda Material Studied: BAMZ 1998-166-003, 1998-166-010, 1998-166-027,
1998-166-074, 1998-166096, 1998-166-100; DMNH 210262, 210272, 210284,
210306, 210314, 210322; FMNH 3640, 8834, 13517, 13535, 13545, 13551,
13564, 13573, 13593, 16459, 41023, 45687, 45778, 100168, 115647, 125209,
153255, 224781, 224800, 224801, 224851, 224867, 224873, 224888, 224894.

Note: Accidentally Introduced - A native of the Bahamas, this species arrived before
the first faunal survey and is one of the few introduced species commonly found
on the smaller island of Bermuda. It is very tolerant of salt spray and is found on

dunes and in other open areas under cover.

December 30, 2000 Bieler & Slapcinsky 67

Figure 43. Lacteoluna selenina. - FMANH 224881, Southampton Parish, Bermuda;

diameter = 5.2 mm (apical and apertural views); diameter = 4.8 mm (basal view).

SAGDIDAE
Lacteoluna selenina (Gould, 1848)
(Map 42, Fig. 43)

Helix selenina Gould, 1848: 38 (Florida).

Helix selenina, - Prime, 1853: 55; Jones, 1859: 106;

Helix vortex Pfeiffer, 1839: 351 [non Helix vortex Linné, 1758] (Cuba);

Helix vortex, — Bland, 1861: table 2, p. 351; Rein, 1867: 77; Jones, 1876: 138;
Bland, 1881: 256; E. A. Smith, 1884: 227;

Microphysa vortex, - Kobelt, 1880: 257, 286;

Helix (group Patula, section Microphysa) vortex, - Tryon, 1887: 98, pl. 19, figs.
25-28;

Helix (Microphysa) vortex, - Heilprin, 1889: 182;

68 NEMOURIA No. 44

Map 42. Lacteoluna selenina.

Patula vortex, - Bartram, 1878: 108; Bland, 1895: 270;

Thysanophora vortex, - Pilsbry, 1900: 495, pl. 62, fig. la, 1b; Verrill, 1902:
728, 229, fig. 70a, b; Gulick, 1904: 413; Peile, 1926: 90; Hughes, post
1969: 1;

Thysanophora selenina, - Vanatta, 1911: 665-668; Vanatta, 1923: 33;

Lacteoluna selenina, - Pilsbry, 1940: 981, 982.

Bermuda Material Studied: BAMZ 1998-166-017, 1998-166-022, 1998-166-043,
1998-166-084, 1998-166-111, 1998-166-122; DMNH 210279, 210329, 210332,
210334; FMNH 3638, 7607, 13486, 13515, 13529, 13540, 13561, 29784, 30198,
30199, 41017, 41018, 45560, 59080, 107033, 224765, 224784, 224807, 224808,
224821, 224834, 224842, 224845, 224856, 224881, 227022, 227031.

Note: Accidentally Introduced - One of the earliest introductions, Lacteoluna selenina
was recorded by Prime (1853) as collected in 1852, and was reported from most
surveys since, partly under its synonym Helix vortex, a preoccupied name. Hughes
(post 1969; citing Gulick, 1904, and Gould, 1969) erroneously included this
species in a listing of fossil species of Bermuda. This calciphile native of southern
Florida and the West Indies is commonly found under stones and debris in
woodlands.

THYSANOPHORIDAE
"Hojeda" hypolepta (Shuttleworth, 1854)
(Map 43, Fig. 44)

Helix hypolepta Shuttleworth, 1854: 129 (Bermuda).
Helix hypolepta, - Jones, 1876: 138;

?Helix hypolepta, - Heilprin, 1889: 182;
Patula hypolepta, - Bland, 1881: 256;

December 30, 2000 Bieler & Slapcinsky 69

Figure 44. "Hojeda" hypolepta. - ANSP 100756, Paget Parish, Bermuda; diameter =

2.4 mm.

Helix (Microphysa) hypolepta, - Pilsbry, 1889: 82, pl. 3, figs. 6-8;

Patula (Thysanophora) hypolepta, - Bland, 1895: 270;

Lacteoluna hypolepta, - Haas, 1952: 104;

Thysanophora hypolepta, - Pilsbry, 1900: 496, pl. 62, fig. 2a, 2b; Verrill, 1902:
728, fig. 68a, b; Gulick, 1904: 413; Verrill, 1906: 169, fig. 52a; Vanatta,
1911: 666-667; Vanatta, 1912: 12: Vanatta, 1923: 33; Peile, 1926: 90;
Hughes, post 1968: 1; Hughes, post 1969: 1; Gould, 1970b: 491, fig. la
(fossil); Gould, 1970a: 572 (fossil); Gould, 197 1a: 88;

Map 43. "Hojeda" hypolepta.

70 NEMOURIA No. 44

Hojeda hypolepta, - Richardson, 1986: 6.

Bermuda Material Studied: ANSP 100756; BAMZ 1995-132021; FMNH
227021; MCZ Gould Station 1 (fossil), Gould Station 4 (fossil), Gould Station 11
(fossil), Gould Station 21 (fossil), Gould Station 21b (fossil), Gould Station 30
(fossil), Gould Station 41 east (fossil), Gould Station 48 (fossil), Gould Station 63
lower (fossil), Gould Station 73 (fossil).

Note: Endemic - The young of this species are very similar to Hawaiia minuscula
and the two species were often confused in early surveys (Pilsbry, 1900). The
anatomy of this species is unknown and shell characters cannot place it in any
known genus. Its traditional familial placement within the Sagdidae or
Thysanophoridae is also suspect. Based on shell characters alone this species
might be equally well placed within the Helicodiscidae or Zonitidae. Pending
anatomical studies, it is here provisionally placed in Hojeda Baker, 1926, following
Richardson (1986). This species is found under rocks in open areas.

HELICOIDEA
BRADYBAENIDAE
Bradybaena similaris (Rang, 1831)
(Map 44, Fig. 45)

Helix similaris “Férussac” Rang, 1831: 15 (numerous localities).

Helix (Helicella) similaris Férussac, 1821: 43 [or p. 47, depending on issue]
[nomen nudum];

Helix (Fruticicola) similaris, - Aldrich, 1889: 9;

Helix hortensis, - Lane, 1891: 2 [non Helix hortensis Muller, 1774: 52];

Eulota simularis [sic], - Olmsted, 1917: 225;

Eulota similaris, - Pilsbry, 1900: 495; Verrill, 1902: 734, fig. 82a-c; Vanatta,
1911: 665-668; Vanatta, 1923: 33;

Eulota (Eulota) similaris, - Honigmann, 1914: 376;

Eulota similis [sic], - Peile, 1926: 90.

Bermuda Material Studied: BAMZ 1998-166-009, 1998-166-030, 1998-166-048,
1998-166-050, 1998-166-059, 1998-166-070, 1998-166-075, 1998-166-093, 1998-
166-119, 1998-166-121, 1998-166-124; DMNH 210243, 210244, 210248,
210253, 210291, 210296, 210299, 210321, 210325; FMNH 3635, 13488, 13491,
13505, 13514, 13522, 13534, 13546, 13562, 13590, 13678, 13685, 13688,
16468, 16469, 16470, 16471, 29792, 56551, 66154, 92867, 147322, 217509,

December 30, 2000 Bieler & Slapcinsky Al

Figure 45. Bradybaena similaris. - FMNH 13579, Smith’s Parish, Bermuda;
diameter = 14.8 mm (apical view); diameter = 15.4 mm (basal and apertural
views).

224158, 224769, 224782, 224802, 224818, 224829, 224877, 224895, 224904,
224915, 224916, 224917.

Note: Accidentally Introduced - First reported in 1889 by Aldrich, this species is
one of the most common snails on Bermuda. Native to southeastern Asia, it is

Map 44. Bradybaena similaris.

iZ NEMOURIA No. 44

widely introduced to subtropical and tropical regions including the southeastern

United States (Dundee, 1970), the Caribbean, and southern Brazil.

COCHLICELLIDAE
Prietocella barbara (Linné, 1758)
(Map 45, Fig. 46)

Helix barbara Linné, 1758: 773 (North Africa).

Bulimus ventrosus (Férussac, 1821), -
Prime, 1853: 55; Redfield,
1853: 17; Jones, 1859: 107;
Bland, 1861: table 2; Jones,
1864: 9; Rein, 1867: 77; Jones,
1876: 139; Moérch, 1878: 3; E.
A. Smith, 1884: 277; Frith,
1891: 15;

Bulimus ventricosus Draparnaud,
1801, - Tristram, 1861: 405
[non Bulimus ventricosus Bru-
guiére, 1792];

Bullimus [sic] ventrosus, - Bartram,
1878: 108;

Helix ventricosa, - Jones, 1876: 138;
Bland, 1881: 256; Bland, 1895:

271;
Cochlicella ventrosa, - Kobelt, 1880: Figure 46. Prietocella barbara. - FMNH
257, 286; 16472, St. George’s Parish, Bermuda;
Helix (Cochlicella) ventricosa, - height = 6.6 mm.

Heilprin, 1889: 182;

Map 45. Prietocella barbara.

December 30, 2000 Bieler & Slapcinsky 73

Helicella (Cochlicella) ventricosa, - Pilsbry, 1900: 494;

Helicella ventricosa, - Verrill, 1902: 732, 733, fig. 79a-d; Vanatta, 1911: 665-
668; Vanatta, 1923: 33;

Cochlicella barbara, - Glaubrecht, 1990: 218;

Prietocella barbara, - Schileyko & Menkhorst, 1997: 54 [for placement in
Prietocella).

Bermuda Material Studied: BAMZ 1998-166-033, 1998-166036, 1998-166044,
1998-166-079, 1998-166095, 1998-166-097, 1998-166-101; DMNH 210245,
210266, 210301, 210307, 210317; FMNH 3636, 3637, 4034, 13487, 13498,
13508, 13513, 13523, 13536, 13543, 13554, 13566, 13570, 13682, 16472,
29789, 41021, 102461, 124907, 125197, 217430, 224756, 224765, 224779,
224797, 224819, 224830, 224871, 224878, 224918, 224919, 227058.

Note: Accidentally Introduced - Recorded in the first survey of Bermuda, and in
nearly every other survey, this snail is very common on Bermuda. Prietocella
barbara is native to the Mediterranean region and has been introduced into North
America to California and the Carolinas. It is often found in large numbers on
bushes and on bark at the base of trees.

HELICIDAE
Otala Iactea (Miller, 1774)
(Map 46, Fig. 47)

Helix lactea Miller, 1774: 19-20 (Europe).

Otala lactea, - Bennett & Hughes, 1959: 432; Hughes, 1962; Hughes, 1963;
Simmonds & Hughes, 1963: 219-221; Hughes (post 1968): 1; Gould,
1969: 419; Monkman, 1982a: 1; Monkman, 1982b: 71; Bennett et al.,
1985: 138; Cock, 1985: 138; Abbott, 1989: 191, fig.; Glaubrecht, 1990:
220, fig. 10.

Bermuda Material Studied: BAMZ 1994-118-002, 1995-133-001, 1998-166-035,
1998-166054, 1998-166064, 1998-166069, 1998-166-106; DMNH 210246,
210289, 210292, 210295, 210315; FMNH 224763, 224775, 224791, 224836,
224867, 224883, 224912, 227002, 227003, 227004, 227005.

Note: Accidentally Introduced - This native of southern Spain and northern Africa is
a common food snail in markets around the world and has also been introduced

74 NEMOURIA No. 44

Figure 47. Otala lactea. - FMNH 224775, Paget Parish, Bermuda; diameter = 33.8

mm (apical view); diameter = 33.9 mm (apertural view).

to North America. In 1928, Otala lactea was introduced to Fairylands, Bermuda,
reportedly by crawling out of a bag purchased from the New York Fulton Fish
Market for food (Gould, 1969: 419, citing personal communication with
Hughes). By the 1950s, it was widespread and considered a horticultural pest by
the Bermuda Department of Agriculture. Otala lactea was the primary impetus for
the release of several predatory snails on Bermuda (Bennett & Hughes, 1959).
Initial very optimistic reports of successful control by the predatory species
(Simmonds & Hughes, 1963; Hughes, post 1968; mirrored by Godan, 1983: 325,
who called it a “great success”) were followed by surveys showing little or no
control of Otala populations (Monkman, 1982a). The current survey finds Otala
lactea to be one of the most common species of snails on Bermuda, easily found
aestivating in bushes throughout the larger islands of Bermuda.

Map 46. Otala lactea.

December 30, 2000 Bieler & Slapcinsky M5

SS

Figure 48. Theba pisana. - BAMZ 1995-133-005, St. George’s Parish, Bermuda;

diameter = 22.2 mm (apical view); diameter = 22.8 mm (apertural view).

Theba pisana (Miller, 1774)
(Map 47, Fig. 48)

Helix pisana Miller, 1774: 60-61 (Europe).

Helix pisana, - Davis, 1904: 126; Vanatta, 1911: 665;
Euparypha pisana, - Peile, 1926: 90.

Bermuda Material Studied: BAMZ 1995-133-005, 1998-166-063; DMNH
210293; FMNH 16467, 29790, 29791, 227018.

Note: Accidentally Introduced - This common Mediterranean species was first
recorded around the city of St. George by Davis (1904) who attributed his record
to the Verrill party of 1898. Theba pisana has also been introduced to Australia,
South Africa and western North America. On Bermuda it is one of the few
introduced species that have a restricted range. To date it has only been found

Map 47. Theba pisana.

76 NEMOURIA No. 44

within a few miles of the city of St. George on St. George's Island. Live individuals
can be found aestivating on the stems of bushes.

Figure 49. Leidyula sloanii. - FMNH 224876, Southampton Parish, Bermuda;
dorsal view; body length = 29 mm.

SYSTELLOMMATOPHORA: RATHOUISOIDEA
VERONICELLIDAE
Leidyula sloanii (Cuvier, 1816)
(Map 48, Fig. 49)

Onchidium Sloanii Cuvier, 1816: 411 (Jamaica).

Vaginulus sp., — Pilsbry 1888: 288;

Vaginulus schivelyae Pilsbry, 1890b: 297, pl. 5, figs. 68 (Bermuda);

Vaginulus schivelyae, - Pilsbry, 1891: 39, pl. 2, figs. 6-8;

Veronicella schivelyae, - Pilsbry, 1900: 502; Verrill, 1902: 728, 735, figs. 84, la-
c; Peile, 1926: 90; Hughes, post 1968: 1;

Veronicella schivelyoe [sic], - Olmsted, 1917: 224;

Veronicella sloanii schivelyae, - H. B. Baker, 1925: 160-163, pl. 3, figs. 14.

Bermuda Material Studied: BAMZ 1998-166-116; FMNH 3642, 13668, 30211,

Map 48. Leidyula sloanii.

December 30, 2000 Bieler & Slapcinsky ial

30212, 59481, 224760, 224868, 224876.

Note: Accidentally Introduced - This species was first recorded from specimens
collected “in the Public Garden at St. George” by the Heilprin party in 1888
(Pilsbry, 1888; 1890b: 298). It is commonly found at night, crawling on walls and
garden fences.

Bermuda specimens were described as Vaginulus schivelyae Pilsbry, 1890, and
thought endemic to Bermuda until compared to specimens of Jamaican Leidyula
sloanii by Baker (1925). Hoffmann (1927: 215) argued in contrast that, pending
further anatomical observations, Pilsbry’s V. schivelyae could be conspecific with
Leidyula floridana (Leidy, 1851), L. moreleti (Crosse & Fischer, 1872), or L. olivacea
Stearns, 1871. The group needs additional study and is here listed under the
earliest name of the complex.

Several singular or doubtful records exist:

Achatina fulica Bowdich, 1822: A snail collected in May of 1971 from Dock Hill,
Devonshire Parish, was identified by staff of the Department of Agriculture as
Achatina fulica from a photograph published in a local newspaper (Hughes, 1971).
However, the snail apparently was not preserved. Specimens of Achatina were not
located in collections at the Bermuda Department of Agriculture or in the
Natural History Museum, Bermuda, and no further specimens have been
reported.

Mesodon zaletus (Binney, 1837): A single series of three specimens of this eastern
North American species were found in the collections of the Natural History
Museum, Bermuda (BAMZ 1995-132-024). However, this large-shelled species was
not found during this survey and no locality information was associated with the
museum material.

Cerion spp.: Cerion maritimum (Pfeiffer, 1839) was listed by Prime (1853: 77, as
Pupa). Jones (1864: 9) referred to Bermuda material “found by Mr. Marrett on the
Port Royal Hills” as “Pupa chrysalis’” (apparently based on Cerion mumia var.
chrysalis Férussac, from Cuba, Guadeloupe and Martinique). One specimen of
Cerion variabile Dall, 1905, was collected by Louis Mowbry in 1926 from beach
drift and reported by Gray (1942). No other specimens of this genus, live or dead,
have been recorded, and no Cerion population seems to be established on the
islands.

Limax maximus Linné, 1758 - Recorded by Hughes (post 1968a), this large slug
does not appear in the collections of either the Bermuda Department of

78 NEMOURIA No. 44

Agriculture or the Natural History Museum, Bermuda. This record is probably a
misidentification of Limacus flavus.

Cepaea nemoralis (Linné, 1758) - The European Garden Snail was reported for
Bermuda by Lane (1891, Essay 1; as Helix nemoralis), but has not been found since.
Verrill (1902: 734) suggested this record to be an error for Poecilozonites
bermudensis, whereas Vanatta (1911: 665) and we interpreted it as an error for
Bradybaena similaris.

Succinea putris, Pupa troquilla pyrenaearia |sic], and Pachystyla mauritiana var.
were listed for Bermuda by Bartram (1878: 108). Bartram misidentified his
material by direct comparison with Chenu’s illustrations of European specimens
in the ‘Manuel Conchyliologie.’ Bartram was probably referring to Succinea

bermudensis, Gastrocopta pellucida, and Poecilozonites bermudensis.
Unavailable names mentioned in the Bermuda literature:

“ Helix somersetit’
Helix Somersetii Prime, 1853: 55 [nomen nudum];
“H[elix] Somersetii (n.s.)” Jones, 1859: 106-107 [based on Prime’s list; nomen

nudum].

“ Bulimus bermudensis”
Bulimus Bermudensis Prime, 1853: 55 [nomen nudum];
“Blulimus] Bermudensis (n.s.)” Jones, 1859: 107 [based on Prime’s list; nomen

nudum].

Neither name was validly introduced and neither can be unequivocally
assigned to any one species.

DISCUSSION

Bermuda’s fauna, with few terrestrial animals, has long been considered
depauperate and disharmonious (Verrill, 1902). However, the 17 species of
terrestrial mollusks native to Bermuda (see Table 1) are slightly more than might
be expected from an island group of such limited size (54 km? or 19.5 mi’) and
low elevation (75 m or 245 ft). Pacific islands of similar size and elevation, for
example, average only 12.5 species (Solem, 1990: table 5). As was pointed out
before, land snail diversity does not seem to be well predicted by island size and
counter-intuitively rises with distance from a colonization source (Solem, 1974).
The high levels of species diversity observed on isolated islands are largely due to

December 30, 2000 Bieler & Slapcinsky 79

extensive species radiations of rarely successful colonists, as can be seen in the
Achatinellidae of Hawaii or Poecilozonites of Bermuda. However, many of
Bermuda’s native land snails, 6 of 17, are native non-endemics and the
abundance of these is perplexing when Bermuda’s isolation and traditional views
of the low dispersal ability of snails are taken into consideration. Several plausible
scenarios have been proposed to explain such dispersal ability of snails, including
wind dispersal (Peake, 1981: 250), rafting (Pilsbry, 1900), and dispersal on
migratory birds (Dundee et al., 1967). Small snails would be more likely to be
transported by these means than larger snails. It is no surprise then, that Gould
(1970b: 490) doubted “that any other factor beyond size is required to explain
why the pulmonate fauna of Bermuda, an assemblage of chance immigrants,
includes such a preponderance of small forms.” However, size may not be the only
factor determining the unusual species composition of Bermuda’s land snail
fauna: Many of Bermuda’s native genera of land snails are known to contain
species having the ability to produce young uniparentally, for example Carychium
(Bulman, 1990) and Vertigo (Pokryszko, 1987), or have anatomy suggestive of self-
fertilization as in Poecilozonites (Pilsbry, 1889: 85).

All eleven species endemic to Bermuda share their closest affinities with
species of eastern North America (see notes under each species), with the possible
exception of “Hojeda” hypolepta whose relationship with other species is unknown.
Antillean affinities reported by Pilsbry (1900) are based in part on several species
now known to have been introduced accidentally. The six native non-endemic
species all have widespread distributions that are remarkably similar and include
the extreme southeast coast of North America, and in most cases the Greater
Antilles as well.

The majority of Bermuda’s terrestrial mollusks are very widely distributed, are
closely associated with human activities, and do not have a fossil history on
Bermuda, leading to the generalized conclusion they are accidentally introduced
through commerce. In fact, at least eleven of these species have also been
accidentally introduced to the Hawaiian Islands (Cowie, 1997) and 21 are known
traveling species often encountered by the Animal and Plant Health Inspection
Service of the United States Department of Agriculture (Robinson, 1999). The
high proportion of synanthropic mollusks on Bermuda (28 of 49 species) mirrors
the high proportions in some of Bermuda’s other terrestrial animal groups, such
as Coleoptera (“nearly all the beetles have been introduced accidentally by human
activity”; Hilburn @ Gordon, 1989: 674) and spiders (53 of 59 species; Sierwald,
1988). Of the 28 introduced snail species, nine or ten are native to Europe and
North Africa, ten are native to Tropical America, four or five are native to North
America, two to Asia, one to Africa, and one to the Pacific Ocean Basin. Because

80 NEMOURIA No. 44
Table 1. Terrestrial mollusk species occurrences on Bermuda by date of survey.

Key to Symbols:

L] Recent

@ Recent Live-Collected
@ Fossil

See notes in species listings for additional information about dates of introduction.

Endemic Species
Carychium bermudense

BHUL HR eee

“Euconulus” turbinatus
“Hojeda” hypolepta

Poecilozonites bermudensis

Poecilozonites circumfirmatus

el Cece:
OOOOSO

Poecilozonites reinianus

Punctum bristoli

i
BeHUO G@

Succinea bermudensis

See PSs ee

O
O
is
| Es
O
O
al
O
ro

Vertigo bermudensis
Vertigo marki

@©@e@0e008e008000 80

fleas
a

Vertigo numellata

Date of survey” fossil 1852 1859 1861 1867 1873 1878 1881 1888 1895 1898 1910 1922 1926 1950 1988

Native Non-Endemic Species
Gastrocopta pellucida @ -"O OO |= = 2
Gastrocopta rupicola @
Pupisoma macneilli ® - = - 2, (ees BSS
Pupoides albilabris ® - HOO - O8F 4 Gee Bae
Striatura meridionalis @

@

Strobilops hubbardi

Date of survey” fossil 1852 1859 1861 1867 1873 1878 1881 1888 1895 1898 1910 1922 1926 1950 1988

*Key to Survey Dates, Collectors and Sources:

Fossil collected by A. Gulick and S. J. Gould (Gulick, 1904; Gould 1970b)

1852 collected by T. Prime (Prime, 1853; Pilsbry 1900)

1859 (pre-1860) collected by T. Prime and J. M. Jones (Jones, 1859)

1861 (pre-1862) collected by Governor F. Murray (Tristram, 1861)

1867 (pre-1868) collected by J. J. Rein and Governor C. B. Ord’s sons (Rein, 1867)

1873 collected by the Challenger Expedition (Smith, 1884; Pilsbry, 1900)

1878 (pre-1879) collected by J. T. Bartram and an un-named gentleman from Hamilton
(Bartram, 1878)

December 30, 2000 Bieler & Slapcinsky
Table 1. (continued)

Accidentally Introduced Species
Opeas hannense

+Streptostele musaecola
Tornatellides sp.

Otala lactea 1928 -
Helicodiscus singleyanus
Allopeas gracile
Allopeas micrum

Theba pisana

Zonitoides arboreus

OM@OOdO

Gastrocopta servilis
Limacus flavus
Bradybaena similaris 1889 -
Gulella bicolor
Leidyula sloanii

CJ
OJ
Obeliscus swiftianus O
a

O
OO 8B

OBR BRB BRBOOBRBO:

Rumina decollata E}.- = is)
Deroceras laeve [Slip ee ve? “(Gl
Milax gagates LJ OO
Oxychilus draparnaudi

a
i

Subulina octona
Ceciliodes acicula

Say ESS) eee ee epee ee) ie

—
BERGE

cay [ea a a Ps
fel Cich eels

Polygyra plana
Prietocella barbara

OO

(= )) ly

a al
ferent

el evel

Hawaiia miniscula
Lacteoluna selenina

al ae
(eal Sh

Oligyra convexa
Patera appressa

Vallonia pulchella

Gyles
Cc
BOUDOOUSODO:
as |
Ci) Gy Osa a ea ae
i
BERHRHREHEUBRRB REE G

Oo.
Ol
LO

Date of survey” 1852 1859 1861 1867 1873 1878 1881 1888 1895 1898 1916 1922 1926 1950 1988

1881 (pre-1882) collected by C. B. Adams, T. Prime, R. Swift, T. Bland (Bland, 1881)
1888 collected by the Heilprin Expedition (Heilprin, 1889; Pilsbry 1900)

1895 (pre-1896) collected by C. B. Adams, T. Prime, R. Swift, T. Bland (Bland, 1895)
1898 collected by the Verrill Expedition (Pilsbry, 1900)

1910 (1905-1910) collected by S. Brown, E. E. and E. G Vanatta (Vanatta, 1911)
1922 collected by H. Hoyt (Vanatta, 1924; Vanatta, 1924)

1926 (pre-1927) collected by A. J. Peile G@ A. Haycock (Peile, 1926)

1950 collected by F. Haas (FMNH collections; Haas 1952)

1988 (1983 & 1988) collected by R. Bieler et al.

+An additional record from the Fahy Collection, 1992.

82 NEMOURIA No. 44

human activities have carried these species throughout the world, tracing a single
point-oforigin for Bermuda’s populations of each species (if only one exists) is
probably impossible. Three of these species are recorded here for the first time
(Streptostele musaecola, Opeas hannense and Tornatellides sp.). Such species
introductions, although nearly tripling the number of land snail species on
Bermuda, have not lead to saturation and subsequent species turnover. Solem’s
(1990: 112) assertion that, given time and appropriate conditions, land snail
species can accumulate sympatrically to a level of more than 60 species (postulated
for natural colonization of Pacific islands) might soon be proved true also for the
in part “artificial” colonization of Bermuda.

Unfortunately, the similarities between Bermuda and the Pacific Islands have
continued with the deliberate introduction of four predatory snail species for
biological control. The earliest reports of declines in abundance in Poecilozonites
came in the late 1960’s (Gould 1969). Subsequent authors (e.g., Glaubrecht 1990)
attributed the reduction in Poecilozonites to competition with the introduced
species Otala lactea and Rumina decollata. However, Rumina had been introduced
nearly a century earlier and Otala more than 40 years before, and noticeable
reductions in population numbers were not observed until the period between
1963 and 1969. It is more likely that the introduction of the predatory species
Euglandina rosea in late 1958 was responsible (see Table 1). Euglandina is now
widespread on the larger islands of Bermuda, with the possible exception of the
southeastern portion of the main island. Gonaxis quadrilateralis is just beginning to
spread beyond its original release sites, and Otala and Rumina are among the most
common and widespread snails on Bermuda, with little evidence of control by
Euglandina. Poecilozonites, on the other hand, has become quite uncommon and
appears to be threatened by extinction. Only two of the three extant species of
this endemic genus were found during this survey and neither of these is
common: P. circumfirmatus was found alive in two localities and fresh-dead
specimens of P. bermudensis were found in another. Several endemic and native
non-endemic species including Poecilozonites reinianus, Vertigo marki, Vertigo
bermudensis and Strobilops hubbardi appear to be restricted to the southeastern
portion of the main island near caves and remnant stands of native vegetation.

It is hoped that this work represents another stepping stone in the informal
series investigating the history of Bermuda’s mollusk fauna, and - most
importantly - that it provides the necessary baseline data for continued
monitoring of a rapidly changing faunal composition.

ACKNOWLEDGMENTS

RB’s 1983 collecting in Bermuda was made possible by a grant from the
Studienstiftung des deutschen Volkes (Federal Republic of Germany) and a

December 30, 2000 Bieler @ Slapcinsky 83

Sydney L. Wright Fellowship from the Bermuda Biological Station. The 1988
survey was supported by a grant of the Bermuda Department of Agriculture &
Fisheries, Paget, Bermuda; and Daniel J. Hilburn’s and Wolfgang Sterrer’s
hospitality during RB’s visit were much appreciated. Daniel J. Hilburn (then with
the Department of Agriculture, now with the Oregon Department of Agriculture,
and Petra Sierwald (then with DMNH, now at FMNH) provided invaluable
assistance during collecting. Computerization and archival-quality rehousing of
FMNH’s land snail collection is supported under National Science Foundation
Grant DEB-9616372 to RB.

We thank the following colleagues for loans of material, assistance during
our visits, help with literature searches, locality data, or information on their
Bermuda holdings: the late Water E. Sage II] (AMNH), David Robinson (ANSP),
Robert J. van Syoc (CAS), Frederick J. Collier and Stephen J. Gould (MCZ),
Wolfgang Sterrer and Penny Hill (BAMZ), John B. Burch (UMMZ), Fred
Thompson and Kurt Auffenberg (FLMNH), as well as Neil Fahy (Daly City,
California) for access to his private collection. We would also like to thank Robert
Cowie (BPBM) for his advice on various publication dates and his comments on
the genus Tornatellides. Jochen Gerber (FMNH) and an anonymous reviewer
provided valuable comments on an earlier draft of the manuscript.

This is Contribution No. 16 of the Bermuda Biodiversity Project (BBP),
Bermuda Aquarium, Natural History Museum and Zoo.

PERATURE GIteD

Abbott, R. T. 1989. Compendium of landshells. A color guide to more than
2,000 of the world’s terrestrial shells. American Malacologists, Inc.,
Melbourne, Florida and Burlington, Massachusetts. viiit 240 pp.

Adams, C. B. 1841. “Ward’s letter.” American Journal of Science, 40: 271.

Adams, C. B. 1845. Specierum novarum conchyliorum, in Jamaica repertorum,
synopsis. Proceedings of the Boston Society of Natural History, 2: 1-17
[January].

Adams, C. B. 1849. Descriptions of supposed new Helicidae from Jamaica.
Contributions to Conchology, no. 3: 33-38.

Albers, J. C. 1850. Die Heliceen, nach natiirlicher Verwandtschaft systematisch
geordnet. T. C. F. Enslin, Berlin. 262 pp.

Albers, J. C. 1860. Die Heliceen, nach natiirlicher Verwandtschaft systematisch
geordnet. Zweite Ausgabe [second edition, edited by E. von Martens]. W.
Engelmann, Leipzig. xviii + 359 pp.

Aldrich, T. H. 1889. H. (Fruticola) similaris, Fer., Triodopsis appressa, Say, Stenogyra
decollata, L., in Bermuda. The Nautilus, 3(1): 9.

84 NEMOURIA No. 44

Ancey, C. F. 1887. Description of new genera or subgenera of Helicidae. The
Conchologists’ Exchange, 1(9-10): 53-54.

Ancey, C. F. 1903. Etudes sur la faune malacologique des Iles Sandwich, 5° Partie.
Journal de Conchyliologie, 51(4): 295-307, pl. 12.

Angas, G. F. 1884. On the terrestrial Mollusca of Dominica, collected during a
recent visit to that island. Proceedings of the Scientific Meetings of the
Zoological Society of London, for the year 1883, part 4: 594-597 [April 1884].

Auffenberg, K. & L. A. Stange. 1986. Snail-eating snails of Florida. Entomology
Circular No. 285, 4 pp.; Florida Department of Agriculture and Consumer
Services, Division of Plant Industry.

Baker, H. B. 1922. Notes on the radula of the Helicinidae. Proceedings of the
Academy of Natural Sciences of Philadelphia, 74: 29-67, pls. 3-7 [8 August].

Baker, H. B. 1925. Nomenclature of Veronicellidae (Vaginulidae). The Nautilus,
39(1): 13-18 [8 July].

Baker, H. B. 1926. North American Veronicellidae. Proceedings of the Academy
of Natural Sciences of Philadelphia, 77: 157-184, pls. 3-6.

Bartram, J. T. 1878. Sea and land shells of Bermuda. Bermuda Almanack and
Directory 1878, pp. 105-108.

Beck, H. 1837-1838. Index Molluscorum praesentis aevi Musei Principis
Augustissimi Christiani Frederici. Pp. 1-100, 1837; pp. 101-124 and appendix
(8 pp.), 1838. Hafniae.

Bennett, F. D. @ I. W. Hughes. 1959. Biological control of insect pests in
Bermuda. Bulletin of Entomological Research, 50(3): 423-434.

Bennett, F. D., M. J. W. Cock, I. W. Hughes, F. J. Simmonds & M. Yaseen. 1985.
Biological control in Bermuda. pp. 119-142 in Cock, M. J. W. (ed.). A review
of biological control of pests in the Commonwealth Caribbean and Bermuda
up to 1982. Commonwealth Institute of Biological Control Technical
Communication No. 9.

Bequaert, J. C. @ W. B. Miller. 1973. The mollusks of the arid southwest with an
Arizona check list. The University of Arizona Press, Tucson. xvi + 271 pp.
Binney, A. 1837-1841. A monograph of the Helices inhabiting the United States.
Boston Journal of Natural History , 1: 466-495 (May 1837); 3: 353-394 (July
1840); 405-438 (“November 1840”) [dates teste A. A. Gould, 1851: 9; but last

part dated as 1841 by Turgeon et al., 1998: 147, 294].

Binney, A. 1842. Descriptions of some of the species of naked, air-breathing
Mollusca, inhabiting the United States. Boston Journal of Natural History,
4(2): 163-175.

Binney, W. G. 1876. On the lingual dentition, jaw, and genitalia of Carelia,
Onchidella, and other Pulmonata. Proceedings of the Academy of Natural
Sciences of Philadelphia for 1876: 183-192, pl.6.

December 30, 2000 Bieler & Slapcinsky 85

Binney, W. G. 1884. Notes on the jaw and lingual dentition of pulmonate
mollusks. Annals of the New York Academy of Sciences, 3: 79-136, pls. 2-17.

Bland, T. 1860. Remarks on certain species of North American Helicidae. Annals
of the Lyceum of Natural History in New York, 7: 69-95.

Bland, T. 1861. On the geographical distribution of the genera and species of
land shells of the West India islands; with a catalogue of the species of each
island. Annals of the Lyceum of Natural History of New York, 7: 335-362.

Bland, T. 1866. Remarks on the origin and distribution of the operculated land
shells which inhabit the continent of America and the West Indies, with a
catalogue of the American species. American Journal of Conchology, 2(1): 54
63, 2(2): 136-143, 2(4): 349-370 [subsequent parts with variant titles].

Bland, T. 1875. Notes on certain terrestrial mollusks, with descriptions of new
species. Annals of the Lyceum of Natural History of New York, 11(1876): 72-
87 [February 1875].

Bland, T. 1881. List of the land shells of Bermuda. Pp. 256-257 in: A. R. Wallace,
Island life; or, the phenomena and causes of insular faunas and floras,
including a revision and attempted solution of the problem of geological
climates. New York, Harper & brothers; xvi + 522 pp.

Bland, T. 1895. List of the land-shells of Bermuda. Pp. 270-271 in: A. R. Wallace,
Island life; or the phenomena and causes of insular faunas and _ floras,
including a revision and attempted solution of the problem of geological
climates. Second and revised edition. London, MacMillan and Co.; xx + 563
pp.

Bland, T. & W.G. Binney. 1873. On the lingual dentition of certain terrestrial
Pulmonata foreign to the United States. Annals of the Lyceum of Natural
History of New York, 10(1872): 219-225 [January 1873].

Boettger, O. 1884. Lebende Vertreter zweier Hochheimer untermiocaner
Landschnecken. Neues Jahrbuch fiir Mineralogie, Geologie und
Palaeontologie, 2: 139.

Bourguignat, J..R. 1925. Dates des livraisons de |’Histoire des Mollusques de
Férussac et Deshayes, vérifées d’aprés le registre de l’éditeur Bailliére, de
Paris. Journal de Conchyliologie, 69: 15-18.

Brown, A. D. 1861. Description of two new species of Helix. Proceedings of the
Academy of Natural Sciences of Philadelphia, 1861: 333.

Brown Jr., W. L. 1987. Punctuated equilibrium excused: the original examples fail
to support it. Biological Journal of the Linnean Society, 31: 383-404.

Bruguiére, [J.-G.]. 1789-1792. Encyclopédie méthodique. Histoire naturelle des
vers. Tome 1. Panckoucke, Paris; Plomteux, Liége. xviii + 757 pp. [pp. 1-344,
June 1789; pp. 345-757, 13 February 1792; teste Cowie 1998].

Bryant, O. 1905. Some notes on Bermudian mollusks. The Nautilus, 18(11): 129-
131.

86 NEMOURIA No. 44

Bulman, K. 1990. Life history of Carychium tridentatum (Risso, 1826) (Gastropoda:
Pulmonata: Ellobiidae) in the laboratory. Journal of Conchology, 33(6): 321-
333;

Chadwick, G. H. 1899. Notes on Polygyra appressa. The Nautilus, 13(5): 54-55.

Clapp, G. H. 1918. New southern forms of Carychium and Thysanophora. The
Nautilus, 31(3): 73-75, pl .8.

Clarke, B., J. Murray & M. S. Johnson. 1984. The extinction of endemic species
by a program of biological control. Pacific Science, 38(2): 97-104.

Clench, W. J. @ R. D. Turner. 1962. New names introduced by H. A. Pilsbry in
the Mollusca and Crustacea. Academy of Natural Sciences of Philadelphia
Special Publication, 4: 1-218.

Cock, M. J. W. 1985. A review of biological control of pests in the
Commonwealth Caribbean and Bermuda up to 1982. Technical
Communication, Commonwealth Institute of Biological Control, 9: xii +
218 pp. (ISBN 0851985505).

Cowan, C. F. 1969. Cuvier’s Régne animal, First Edition. Journal of the Society
for the Bibliography of Natural History, 5(3): 219.

Cowie, R. H. 1997. Catalog and bibliography of the nonindigenous nonmarine
snails and slugs of the Hawaiian Islands. Bishop Museum Occasional Papers,
50: 1-66.

Cowie, R. H. 1998. Catalog of the nonmarine snails and slugs of the Samoan
Islands. Bishop Museum Bulletin in Zoology, 3: viii + 122 pp. [13 January].

Cuvier, [G.] [“1817”] 1816. Le régne animal distribué d’aprés son organisation,
pour servir de base a l'histoire naturelle des animaux et d’introduction a
l'anatomie comparée. Tome II, cotenant les reptiles, les poissons, les
mollusques et les annélides. Deterville, Paris. xviii + 532 pp. [7 December
1816, teste Cowan, 1969].

da Costa, E. M. 1778. Historia naturalis testaceorum Britanniae, or, the British
conchology; containing the descriptions and other particulars of natural
history of the shells of Great Britain and Ireland: illustrated with figures. In
English and French. Da Costa, London. xii + 254 + viii pp., 17 pls.

Davis, C. A. 1904. Notes on the Mollusca of the Bermuda Islands. The Nautilus,
17(11): 125-130, pl. 4.

Davis, C. J. &@ G. D. Butler, Jr. 1964. Introduced enemies of the Giant African
Snail, Achatina fulica Bowdich, in Hawaii (Pulmonata: Achatinidae).
Proceedings of the Hawaiian Entomological Society, 18(3): 377-389.

Deisler, J. E. & R. T. Abbott. 1984. Range extensions of some introduced land
mollusks in the Bahama Islands, with first reports for four species. The
Nautilus, 98(1): 12-17.

Deshayes, G. P. 1832. Encyclopédie méthodique. Histoire naturelle des vers..., 2
(FRLMYT): 145-594 [29 September].

December 30, 2000 Bieler & Slapcinsky 87

Draparnaud, J. 1801. Tableau des mollusques terrestres et fluviatiles de la France.
Renaud, Montpellier; Bossange, Masson & Besson, Paris. 116 pp. [14 July].

Draparnaud, J.-P.-R. 1805. Histoire naturelle des mollusques terrestres et
fluviatiles de la France. Louis Colas, Paris. iviii, 1-164 pp., pls. 1-13
[posthumously published].

Drouét, H. 1859. Essai sur les mollusques terrestres et fluviatiles de la Guayane
Francaise. J.-B. Bailliére, Paris. 109 pp., 4 pls.

Dundee, D. S. 1970. Introduced Gulf Coast molluscs. Tulane Studies in Zoology
and Botany, 16(3): 101-115.

Dundee, D. S., P. H. Philipps & J. D. Newsom. 1967. Snails on migratory birds.
The Nautilus, 80(3): 89-91.

Eldredge, N. & S. J. Gould, 1972. Punctuated equilibria: an alternative to
phyletic gradualism. pp. 82-115. In: T. J. M. Schopf (ed.), Models in
Paleobiology. Freeman, Cooper & Company, San Francisco. 250 pp.

Ellis, A. E. 1951. Proposed addition to the “Official List of Specific Trivial Names
in Zoology” of the names of certain non-marine species in the phylum
Mollusca. Bulletin of Zoological Nomenclature, 2(4): 125-128.

Fechter, R. @ G. Falkner. 1990. Weichtiere. Europaische Meeres- und
Binnenmollusken. Steinbachs Naturftihrer (G. Steinbach, ed.), Mosaik
Verlag, Munich. 287 pp.

Ferguson, D.C., D.J. Hilburn & B. Wright. 1991. The Lepidoptera of Bermuda:
their food plants, biogeography, and means of dispersal. Memoirs of the
Entomological Society of Canada, 158: iii, 1-105.

Férussac, D. de. 1821-1822. Tableaux systématiques des animaux mollusques
classés en familles naturelles, dans lesquels on a établi la concordance de tous
les systemes; suivis d’un prodrome générale pour tous les mollusques
terrestres ou fluviatiles, vivants ou fossiles. A. Bertrand, Paris, and J. B.
Sowerby, London. ixlviii, 1-27, 1-110 [+i] pp. [published in issues; see
Sherborn & Woodward, 1901, and Cowie, 1998, for dates].

Férussac, de [A. E. J. P. J. F. d’A.] 1823. P. 966 in de Férussac & G.-P. Deshayes,
1820-1851. Histoire naturelle générale et particuliére des mollusques
terrestres et fluviatiles tant des espéces que l’on trouve aujourd’hui vivantes,
que des dépouilles fossiles de celles qui n’existent plus; classés d’aprés les
caractéres essentiels que présentent ces animaux et leurs coquilles, Tome 2.,
Paris, J.-B. Bailliére; xvi + 260 pp., 247 pls.

Férussac, D. de & G-P. Deshayes. 1819-1851. Histoire naturelle générale et
particuliére des mollusques terrestres et fluviatiles tant des espéces que l’on
trouve aujourd’hui vivantes, que des dépouilles fossiles de celles qui
n’existent plus; classés d’aprés caractéres essentiels que présentent ces
animaux et leurs coquilles. Bailliére, Paris. Issued in 42 parts. [see Sherborn

88 NEMOURIA No. 44

& Woodward, 1901; Bourguignat, 1925, and Kennard, 1942, for individual
dates].

Fisher, T. W., R. E. Orth & S. C. Swanson. 1980. Snail against snail. California
Agriculture (Nov. - Dec.): 3 pp.

Fischer [von Waldheim], G. 1807. Museum Demidoff, ou, Catalogue
systématique et raisonnée des curiosités de la nature et de l’art: données a
l'Université Impériale de Moscou par son ecellence Monsieur Paul de
Demidoff. Volume 3. Imprimerie de l'Université Imperiale, Moscow.
Frontispiece, ix + 330 pp., 6 pls.

Frith, M. A. M. 1891. Essay [3]. Pp. 14-18 (with copies of letters by Jones and
Bartram), in: Five essays as furnished to the Board of Agriculture in response
to an advertisement offering prizes for the descriptions of and the history of
the Spiral Snails, and the most efficacious, expeditious, and economical
methods to effect their extermination. Gregory V. Lee, Queen’s Printer,
Hamilton, Bermuda.

Gerber, J. 1996. Revision der Gattung Vallonia Risso 1826 (Mollusca:
Gastropoda: Valloniidae). Schriften zur Malakozoologie aus dem Haus der
Natur - Cismar, 8: 1-227.

Giusti, F., G. Manganelli & P. J. Schembri. 1995. The non-marine molluscs of the
Maltese Islands. Museo Regionale de Scienze Naturali, Torino, Monografie
XV: 1-607.

Glaubrecht, M. 1990. Bermuda - ein evolutionarer Mikrokosmos im Atlantik?
Zur Naturgeschichte einer isolierten Inselgruppe. Natur und Museum,
120(7): 209-222.

Godan, D. 1983. Pest slugs and snails - biology and control. Springer-Verlag,
Berlin, Heidelberg, New York. x + 445 pp.

Gould, A. A. 1843. Monograph of the species of Pupa found in the United States
[continued]. Boston Journal of Natural History, 4(3): 350-360, pl. 16.

Gould, A. A. 1848. Untitled. [“Dr. Gould had recently examined the Shells
collected by Mr. J. Bartlett, in the south-western States, for the late Dr.
Binney... He gave descriptions of the following...” (p. 37)]. Proceedings of the
Boston Society of Natural History, 3: 37-41.

Gould, A. A. (editor). 1851. The terrestrial air-breathing mollusks of the United
States, and the adjacent territories of North America: described and
illustrated by Amos Binney. Vol. 1. C.C. Little & J. Brown, Boston. xxix +
366 pp.

Gould, S. J. 1966. Allometry in Pleistocene land snails from Bermuda: the
influence of size upon shape. Journal of Paleontology, 40(5): 1131-1141.
Gould, S. J. 1968. Ontogeny and the explanation of form: an allometric analysis.

Journal of Paleontology, 42(5), part 2 of 2: 81-98, pl. 10.

December 30, 2000 Bieler & Slapcinsky 89

Gould, S. J. 1969. An evolutionary microcosm: Pleistocene and Recent history of
the land snail P. (Poecilozonites) in Bermuda. Bulletin of the Museum of
Comparative Zoology, 138(7): 407-531, frontispiece, pls. 1-5 [December].

Gould S. J. 1970a. Coincidence of climatic and faunal fluctuations in Pleistocene
Bermuda. Science, 168: 527-573 [May].

Gould, S. J. 1970b. Land snail communities and Pleistocene climates in Bermuda:
a multivariate analysis of microgastropod diversity. Proceedings of the North
American Paleontological Convention (Field Museum of Natural History,
Chicago, September 5-7, 1969), 1(part E): 486-521 [October].

Gould S. J. 1971la. Environmental control of form in land snails: a case of
unusual precision. The Nautilus, 84(3): 86-93 [January].

Gould S. J. 1971b. Precise but fortuitous convergence in Pleistocene land snails
from Bermuda. Journal of Paleontology, 45(3): 409-418 [May].

Gould S. J. 1991. Unenchanted evening. Natural History, 9/91: 4-14.

Gray, A. F. 1942. A Cerion found in Bermuda. The Nautilus, 55(3): 104.

Guilding, L. 1828. Observations on the zoology of the Caribaean islands
[continued]. The Zoological Journal, 3: 527-544, pls. 15, 26 (Supp.), 27
(Supp.), 28 (Supp.).

Gulick, A. 1904. The fossil land shells of Bermuda. Proceedings of the Academy
of Natural Sciences of Philadelphia, 56: 406-425, pl. 36.

Haas, F. 1952. On the mollusk fauna of the landlocked waters of Bermuda.
Fieldiana, Zoology, 34(8): 101-105.

Hadfield, M. G. & B. S. Mountain. 1981. A field study of a vanishing species,
Achatinella mustelina (Gastropoda, Pulmonata), in the Waianae Mountains of
Oahu. Pacific Science, 34(4) [1980]: 345-358.

Hanna, G. D. 1966. Introduced mollusks of western North America. Occasional
Papers of the California Academy of Sciences, 48: 1-108, pls. 14.

Hastings, G. R. 1892. The broken-tail snail in Bermuda [and reply by C. V. Riley
and L. O Howard]. Insect Life, 4(9-10): 334.

Heilprin, A. 1889 [“With additions by Prof. J. Playfair McMurrich, Mr. H. A.
Pilsbry, Dr. George Marx, Dr. P. R. Uhler, and Mr. C. H. Bollman”]. The
Bermuda Islands: a contribution to the physical history and zoology of the
Somers Archipelago: with an examination of the structure of coral reefs:
researches undertaken under the auspices of the Academy of Natural
Sciences of Philadelphia. Heilprin, Philadelphia. 231 pp., 17 pls.

Hilburn, D. J. @ R. D. Gordon. 1989. Coleoptera of Bermuda. Florida
Entomologist, 72: 673-692.

Hoffmann, H. 1927. On some North American Vaginulidae. Proceedings of the
Academy of Natural Sciences of Philadelphia, 79: 209-212.

Honigmann, L. 1914. Beitrag zur Landmolluskenfauna der Bermudas.
Zoologischer Anzeiger, 44(8): 375-377 [June].

90 NEMOURIA No. 44

Hubricht, L. 1985. The distribution of the native land mollusks of the eastern
United States. Fieldiana, Zoology, (NS) 24: iviii, 1-191.

Hughes, I. W. (unpublished report, 1962). Annual Report of the Assistant
Director of Agriculture for the year ending 31st December, 1962.

Hughes, I. W. (unpublished report, 1963). Annual Report of the Assistant
Director of Agriculture for the year ending 31st December, 1963.

Hughes, I. W. (post 1968). Recent terrestrial mollusks of Bermuda. Department
of Agriculture and Fisheries, Bermuda; 3 pp. handout.

Hughes, I. W. (post 1969). Fossil land snails of Bermuda. Department of
Agriculture and Fisheries, Bermuda; 2 pp. handout.

Hughes, I. W. 1971. Giant African snail. Monthly Bulletin of the Bermuda
Department of Agriculture and Fisheries, 41(6): 48.

Hutton, T. 1834. On the land shells of India. Journal of the Asiatic Society of
Bengal, 3: 81-93.

ICZN (International Commission on Zoological Nomenclature). 1955. Opinion
336. Additions to the “Official List of Specific Names in Zoology” of the
specific names of one hundred and twenty-two non-marine species of the
phylum Mollusca. Opinions and Declarations rendered by the International
Commission on Zoological Nomenclature (F. Hemming, ed.), 10(3): 77-108.

ICZN (International Commission on Zoological Nomenclature). 1994. Opinion
1766. Tortaxis Pilsbry, 1906 and Allopeas Baker, 1935 (Mollusca, Gastropoda):
conserved by the designation of a neotype for Achatina erecta Benson, 1842.
Bulletin of Zoological Nomenclature, 51(2): 162-163.

Johnson, C. W. 1900. Some notes on Rumina decollata Linn. The Nautilus,
13(10): 117.

Jones, J. M. 1859. The naturalist in Bermuda; a sketch of the geology, zoology,
and botany, of that remarkable group of islands; together with meteorological
observations. Reeves & Turner, London. xii + 200 pp., 1 map [we have been
unable to locate an “1852” edition of this work as cited by some authors].

Jones, J. M. 1864. Contributions to the natural history of the Bermudas. Part I.
Mollusca. J. Bowes & Sons, Halifax. 13 pp. [also found cited as Transaction
Nova Scotian Institute of Natural Science of Halifax, N.S. 1(2): 14-26; not
seen].

Jones, J. M. 1876. Mollusca. The visitor’s guide to Bermuda, with a sketch of its
natural history. 156 pp., 5 woodcuts, Reeves & Turner, London.

Karlin, E. J. & J. A. Naegele. 1966. Biology of the Mollusca of Greenhouses in
New York State. Cornell University Agricultural Station Memoir, 372: 1-35.

Kennard, A. S. 1942. The Histoire and Prodrome of Férussac. Proceedings of the
Zoological Society of London, 25(1): 12-17.

Kerney, M. P. & R. A. D. Cameron. 1979. A field guide to the land snails of
Britain and North-West Europe. Collins, London; 288 pp., 24 pls.

December 30, 2000 Bieler & Slapcinsky 91

Kinzie III, R. A. 1992. Predation by the introduced carnivorous snail Euglandina
rosea (Férussac) on endemic aquatic lymnaeid snails in Hawaii. Biological
Conservation, 60: 149-155.

Kobelt, W. 1880. Die geographische Verbreitung der Mollusken (Schluss).
Jahrbticher der Deutschen Malakozoologischen Gesellschaft, 7(3): 241-286
[July).

Krauss, N. L. H. 1964. Investigation on biological control of Giant African
(Achatina fulica) and other land snails. The Nautilus, 78(1): 21-27.

Kutchka, G. M. 1934. Notes on the fossil land shells from Bermuda in the
collection of the Carnegie Museum. Annals of the Carnegie Museum, 22(2-
4): 293-302, pls. 19-21.

Lamarck, [J. B. P. A. de Monet de] 1822. Histoire naturelle des animaux sans
vertébres, présentant les caractéres généraux et particuliers de ces animaux,
leur distribution, leurs classes, leurs familles, leurs genres, et la citation des
principales espéces qui s'y rapportent ..., 6(2): 232 pp.; Lamarck, Paris [April].

Lane, W. G. 1891. The spiral snails of Bermuda. [Essay 1] Pp. 1-8, in: Five essays
as furnished to the Board of Agriculture in response to an advertisement
offering prizes for the descriptions of and the history of the Spiral Snails, and
the most efficacious, expeditious, and economical methods to effect their
extermination. Gregory V. Lee, Queen’s Printer, Hamilton, Bermuda.

Lea, I. 1841. On fresh water and land shells. Proceedings of the American
Philosophical Society, 2(17): 30-35.

Leidy, J. 1851. Special anatomy of the terrestrial Gasteropoda of the United
States. Pp. 198-248, pls. 1-16 in: Gould, A. A. (ed.), The terrestrial air-
breathing mollusks of the United States, and the adjacent territories of North
America: described and illustrated by Amos Binney. Vol. I. Charles C. Little
& James Brown, Boston.

Linné, C. 1758. Systema naturae per regna tria naturae, secundum classes,
ordines, genera, species, cum characteribus, differentiis, synonymis, locis. 12"
ed. Volume 1. Stockholm. 824 pp.

Manganelli, G. & F. Giusti. 1997. Helix draparnaudi Beck, 1837 (currently
Oxychilus draparnaudi; Mollusca, Gastropoda): proposed conservation of the
specific name. Bulletin of Zoological Nomenclature, 54(3): 148-151.

Martens, E. von. 1889. Eine ausgestorbene Landschnecke von den Bermuda-
Inseln. Sitzungsberichte der Gesellschaft naturforschender Freunde zu Berlin,
17 December 1889, pp. 201-202.

Mazyck, W. G. 1889. Note on Helix Bermudensis, Pfeiffer. Proceedings of the
Elliott Society, 1888: 210-211 [19 March 1889].

Monkman. D. 1982a. Distribution of predaceous snails Euglandina rosea and
Gonaxis quadrilateralis in Bermuda. Unpublished report, Department of
Agriculture and Fisheries, Bermuda.

92 NEMOURIA No. 44

Monkman, D. 1982b. On the trail of the snail. Monthly Bulletin of the
Department of Agriculture and Fisheries, Bermuda, 53(9): 70-72.

Moquin-Tandon, A. 1855. Histoire naturelle des mollusques terrestres et
fluviatiles de France contenant des études générales sur leur anatomie et leur
physiologie et la description particuliére des genres, des espéces et des
variétés. Atlas. J-B. Bailliére, Paris. 54 pls.

Morch, O. A. L. 1878. Catalogue of West-India shells in the collection of Dr. C.
M. Poulsen. Bianco Luno, Copenhagen. 16 pp.

Morelet, A., 1860. Description de nouvelles espéces de l'Afrique occidentale,
rapportées par M. le capitaine Vignon. Journal de Conchyliologie, 8: 189-191
[April].

Morrison, J. P. E. 1953. Two new American species of Strobilops. The Nautilus,
67(2): 53-55, pl. 6 [October].

Miller, O. F. 1774. Vermium terrestrium et fluviatilium, seu animalium
infusoriorum, helminthicorum, et testaceorum, non marinorum, succincta
historia. Volume 2. Mdller, Copenhagen and Leipzig. [i-xxxv], 1-214 pp.,
index [215-224].

Murray, J., E. Murray, M.S. Johnson & B. Clarke. 1989. The extinction of Partula
on Moorea. Pacific Science, 42(3-4) (1988): 150-153 [January].

Naggs, F. 1989. Gulella bicolor (Hutton) and its implications for the taxonomy of
streptaxids. Journal of Conchology, 33(3): 165-168, pl. 18 [June].

Naggs, F. 1992. Case 2833. Tortaxis Pilsbry, 1906 and Allopeas Baker, 1935
(Mollusca, Gastropoda): proposed conservation by the designation of a
neotype for Achatina erecta Benson, 1842. Bulletin of Zoological
Nomenclature, 49(4): 258-260.

Naggs, F. 1994. The reproductive anatomy of Paropeas achatinaceum and a new
concept of Paropeas (Pulmonata: Achatinoidea: Subulinidae). Journal of
Molluscan Studies, 60(2): 175-191.

Olmsted, J. M. D. 1917. Notes on the locomotion of certain Bermudian mollusks.
The Journal of Experimental Zoology, 24(2): 223-236.

Orbigny, A. d’. 1835. Synopsis terrestrium et fluviatilium molluscorum, in suo
per Americam merdionalem itinere, ab A. D’Orbigny, collectorum. Magasin
de Zoologie, 5, Classe V, no. 61 [terrestrial mollusks]: [1]-22.

Orbigny, A. d'. 1841-1846. Mollusques. In: Ramon de la Sagra: Histoire physique,
politique et naturelle de I'Ile de Cuba. - 1: 208, pls. 1-14 (1841); 209-264, pls.
15-17 (1842); 2: 1-112, pls. 1-7 (1842); 113-380, pls.8-24 ("1846", 1853?);
Atlas (1842) [fide Keen, 1971]; A. Bertrand, Paris.

Oughton, J. 1948. A zoogeographical study of the land snails of Ontario.
University of Toronto Studies, Biological Series, no. 57: ixi, 1-128, 2 tables.

Peake, J. F. 1981. The land snails of islands - a dispersalist’s viewpoint (chapter
19). Pp. 247-263 in: The evolving biosphere (P. L. Forey, ed.), Chance,

December 30, 2000 Bieler & Slapcinsky 93

change & challenge (P. H. Greenwood, gen. ed.). British Museum (Natural
History), Cambridge University Press.

Peile, A. J. 1924. Notes on the genus Poecilozonites; with descriptions of new
species. Proceedings of the Malacological Society of London, 16(1): 16-23.
Peile, A. J. 1925. Further notes on Poecilozonites. Proceedings of the Malacological

Society of London, 16(5): 257-258.

Peile, A. J. 1926. The Mollusca of Bermuda. Proceedings of the Malacological
Society, London, 17(2-3): 71-98.

Peniston, A. 1891. Essay [5]. Pp. 21-22, in: Five essays as furnished to the Board of
Agriculture in response to an advertisement offering prizes for the
descriptions of and the history of the Spiral Snails, and the most efficacious,
expeditious, and economical methods to effect their extermination. Gregory
V. Lee, Queen’s Printer, Hamilton, Bermuda.

Pfeiffer, L. 1839. Bericht tber die Ergebnisse meiner Reise nach Cuba im Winter
1838-1839. [Wiegmann’s] Archiv fiir Naturgeschichte, 5(1): 346-358.

Pfeiffer, L. 1840. Uebersicht der im Januar, Februar und Marz 1839 auf Cuba
gesammelten Mollusken. [Wiegmann’s] Archiv fiir Naturgeschichte, 6(1):
250-261.

Pfeiffer, L. 1841a. Symbolae ad historiam heliceorum. T. Fischer, Kassel. 100 pp.

Pfeiffer, L. 1841b. Beitrage zur Molluskenfauna Deutschlands, insbesondere der
Osterreichischen Staaten. Archiv fiir Naturgeschichte, 7(1): 215-230.

Pfeiffer, L. 1845. Description of twenty-two new land-shells, belonging to the
collection of Mr. H. Cuming. Proceedings of the Zoological Society of
London, 13: 63-69 (September).

Pfeiffer, L. 1847-1848. Monographia Heliceorum viventium. Sistens descriptiones
systematicas et criticas omnium huius familiae generum et specierum hodie
cognitarum. Vol. 1. F. A. Brockhaus, Leipzig. xxxii + 484 pp.

Pfeiffer, L. 1849. Description of twenty-nine new species of Helicina, from the
collection of H. Cuming. Proceedings of the Zoological Society of London,
16 (1848): 119-125 [25 April 1849].

Pfeiffer, L. 1852. Uebersicht des gegenwartigen Zustandes der Heliceenkunde
(Schluss). Zeitschrift fiir Malakozoologie, 9(10): 145-151.

Pfeiffer, L. 1853. Diagnosen neuer Heliceen. Zeitschrift fiir Malakozoologie,
10(4): 51-58.

Pfeiffer, L. 1857. Descriptions of thirty-one new species of land-shells, from Mr.
Cuming’s Collection. Proceedings of the Zoological Society of London, 25:
107-113 [August-September].

Pfeiffer, L. 1859. Monographia Heliceorum viventium. Sistens descriptiones
systematicas et criticas omnium huius familiae generum et specierum hodie
cognitarum. Vol. 4. F. A. Brockhaus, Leipzig. 920 pp.

94 NEMOURIA No. 44

Pfeiffer, L. 1863. Zwei neue Landschnecken von Bermuda. Malakozoologische
Blatter, 11(1864): 1-2.

Pfeiffer, L. 1868. Monographia Heliceorum viventium. Sistens descriptiones
systematicas et criticas omnium huius familiae generum et specierum hodie
cognitarum. Vol. 5. F. A. Brockhaus, Leipzig. xii + 565 pp.

Pfeiffer, L. 1879-1881 [posthumous, edited by S. Clessin]. Nomenclator
Heliceorum viventium quo continetur nomina omnium hujus familiae
generum et specierum hodie cognitarum disposita ex affinitate naturali. T.
Fischer, Kassel. Parts 1-6, pp. 1-384 (1879); 617 pp. (1881).

Philippi, R. A. 1843. Helix Tab. HI. Abbildungen und Beschreibungen neuer oder
wenig gekannter Conchylien, unter Mithilfe mehrerer deutscher
Conchyliologen herausgegeben, 1(3): 47-52, pl. 3.

Pilsbry, H. A. The following dates of publication for H.A. Pilsbry are all teste Clench &
Turner (1962):

Pilsbry, H. A. 1888. On the helicoid land molluscs of Bermuda. Proceedings of
the Academy of Natural Sciences of Philadelphia, 40: 285-291, pl. 17
(December) [also printed in Heilprin, 1889: 191-201, pl.16].

Pilsbry, H. A. 1889. New and little-known American molluscs. No. 1. Proceedings
of the Academy of Natural Sciences of Philadelphia, 41(19): 81-88 [May], 89,
pl. 3 June}.

Pilsbry, H. A. 1890a. Poecilozonites circumfirmatus var. corneus. The Nautilus, 3(8):
95 [January].

Pilsbry, H. A. 1890b. New and little known American mollusks, No. 3.
Proceedings of the Academy of Natural Sciences of Philadelphia, 42(20): 296-
302, pl. 5 [October].

Pilsbry, H. A. 1891. On Helix harfordiana Cooper, and other shells. The Nautilus,
5(4): 39-42, pl. 2 [19 August].

Pilsbry H. A. 1900. The air-breathing mollusks of the Bermudas. Transactions of
the Connecticut Academy of Sciences, 10(2): 491-509, pl. 62 [September].
Pilsbry, H. A. 1916-1918. Pupillidae (Gastrocoptinae). Manual of Conchology
(Second Series: Pulmonata), 24 (part 93): 1-112, pls. 1-13 [December 1916];
(part 94): 113-176, pls. 14-29 [July 1917]; (part 95): 177-256, pls. 30-38

[November 1917]: (part 96): 257-380, i-xii, pls. 39-49 [April 1918].

Pilsbry, H. A. 1919. Pupillidae (Gastrocoptinae, Vertigininae). Manual of
Conchology (Second Series: Pulmonata), 25 (part 98): 65-144, pls. 6-10
[February]; 25 (part 99): 145-224, pls. 11-18 [June].

Pilsbry, H. A. 1924. Recent and fossil Bermudian snails of the genus Poecilozonites.
Proceedings of the Academy of Natural Sciences of Philadelphia, 76: 1-9
[June].

Pilsbry, H. A. 1934. Notes on the fossil land shells from Bermuda. The Nautilus,
48(1): 34.

December 30, 2000 Bieler & Slapcinsky 95

Pilsbry, H. A. 1939-1948. Land Mollusca of North America (north of Mexico).
The Academy of Natural Sciences of Philadelphia Monographs, 3, volume
1(1): ixvii, 1-573, i:ix [December 1939]; 1(2): i-viii, 575-994, iix [August
1940]; 2(1): frontispiece, ivi, 1-520 [December 1946]; 2(2): ixlvii, 521-1113
[March 1948].

Pilsbry, H. A. & J. H. Ferriss. 1906. Mollusca of the southwestern states. II.
Proceedings of the Academy of Natural Sciences of Philadelphia, 58: 123-
175, pls. 5-9 [June].

Pilsbry, H. A. & E. G. Vanatta. 1909. A new land shell from Bermuda. The
Nautilus, 23(5): 63-64 [October].

Pilsbry, H. A. @ E. G. Vanatta 1924. Polygyra appressa linguifera and sancta
georgiensis. The Nautilus, 38(1): 4-6 [July].

Poey, F. 1851. Memorias sobre la: historia natural de la isla de Cuba,
acompaniadas de sumarios latinos y extractos en frances. Havana, Viuda de
Barcina. Vol. 1: 1461 pp., pls. 1-34.

Pokryszko, B. M. 1987. On the aphally in the Vertiginidae (Gastropoda:
Pulmonata: Orthurethra). Journal of Conchology, 32: 365-375

Preston, H. B. 1910. Additions to the non-marine molluscan fauna of British and
German East Africa and Lake Albert Edward. The Annals and Magazine of
Natural History, (8)6: 526-536, pls. 7-9.

Prime, T. 1853. List of shells and corals collected at Bermuda, by Temple Prince
[sic], L.L.B. of New York. 1852. The Bermuda Pocket Almanack for 1853, p.
55.

Proschwitz, T. von. 1983. Vaxthussnackan Hawaiia minuscula (Binney) funnen i
Sverige, samt nagot om tankbara efterféljare. Fauna och flora, 78(6): 277-284.

Proschwitz, T. von. 1994. Lamellaxis clavulinus (Potiez & Michaud) as a greenhouse
species in Sweden. Journal of Conchology, 35(2): 184-185.

Pulteney, R. 1799. Catalogues of the birds, shells, and some of the more rare
plants of Dorsetshire. From the new ... edition of Hutchin’s history of that
county. 92 pp. London.

Rang, S. 1831. Description des coquilles terrestres recueillies pendant un voyage a
la cOte occidentale d’Afrique, et au Brésil. Annales des Sciences Naturelles,
24: 5-63, pl. 3 [September].

Redfield, J. H. 1853. Descriptions of new species of Helicidae. Annals of the
Lyceum of Natural History of New York, 6(4): 14-18.

Reeve, L. A. 1851-1854. Conchologia iconica: or, illustrations of the shells of
molluscous animals. Vol. VII. Containing a monograph of the genus Helix. L.
Reeve, London. 210 pls. + unpaginated text [see Cowie, 1998, for dates of
individual plates].

Rein, [J. J.] 1867. Bermuda shells. The Bermuda Pocket Almanack for 1867, pp.
75-77.

96 NEMOURIA No. 44

Richardson, L. 1986. Polygyracea: catalog of species (Part 3, Sagdidae). Tryonia,
13: 1-38.

Richardson, C. L. 1988. Streptaxacea: catalog of species. Part I Streptaxidae.
Tryonia, 16: i, 1-326.

Riedel, A. 1980. Genera zonitidarum - Diagnosen supraspezifischer Taxa der
Familie Zonitidae (Gastropoda, Stylommatophora). W. Backhuys,
Rotterdam, 197 pp. incl. 2 pls.

Robinson, D. G. 1999. Alien invasions: the effects of the global economy on non-
marine gastropod introductions into the United States. Malacologia, 41(2):
413-438.

[Say, T.] 1816. Conchology. p. [1-15], pls. 14. In: Nicholson, W., ed., American
edition of the British Encyclopedia, or dictionary of arts and sciences,
comprising an accurate and popular view of the present improved state of
human knowledge. Volume IV. Mitchell, Ames & White, Philadelphia. [not
seen]

Say, T. 1821. Descriptions of univalve shells of the United States. Journal of the
Academy of Natural Sciences of Philadelphia, 1(1): 149-179.

Say, T. 1822. Descriptions of univalve terrestrial and fluviatile shells of the
United States. Journal of the Academy of Natural Sciences of Philadelphia,
2(2): 370-381.

Say, T. 1825. Descriptions of some new species of fresh water and land shells of
the United States. Journal of the Academy of Natural Sciences of
Philadelphia, 5(1): 119-131.

Schileyko, A. A. @ H. P. M. G. Menkhorst. 1997. Composition and phylogenetic
relations of the Cochlicellidae (Gastropoda, Pulmonata). Ruthenica, 7(1): 51-
60.

Seddon, M. B. & D. T. Holyoak. 1993. Land Gastropoda of NW. Africa: New
distributional data and nomenclature. Journal of Conchology, 34(5): 321-
Sie) le

Sheppard, R. 1823. Description of seven new British fresh-water shells, with
observations upon many other species. Transactions of the Linnean Society
of London, 14: 148-170.

Sherborn, C. D. & B. B. Woodward. 1901. On the dates of publication of the
‘Histoire naturelle générale et particuliére des mollusques terrestres et
fluviatiles’ and the ‘Tableaux systématiques des animaux mollusques,’ by the
Barons Férussac and G. P. Deshayes. The Annals and Magazine of Natural
History, (7)8: 74-76.

Shuttleworth, R. J. 1853. Diagnosen neuer Mollusken. No. 5. Critische
Beleuchtung der zur Gruppe Sagda Beck, Albers gehoérenden westindischen
Heliceen. Berner Mittheilungen, Dec. 1853: 297-335.

December 30, 2000 Bieler & Slapcinsky 97

Shuttleworth, R. J. 1854. Diagnosen neuer Mollusken. No. 6. Beitrage zur
naheren Kenntniss der Land- und Siisswasser-Mollusken der Insel Portorico.
Berner Mittheilungen, Mar. 1854: 33-56.

Sierwald, P. 1988. Spiders of Bermuda. Nemouria, Occasional Papers of the
Delaware Museum of Natural History, 31: 1-24.

Simmonds, F. J. & I. W. Hughes. 1963. Biological control of snails exerted by
Euglandina rosea (Férussac) in Bermuda. Entomophaga, 8(3): 219-222.

Smith, B. J. @ J. Stanisic. 1998. Pulmonata. Pp 1037-1125 in Beesley, P.L., G. J.
B. Ross, & A. Wells (eds.), Mollusca: The southern synthesis. Fauna of
Australia. Vol. 5. CSIRO Publishing: Melbourne.

Smith, E. A. 1884. An account of the land and freshwater Mollusca collected
during the voyage of the ‘Challenger’ from December 1872 to May 1876.
Proceedings of the Scientific Meetings of the Zoological Society of London,
52(18-19): 258-281, pls. 22, 23.

Smith, E. A. 1894. A list of the land and fresh-water Mollusca collected by Dr. J.
W. Gregory in East Africa during his expedition to Mount Kenia, with
descriptions of a few new species. Proceedings of the Malacological Society of
London, 1(4): 163-168 [25 October].

Smith, H. H. 1912. Rumina decollata in Mobile and New Orleans. The Nautilus,
6(1): 4-6.

Solem, A. 1964. New records of New Caledonian nonmarine mollusks and an
analysis of the introduced mollusks. Pacific Science, 18(2): 130-137.

Solem, A. 1974. Island size and species diversity in Pacific Island land snails.
Malacologia 14 (“1973”; Proceedings of the Fourth European Malacological
Congress): 397-400.

Solem, A. 1989. Non-camaenid land snails of the Kimberley and Northern
Territory, Australia. I. Systematics, affinities and ranges. Invertebrate
Taxonomy, 2(4)(1988): 455-604 [22 March].

Solem, A. 1990. Limitations of equilibrium theory in relation to land snails.
International Symposium on Biogeographical Aspects of Insularity (Rome,
18-22 May 1987). Atti dei Convegni Lincei, 85: 97-116.

Spix, J. B. 1827. Testacea fluviatilia quae in itinere per Brasiliam annis
MDCCCXVII - MDCCCXX jussu et auspiciis Maximiliani Josephi I.
Bavariae regis augustissimi ... Digessit, descripsit et observationibus illustravit
Dr. J. A. Wagner. C. Wolf, Munich. pp. [i-iii] iv, 1-36 (text), [iii] (plate
explanations), pls. 1-29.

Sterki, V. 1893. Observations on Vallonia. Proceedings of the Academy of Natural
Sciences of Philadelphia, 1893: 234-279, pl. 8.

Thiele, J. 1929-31. Handbuch der systematischen Weichtierkunde. Vol. 1. 778
pp.; Gustav Fischer, Jena. [Part 1 (Loricata; Gastropoda: Prosobranchia): 1-
376, September or October 1929; Part 2 (Gastropoda: Opisthobranchia and

98 NEMOURIA No. 44

Pulmonata; additions; index for parts 1 and 2): 377-778, before November
1931).

Tomlin, J. R. le B. 1932. A supposed new Succinea from Bermuda. Proceedings of
the Malacological Society of London, 20(2): 98-100, pl. 9A.

Tristram, H. B. 1861. Catalogue of a collection of mollusks from Bermuda.
Proceedings of the Zoological Society of London, 1861: 403-405.

Tryon, G. W. 1887. Manual of Conchology; structural and systematic. With
illustrations of the species. Second series: Pulmonata, Vol. 3 (Helicidae: Vol.
1): G. W. Tryon, Philadelphia; pp. 1-313, pls. 1-63.

Turgeon, D. D., J. F. Quinn, Jr., A. E. Bogan, E. V. Coan, F. G. Hochberg, W. G.
Lyons, P. M. Mikkelsen, R. J. Neves, C. F. E. Roper, G. Rosenberg, B. Roth,
A. Scheltema, F. G. Thompson, M. Vecchione, & J. D. Williams. 1998.
Common and Scientific Names of Aquatic Invertebrates from the United
States and Canada: Mollusks, 2nd ed. American Fisheries Society, Special
Publication 26, Bethesda, Maryland, 526 pp.

[UCIPM] University of California, Statewide Integrated Pest Management
Project. UC Pest Management Guidelines. Citrus - Brown Garden Snail.
Web Site <http://axp.ipm.ucdavis.edu/PMG/r107500111.html> [update
5/99; last viewed February 2000].

Vanatta, E. G. 1911. Bermuda shells. Proceedings of the Academy of Natural
Sciences of Philadelphia, 62: 664672 [“Dec., 1910”; published January
1911).

Vanatta, E. G. 1912. Bermuda shells. The Nautilus, 26(1): 12.

Vanatta, E. G. 1923. Bermuda shells. The Nautilus, 37(1): 32-33.

Vanatta, E. G. 1924. Land shells of Admiral’s Cave, Bermuda. The Nautilus,
38(1): 6-7 [July].

Vanatta, E. G. 1925. Four new species of shells. Proceedings of the Academy of
Natural Sciences of Philadelphia, 76(1924): 423-425 [February 1925].

van der Schalie, H. 1948. The land and fresh-water mollusks of Puerto Rico.
Miscellaneous Publications, Museum of Zoology, University of Michigan, No.
70: 1-134, pls. 1-14.

Verrill, A. E. 1902. The Bermuda Islands: their scenery, climate, productions,
physiography, natural history, and geology; with sketches of their early history
and the changes due to man. Transactions of the Connecticut Academy of
Arts and Sciences, 11(2) (1901-1902): 413-896, pls. 65-104 [“Introduction of
land snails and slugs,” pp. 727-735; see also Verrill, 1903].

Verrill, A. E. 1903. The Bermuda Islands: lists of figures in the text; explanation
of plates; index; contents. Transactions of the Connecticut Academy of Arts
and Sciences, 11(2): 897-957, v-x [February 1903; teste Ferguson et al., 1991).

Verrill, A. E. 1905-1907. The Bermuda Islands. Part IV. - Geology and
Paleontology, and Part V. - An account of the coral reefs. Transactions of the

December 30, 2000 Bieler & Slapcinsky 99

Connecticut Academy of Arts and Sciences, 12 (1904-1907): 4-92 (June
1905), 93-140 (Nov. 1905), 141-188 (Feb. 1906), 189-236 (Mar. 1906), 237-
284 (Apr. 1906), 285- 300 (June 1906), 301-332 (Mar. 1907), 333-348 (May
1907) 348, pls. 16-40 [“Land shells,” pp. 158-172].

Wagner, A. 1911. Die Familie der Helicinidae [Section 2]. Neue Folge.
Systematisches Conchylien-Cabinet von Martini und Chemnitz, Band 1,
Abtheilung 18, Theil 1, Lieferung 547: 329-391, pls. 67-70 [May].

Wallace, A. R. 1895. Island life; or the phenomena and causes of insular faunas
and floras, including a revision and attempted solution of the problem of
geological climates. Second and revised edition. London, MacMillan and
Co.; xx + 563 pp.

Wiktor, A. 2000. Agriolimacidae (Gastropoda: Pulmonata) - a systematic
monograph. Annales Zoologici, 49(4): 347-590.

Woodley, N. E. & D. J. Hilburn. 1994. The Diptera of Bermuda. Contributions
of the American Entomological Institute, 28(2): 1-64.

Zilch, A. 1959-1960. Gastropoda von Wilhelm Wenz. Teil 2. Euthyneura.
Handbuch der Palaozoologie (O. H. Schindewolf, ed.), 6(2): i-xii, 1-835.
Borntraeger, Berlin [pp. 1-200, 17 July 1959; 201-400, 25 November 1959;
401-600, 30 March 1960; 601-835, i-xii, 15 August 1960].

100 NEMOURIA No. 44

Founding Editor Managing Editor
JOHN E. DU PONT TIMOTHY A. PEARCE

Consulting Editors

M.G. HARASEWYCH, National Museum of Natural History, Smithsonian
Institution, Washington, DC.

KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh, PA.

RICHARD W. THORINGTON, JR., National Museum of Natural History,
Smithsonian Institution, Washington, DC.

Published by the
DELAWARE MUSEUM OF NATURAL HISTORY

P.O. Box 3937, Wilmington,
Delaware 19807-0937, U.S.A.

‘Founding Editor Managing Editor
JOHN E. DU PONT TIMOTHY A. PEARCE

Consulting Editors

M.G. HARASEWYCH, National Museum of Natural History, Smithsonian
Institution, Washington, DC.

KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh, PA.

RICHARD W. THORINGTON, JR., National Museum of Natural History,
Smithsonian Institution, Washington, DC.

Published by the
DELAWARE MUSEUM OF NATURAL HISTORY

P.O. Box 3937, Wilmington,
Delaware 19807-0937, U.S.A.

eS he ee ee

HECKMAN
BIND ER y, INC.
\ Bound-To-Please*

AUG 01

N. MANCHESTER, INDIANA 46962

3 9088 01228 7181

SMITHSONIAN INSTITUTION LIBRARIES

Parle. vw n

DANS CRRA iy,

Bee er ea

axes

SS ny

mm na é

PAKS ha,

Tika Ete ae

Wea 5

aay

Oe Ghee ae

Cea

PADI 9 PUN eneriny -

F Oe
Styaiaeey spa
on Web,
use

*S ee hay Rene
RES BWW Gay

icidat E teen
ay

ors
wBeLSesaveen
hr od ~ ie

<paatesy chee

a eet
PONV aac

Shegra eve Ss

tre
Pesan

PP ge AS

CUE

A

Say neva
ais

Fa
PERERS WG

oY Ri ae

BREMCP snap
RSs F ae

2 vesuiny

SPALL A RAs ong
eee

LOD pay :

PERM Le ay
PEAS

REE Ase

Por ae

RAL PR Dea Gavi: