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Department of Invertebrates

Field Museum of

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Arthur E. Bogan

North Carolina State Museum of

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Philippe Bouchet

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Muséum National d'Histoire Naturelle

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Center for Conservation Research

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Swedish Museum of Natural History

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Delaware Museum of

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CONTENTS

John Slapcinsky

Shugian Zhang

Suping Zhang

Anna C.A. Salles

Cléo D.C. Oliveira

Ricardo S. Absalao

Research Note

Jeremy Tiemann

Clarissa Lawlis

Sarah Douglass

Robert Burn

INGEICE Retr ee es es eee

NAUTILUS

Volume 132, Number 1

April 5, 2018

ISSN 0028-1344

Vitrea clingmani Dall in Pilsbry, 1900, a snail endemic to the summits of

the Black Mountains and Great Craggy Mountains of North Carolina is

now assigned to the genus Pilsbryna (Gastropoda: Stylommatophora:

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Redescription of the jumping snail Ovachlamys fulgens (Gude, 1900)

(Gastropoda: Helicarionoidea: Helicarionidae): An anatomical and

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Flabellina McMurtrie, 1831: The earliest valid Latinization? ..................cc0c000ee 33

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Sponsored in part by the State of

Florida, Department of State,

Division of Cultural Affairs and the

Florida Council on Arts and Culture

CULTURE

BUILDS

FLORIDA

FLORIDA DEPARTMENT of STATE

DIVISION of CULTURAL AFFAIRS

™

THE NAUTILUS 132(1):1-12, 2018

Page |

Vitrea clingmani Dall in Pilsbry,

1900, a snail endemic to the

summits of the Black Mountains and Great Craggy Mountains of

North Carolina is now assigned to the genus Pilsbryna (Gastropoda:

Stylommatophora: Oxychilidae)

John Slapcinsky

Florida Museum of Natural History

University of Florida

Gainesville, FL 32611 USA

[email protected]

ABSTRACT

Collecting in wet leaf-litter microhabitats in the Great Craggy

Mountains and Black Mountains of western North Carolina

uncovered juvenile shells and live adults of the rare snail

Vitrea clingmani Dall in Pilsbry, 1900. This newly available

material Allows the description of the radula, juvenile shell,

and reproductive system of this species, providing characters

fora phylogenetic analysis, which suggests reassignment of

the species from the genus G lyphyalinia (Glyphyalus) to

Pilsbryna. This new placement is corroborated by COI se-

quence data. Surveys near the type locality expand the known

range of Pilsbryna clingmani and suggest this species is en-

demic to spruce-fir aml northern hardwood forests at eleva-

tions above 1500 m in the Black Mountains and contiguous

Great Craggy Mountains of western North Carolina where it is

further restricted to wet leaf litter microhabitats, making it

one of the most narrowly distributed snail species in eastern

North America.

Additional Keywords: Terrestrial snail, Pilsbryna, Appalachian

Mountains, spruce-fir forest, seeps

INTRODUCTION

Pilsbryna clingmani, under the name Glyphyalinia clingmani,

is listed as a species of concern by the U.S. Fish and

Wildlife Service and a threatened species ranked as

critically imperiled in North Carolina (LeGrande et al.,

2008), but very little is known about its range and habitat.

What little is known is based lar gely on reports from two

brief surveys of Mount Mitchell. North Carolina, by ex-

perienced malacologists (Pilsbry, 1900; Walker and

Pilsbry, 1902) which resulted in a handful of specimens

now scattered among several museum collections. This

scarcity of records suggests the species is not widely

distributed or may require or prefer microhabitats not

commonly or easily sample -d. In the past century the

forests of Mount Mitchell have undergone profound

change and during that time Pilsbryna clingmani has been

reported only once (Hubricht, 1970).

Non-marine mollusks have the unfortunate distinction

of having the highest number of recent extinctions

documented in any animal group (Régnier et al., 2009:

Cowie et al., 2017). Many of these were extinctions of

terrestrial snails endemic to islands but snails restricted to

other geographically isolated habitats are vulnerable to

habitat loss that might lead to extinction. Several of

eastern North America’s most narrowly distributed snail

species (Hubricht, 1985) occur in southern spruce-fir

forest (Pilsbry, 1940; 1946; 1948). This community is

dominated by c canopy species Abies fraseri (Pursh, 1814)

and Picea rubens Sargent, 1898, and is limited to seven

disjunct stands at peaks above 1500 m elevation in the

southern Appalachian Mountains. Pilsbryna clingmani is

known from one of these stands of southern spruce-fir

forest at the summits of the Black Mountains (Walker and

Pilsbry, 1902). All published records were based on

specimens collected prior to 1902 within 5 km of Mount

Mitchell at elevations above 1900 m. Shortly afterwards

approximately 50-65% of the spruce-fir forest on the

Black Mountains was logged between 1912 and 1927

(Pyle and Schafale, 1988). Loss of forest cover prompted

reforestation efforts, which unfortunate ly included non-

native trees (Minckler, 1940). These non-native species

are probably the source of the catastrophic introduction of

the balsam woolly adelgid, Adelges piceae (Ratzeburg,

1844), which was first reported on Mount Mitchell in

1957 (Speers, 1958). Nearly all mature fir trees in the

Black Mountains had succumbed to A. piceae by 1988

(Dull et al., 1988), opening large portions of the forme tly

dense canopy and exposing re al spruce, Picea rubens, to

damage similar to thinning mortality (Nicholas et al.,

1992). Acid mists eoeted with air Bees reported

from Mount Mitchell (DeFelice, 1997) probably cause

additional reduction in growth of red spruce by lowering

availability of calcium (Jacobson et al., 1990). The more

open canopy probably produces changes in soil moisture

>. )

Page 2

and leaf litter cover that may negatively impact moisture-

dependent land-snails.

Recent land snail surveys in wet leaf litter micro-

habitats in the Black Mountains and contiguous Great

Craggy Mountains in western North Caroline found

specimens of Pilsbryna clingmani. Rediscovery of this

species permits its redescription, comparison with

other species, reassignment to the genus Pilsbryna,

and clarification of its geographic range and habitat

requirements.

MATERIALS AND METHODS

Twenty-nine survey sites that were likely to contain snails

were chosen along easily accessible roadways and trails in

the Black Mountains and Craggy Mountains near Mount

Mitchell, with special attention given to seeps in both

open and forested habitats. An additional 41 sites were

sampled in other mountain ranges in western North

Carolina. Specimens were hand-collected from the leaf-

litter soil interface during 1 hour searches within sites of

approximately 100 m> - Aaldktioreall specimens were picked

from approximately 1 liter samples of leaf-litter sievings

gathered from each site. Live collected animals were

mereoted overnight in a suspension of water containing

a 1 cm length of mentholated cigarette and then preserved

in 75% ethanol. Gross anatomical dissections were made

under 75% ethanol using a dissecting microscope. Iso-

lated reproductive systems were stained with Harris’

haematoxylin and Semichon’s aceto-carmine stain, cleared

with glycerin, and mounted on slides. Drawings of the shell

and reproductive anatomy were made with the assistance

of a camera lucida, and measurements taken using an

ocular micrometer. Greater and lesser shell widths, shell

height, aperture width, aperture height, umbilical width,

spire width, and number of whorls were measured as

described by Slapcinsky and Coles (2004) and shown in

Figures 1— 3. Means and standard deviations are reported

Fan all measurements of 14 undamaged adult specimens.

Shells with rapidly expanding body whorls and 4.9 or more

THE NAUTILUS, Vol. 132, No. 1

whorls were presumed to be mature and two were

dissected and confirmed to have adult reproductive

structures. Radulae were isolated from dissected buccal

masses using a saturated KOH solution. Scanning elec-

tron micrographs of shells and radulae were made using

a HITACHI S-4000 FE-SEM. All available specimens a

Pilsbryna clingmani in the collections of the Academy of

Natural Sciences, Philadelphia (ANSP); Carnegie Museum

of Natural History, Pittsburg (CMNH); Field Museum of

Natural History, Chicago (F MNH); National Museum of

Natural History (USNM): Schiele Museum of Natural

History, Gastonia, North Carolina (SMNC); University of

Florida, Florida Museum of Natural History, Gainesville

(UF); and in the private collections of Harry G. Lee,

Jacksonville, Florida (HGL) and Amy Van Devender (ASV)

were examined. Anatomical terminology follows Tompa

(1984). Higher-level systematics follows Bouchet et al.

(2017).

Morphological characters derived from dissections

combined with those gleaned from published figures

of reproductive systems, radulae, and shells (Baker,

1929a, b; Pilsbry, 1946; Slapcinsky and Coles, 2004)

were used to build a matrix of 28 characters (Table 1)

for 47 oxychilid species (Table 2). Characters were treated

as unordered. A phylogenetic analysis was performed

using the optimality criterion of parsimony in Mesquite

version 3.2 (Maddison and Maddison, 2017).

DNA was extracted from 1 mm?® samples of foot tissue

using a solution of 10% Chelex beads (Bio-Rad Labora-

tories) in sterile water heated to 65°C for 4 hours. DNAses

and other protein contaminants were digested using

Proteinase K. A 655 bp nucleotide fragment of cyto-

chrome oxidase I (COI) was amplified using the primer

pairs LCO1490/HCO2198 (Folmer et all, 1994) and

GoTaq DNA Polymerase (Promega, Madison, Wisconsin).

Sequencing was performed in both directions at the

University of Florida, Interdisciplinary Center for

Biotechnology Research or the Canadian Centre for

DNA Barcoding (CCDB), using their standard pro-

cedures. Primer regions were acted and sequences

edited and aligned using Geneious version 6.1.7 (Kearse

1 AE Z

phone 1-3.

Shell measurements. 1. Spire width (line a-b); whorl count (line 14.9). 2. Shell height (line c—d); aperture height (line

e-f); aperture width (line g—f). 3. Greater width (line h-i); lesser width (line i-j); umbilical width dine k-l).

J. Slapcinsky, 2018

Die sys

Page 3

Table 1. Morphological characters of the shell, radula, and

reproductive system used in the phylogenetic analysis.

1. shell size: 0=large, >12 mm; 1=small, <12 mm

2. whorls: 0=<3.5; 1=3.8-4.2: 2=4.5-5.5; 3=>6

3. umbilicus: 0= umbilicate; 1= perforate/imperforate

4. axial grooves: 0=absent; 1=close and irregular; 2 distant and

regular

5. body whorl: 0=evenly expanding; 1=body whorl flared

6. parietal lamella: 0=absent: |=present

7. columellar lamellae: 0=absent; 1=present

8. subbasal lamellae: 0=absent; 1=present

9. basal lamellae: O=absent; 1=present

10. subpalatal lamellae: O=absent; 1=present

11. palatal lamellae: 0=absent; 1=present

12. suprapalatal lamellae: O=absent; 1=present

13. sutural lamellae: 0=absent; 1=present

14. axial fusion: 0O=absent; 1=present, ?=no lamellae

15. spiral fusion: 0=absent; 1=present, ?=no lamellae

16. lamellae axial spacing: 0=not widely spaced; 1=spaced

1/4-1/2 whorl: 2=no lamellae

17. periuterine gland: 0=absent; 1=present

18. perivaginal gland: 0=absent; 1=present

19. vas deferens: 0=long; 1 short

20. epiphallar caecum: 0=absent; 1=present

21. epiphallar flagellum: O=absent; 1=present

22. epiphallar insertion: 0=apical; 1=subapical

23. oval subapical penial pilasters: 0=absent; 1=present

24. central mesocones: 0=attenuate; 1=diamond shaped

25. lateral teeth: 0=tricuspid; 1=bicuspid

26. marginal teeth: 0=smooth; 1=serrate

27. jaw cusps: 0=three cusps; 1=four or five cusps

28. spacing of jaw cusps: 0=cusps evenly spread; 1=cusps

central

et al., 2012) and deposited in GenBank under acces-

sion numbers MG648722-MG648731. Phylogenetic anal-

ysis using maximum likelihood and the GTR +1+G

model ae evolution was performed in MEGA6 (Tamura

et al., 2013).

SYSTEMATICS

Family Oxychilidae Hesse in Geyer, 1927

Family Placement: Traditionally, most North Ameri-

can limacoid species have been placed in a morphologi-

cally heterogeneous Zonitidae (Baker, 1931; Pilsbry, 1946;

Riedel, 1980). A cladistic analysis of Limacoidea using

reproductive, muscular, nervous, lung, digestive, edemal

body, and shell characters cast doubt on nine monophyly

of Zonitidae (Hausdorf, 1998) and elevated most of its

component subfamilies. Paravitrea and its then syno-

nym Pilsbryna (see Riedel, 1980), were excluded

from Vitrininae, now Pristilomatidae, and moved to

Daudebardiidae Kobelt, 1906. The genera Glyphyalinia,

Nesovitrea, Mesomphix, and Vitrinizonites were also

placed in Dauderbardiidae. More recently, Daudeberdiinae

has been treated as a subfamily within Oxychilidae

(Bouchet et al., 2017). However, family level relation-

ships of taxa formerly treated as Zonitidae are not fully

resolved and remain controversial (Schileyko, 2003,

Welter-Schultes, 2012).

Genus Pilsbryna Baker, 1929

Type Species: — Pilsbryna aurea Baker, 1929 by monotypy.

Pilsbryna clingmani (Dall in Pilsbry, 1900) new

combination

(Figures 4-18)

Zonites wheatleyi— Binney, 1885: 223. [not Zonites wheatleyi

Bland, 1883].

Viitrea]. clingmani Dall, 1898: 100-101 [nomen nudum

as a variety of Zonites wheatley/i].

Vitrea clingmani Dall in Pilsbry, 1900: 149-150, Figure 2;

vyalleee and Pilsbry, 1902: 431, pl. 24, Figures 1 3.

Vitrea approxima W alker and Pilsbry, 1902: 431-432, pl.

24, Figures 7-9.

Retinella clingmani.—Pilsbry, 1946: 275-276, Figure 136a.

Retinella approxima. —Pilsbry, 1946: 276-277, Figure 137.

Glyphyalinia clingmani. —Hubricht, 1970: 13: Hubricht,

1974: 34 [synonymizes Retinella approxima with

Glyphyalinia clingmani|; Hubricht, 1985: 23.

Diagnosis: A large Pilsbryna with an adult shell of

4.9-5.4 whorls that is 5.0-7.8 mm in width and

2.6-3.9 mm in height. Early whorls increase slowly and

regularly in size but whorl expansion increases in the body

salar of the adult. Shells with fewer than 2.5 whorls

contain a sinuous parietal lamella and two spirally

arranged series of basal and palatal nodules that can ex-

tend a full whorl into the aperture.

Description: — Body coloration dark blue-grey (Figure 4).

Shell is depressed- helicoid, umbilicate, glossy, faneincent

brown, and sculptured with dense and irregularly spaced,

indented axial lines that become more fre quent on the last

quarter of the body whorl of the adult (Figures 5—8). Adult

shell (n= 14) 5.0-7.8 mm (mean=6.2+0. 9 mm) in greater

width, 4.1-6.3 mm (mean=5.0+0.7) in lesser sarki and

2.6-3.9 mm (mean=3.1+£0.5 mm) in height with 4.9-5.4

(mean = 5.1+0.1) slowly expanding eee The final 1/3 of

the Tyadhy ine of the adult expands more rapidly than

previous whorls; the lesser/greater width ratio is 0.80—0.87

(mean=0.85). The arlene also expands rapidly in the

body whorl of the adult, reaching a width of 0.7 to 1.0 mm

(mean = = 0.9+0.1). Spire width is 2.4-3.9 mm (mean=3.0+

0.4). The aperture is ovate; aperture width is 2.74.0 mm

(mean=3.2=+0.5), aperture height is 2.1—3.0 mm

(mean=2.5+0.3). Shells of immature animals with fewer

than 2.5 whorls have a sinuous parietal lamella and a series

of paired basal and palatal nodules (Figures 9-10). All

traces of nodules and lamellae are resorbed in adults.

Reproductive Anatomy: Two specimens were exam-

ined (UF 292689). The basal half of the penis is smooth

while the apical half bears numerous small thorn shaped

papillae (Figure 11). The penial retractor muscle is apically

Page 4

THE NAUTILUS, Vol. 132, No. 1

Table 2. Matrix of 28 characters and 47 species used in the morphological analysis (see Table 1 for explanation of characters).

Characters

Taxa Les 45 6 7 & OF IO Il 1 IS 14 15 16 l7 1B IG B Vl BD 98 94 95 9B O97 OB

Glyphalinia burringtoni I oO@di@ogo@od@o@oq@goqogoe@ore?rrerrpriliil i @oo@onhii@o®P?

Glyphalinia carolinensis t2?it2?@ooood®oe®ogo@o@orPperrr?Pri dt gi @oioii op? ® ®P

Glyphalinia cryptomphala litrk2?oo@o oo oo oqo Pp PPTL id Orto@oi@o Pp? Be?

Chynwiveacmwonmmnme lo O20 0 0 00 0 OM © ® PP Pt_LtIt® OOO iO O P ®

Glyphalinia indentata i @®@il ®¥oeoeqodoeoqodoqo@orprrperpiadi?ep@oioii@i®»?®

Glyphalinia lewisiana l@®@@oitrogo®o®odogoogo®orprpr? P ri i@oogodiinii ink oOo P B

Glyphalinia paucilirata lr@OirtZzgqooooooe@oo@oorprprPpii_ itil @Oiaoigdi @i

Glyphalinia pentadelphia Ll@o@oirogoodod@ododooo®drprrprrPprirhi i GOOO®O i ©O O BP?

Glyphalinia praecox liriigoegododdododgodo@o@orpprpriiinh P@OIt Oi @goi®@ yj

Glyphalinia rhoadsi l@®O@o2oOooo@oododoo@o rprPpPiiia @o@gQgogonriogdao @

Glyphalinia roemeri roo?zxoo®eddododo@odo@o®orrprrpi i i @g@oii@oin ii @ 2 @

Glyphalinia sculptilis lZit2z2qooqoQqoqogoooooogrrpepiraii@oi@onk@d i ® ?P

Glyphalinia solida lii%@Oogododo@od@o@o@orpr?P? PL dt ik @Oilo@oigd? oO ii

Glyphalinia wheatleyi tt @2 09 OOOO OO OO P PPT ik@oogod®oi@od@o ®? >?

Mesomphix andrewsae Olt @QgQ@oodoododd@odovo@og®edr?rrPrPOHOOD®”A DiI ODO OOD P ?

Mesomphix capnodes Ol OO @®@OdO®*OOOOOO®MD P P POit@oooi@ooo oO DP PB

Mesomphix cupreus Ologogqooqoqogoo®oedod@o@orrprrd@®”il@gdogoiikhoo@oq@d P ®P

Mesomphix friabilis Ol too Od@OOODOOOAOOAP P POlOOOD i OO O O P P

Mesomphix inornatus Oli @@gdgoaegog®eg®o*ddsvs@O@Oo@orprr?Prr»?Pg*oew®eogedoii@®m@o@o@o 2 B

Mesomphix latior Ol itl Oooo ®ooo@o@ogorrppeoegoedeoedgedi @@o@e@od P OP

Mesomphix perlaevis Oil hk OD@*dgaogodgdgg@gdgedogo@orre@oaedda@adeoOoii@®dO®O®O QO @

Mesomphix pilsbryi OrodDgDOODOADYVOOADOA ODO P PPOokOOMl OOM OO P ®

Mesomphix rugeli Otil ®©9@OOO ODD OO®*™ PP P POODDAO i OOO @® PR P

Mesomphix subplanus 0o?Itl@Oooo ooo Ooo Pp P POO OOHOl OO O MO P P

Mesomphix vulgatus Oli O9OOODDAODHOADAO P P POW OHO il ®© OOO P FP

Nesovitrea binneyana l@®@@oiogodgogdgdgoq@ogo@oorprrprpri@®i@®@o@oiiii © P P

Nesovitrea electrina l@®@@oi@ogoe@o@ododd@odg®?@®o@or»?perprRi@il@ogog@oii ii i @ PB P

Oxychilus cellarius ii @O@@og@od@oggod@o®eoqog@o@d@orpeeperrPil@oegood®od@or? @®iio Pp P

Paravitrea andrewsae l%@oi@ogo@dogih ii gtrkrig@oggo@goitk@og@oiih@oitiogi i @O i

Paravitrea calcicola t?@di@ogodg@og@oi @mair@ogoeg@goeoiiil@ogoi @oi#krte@od@oiinii@ P P

Paravitrea clappi l2@goitodogoiii#ida#dri@ggo@oi @®@@@di@og@o@oh ii @ P P

Paravitrea lamellidens l?@Oitogooiinttltiili@oii@gdi@doiig@®oo@oq@ogoiii @ii dj

Paravitrea multidentata Ll2?@qoigogogoirtiadtildri@mgig@oinr@oaqoi@goo®o@o i id O P P

Paravitrea petrophila l?oi#rhe@og@o@ood@o®o@o@orperprpere@madgmi@O@oo oi ih@o Pe?

Paravitrea pilsbryana i? @oit@@ood@odik @gH@ooOogdeoaiIt Oot Ot @®@oOi itl @® PF

Paravitrea placentula l?@ore@ooo®o@®o#hhoinh@godg@qooqoin@ogoiiog®oooi iio Pe?

Paravitrea pontis l?@dint@oeo®od@odd@oi@oiiklh@g@m®@odoeiung@od@oigog®go@oiitl@o Pp P

Paravitrea tridens r2@oir@ooi@oiilheigqogrgdggoogoikgqgooit @®mi ©@@® i hk O DP

Paravitrea variabilis r?2q@oaitoeodgo#it ils kkti @odoigoigogoiogog®dd@ i it@ Pe? P

Paravitrea walkeri l2@miogdoirtikhiiktkhigdgi@dai@oegoihgogaogeo@oiii © P P

Pilsbryna aurea ri@oirtiiaiik@o#q@ooegododoegoilheg@oog@oii@ogdgogoii ii © P P

Pilsbryna castanea Hi @mgwxtikt O@O@kr@OOgOQgODaODAk ODA ODL OA OOOH it I Ol d

Pilsbryna clingmani lt i @ it it dogo O@il @ogdo#rhgqgegqogokhd@oe@ogoihri @r Pp

Pilsbryna nodopalma ri Ori lk@ooirg@oreriq@oao@oonhkh@dogoirit@oedogo@gonh ii @® P P

Pilsbryna quadrilamellata li @®@ititii @Oir @@M Oi @otegqogoo®oiik @o oo ii i @O P P

Pilsbryna vanattai rir@ogi?rig#rroge@gdgtrh@odi@oo@oitegogooiti@ooogoiini@d ? Pp

Vitrinizonites latissimus oO il OOOO DOdOOOOdODAOP P PHOoOHODOH OO OO PP?

inserted. The epiphallus is about the same length and Radula: Two specimens were examined (UF 292689).

roughly 2/3 the diameter of the penis and inserts laterally at

the apex of the penis. The epiphallus is sculptured with

longitudinal folds and tapers slightly before widening at the

junction with the narrow vas deterens. The spermatheca i is

ovate, and the narrow spermathecal duct expands slightly

basally. No sign of perioviducal or perivaginal glands were

observed. While most characters varied little in size, the

albumen gland and oviduct were larger in one of the two

specimens that were dissected.

The central tooth of the radula is symetrically tricuspid,

the mesocone expands slightly above the ectocones, then

tapers slowly to the apex (Figure 12). The ectocones are

short and sy mmetrical, less than % the total height of the

tooth. The three laterals are tall, slender, and asymmet-

rically tricuspid and slightly convex proximally and con-

cave distally. The endocone is tall, about % the height of

the mesocone while the ectocone is less than 4% the height

of the mesocone. The marginals are tall, slender, emooths

J. Slapcinsky, 2018

Page 5

Figure 4.

Pilsbryna clingmani (Dall in Pilsbry, 1900). Live

individual from Mount Mairehelll Yancey County, North Caro-

lina. Photographed by R. Wayne Van Devender.

and unicuspid, with concave peripheral edges and convex

distal edges.

Holotype:

locality.

USNM 38910 (Figures 13-15), from type

Type Locality: USA, North Carolina, Yancey County,

Black Mountains near Clingman’s Peak, coll. H. Hemphill,

date unknown but hetere the publication of Binney

(1885).

Other Material Examined: USA, North Carolina,

Yancey County, Clingman’s Peak, coll. J]. Ferriss, May

1901: ANSP 84060, 7 specimen; CM 73964, January

1905 (month and day reversed on label, probably May

1901), 1 specimen; FMNH 58325, 4 specimens. USA,

North Carolina, Yancey County, Mount Mitchell: CM

73963, coll. H. Hemphill, November 1895, 1 specimen,

identified as Retinella approxima on the label; FMNH

240953, coll. L. Hubricht, 26 May 1962, 9 specimens;

FMNH 240954, coll. L. Hubricht, 26 May 1962, 1

specimen. USA, North Carolina, Yancey County, Mount

Mitchell State Park: near summit parking area, ASV-

2007-184, 35.7665° N, 82.2644° W, 2001 m, coll. A. and

W. Van Devender, 3 November 2007, 1 specimen; near

family camping area, ASV-2008-14, 35.7599° N,

82.2711° W, 1920 m, coll. A. and W. Van Devender, 3

November 2007, 1 specimen; near campground, ASV-

2011-99, 35.7600° N, 82.2709° W, 1914 m, coll. A

Gettleman and Denise Furr, 1 October 2011, 2 specimens;

SMNC MAL666-01, 35.760701° N, 82.270285° W, 1915 m,

coll. Denise Furr, 1 specimen. USA, North Carolina,

Yancey County, Mount Mitchell, Wilson’s Cove, ANSP

83260, coll. B. Walker, 1901, Lectotype (Baker,

Vitrea approxima Walker and Pilsbry, 1902 (Figures

16-18). USA, North Ce Yancey County, Potato

Top: ANSP 83262, coll. J. Ferriss, 8 July 1902, 5

specimens; UF 107824, call J. Ferriss, 1901, 1 specimen.

USA, North Carolina, Buncombe C ounty, Bear Pen Gap

1962) of

near Craggy Gardens Picnic Area, 35.701° N, 82.391° W,

coll. J. Slapcinsky: UF 292689, 29 May 2001, 6 specimens:

UF 299560, 12 Septe »mber 2002, 3 specimens; UF 341346,

19 June 2004, 1 specimen; UF 347735, 21 August 2004, 5

specimens.

Remarks: Binney (1885) identified Hemphill’s speci-

men from Clingham’s [sic] Peak as Zonites wheatleyi,

which is treated as a species of Glyphyalinia (Hubricht,

1985). Dall (1898) gave the varietal name Vitrea clingmani

to Hemphill’s shell but did not provide a figure or de-

scription. Pilsbry (1900) indicated that Dall’s new name

was a nomen nudum and included a description, a refer-

ence to a type lot, USNM 38910, and a figure, all of which

were provided by Dall. In addition to the holotype, which

is consistent with Dall’s description, measurements, and

figure (Pilsbry, 1900, Figure 2), lot USNM 38910 also

emiteinadl a specimen of Paravitrea petrophila (Bland,

1883). This is probably the specimen Dall had compared

to his new species in the original description and this shell

is now USNM 38910a.

Synonymy of Vitrea approxima Walker and Pilsbry,

1902: Vitrea approxima was described based on two

specimens collected by James Ferris and Bryant Walker

in Wilsons Goveon Mount Mitchell (Walker and Pilsbry,

1902) and does not appear to have been collected since.

One of these specimens (Figures 16-18) was deposited

by Walker (ANSP 83260) and selected as the lectotype

by Baker (1962) based on original measurements, but

cited incorrectly as ANSP 82360. The fate of the second

specimen is unknown. Vitrea approxima differs from

Pilsbryna_ clingmani, which is larger, and Pilsbryna

vanattai, which is smaller, by having a relatively small

umbilicus and lacking a rapidly expanding body whorl.

Hubricht (1974) synonymized Vitrea approxima with

Pilsbryna clingmani (as Glyphyalinia), suggesting unique

attributes of “the lectotype were the result of shell

damage early in life. While the lectotype (Figures 16-18)

has shell damage between the second and third whorls, it

does not appear to be seriously deformed. It is a juvenile

of only 4.6 whorls that has not reached the stage of rapid

body whorl expansion that would result in a fared body

whorl and relatively large umbilicus that is common in

adults of all Pilsbryna species (Slapeinsky and Coles,

2004) and it appears to be a juvenile specimen of Pils-

bryna clingmani.

Habitat and Distribution: All previous records of

Pilsbryna clingmani were from spruce-fir forest above

1800 m elevation on the peaks of the Black Mountains

near Mount Mitchell, North Carolina (Walker and Pilsbry,

1902). My surveys of the Craggy Mountains (Figure 19)

extend the range of P. dlinenvan south to near Craggy

Gardens picnic area. This also extends the ele eatonal

range of the species down to 1500 m and into northern

hardwood forest. Like other Pilsbryna species, P. clingmani

is common in wet leaf-litter in seep runs and ditches on wet

soil or under wood. However, at higher elevation sites on

Mount Mitchell, it is also common among grasses and

Page 6 THE NAUTILUS, Vol. 132, No. 1

Figures 5-12. Pilsbryna clingmani (Dall in Pilsbry, 1900). 5-7. photographs of shell SMNC MAL666-01, width 6.2 mm, photo-

graphed by R. Wayne Van Devender. 8. Scanning electron micrograph of apical view of adult shell UF 299560, width 6.3 mm. 9.

Scanning electron micrograph of apertural view of juvenile shell of 2.5 whorls with body whorl broken to expose internal lamellae, UF

360436, width 1.4 mm. 10. Line drawing of juvenile shell of 2.9 whorls, UF 360436, width 1.6 mm. 11. Camera lucida drawing of

genitalia, UF 292689, scale bar = 5.0 mm. 12. Scanning electron micrograph of radula, UF 292689, scale bar = 50 pm. Abbreviations:

AG = albumen gland, EP = epiphallus, HD = hermaphroditic duct, OV = free oviduct, PE = penis, PG = prostate gland, PP = penial

papillae, PR = penial retractor muscle, BC = bursa copulatrix, UT = uterus, VD = vas deferens.

J. Slapcinsky, 2018

Page 7

Figures 13-18. Pilsbryna clingmani (Dall in Pilsbry, 1900).

Shells of type material. 13-15. USNM 38910, Holotype of Vitrea

clingmani Dall in Pilsbry, 1900, tata 6.3 mm, photographed by Paige Billin-Frye. 16-18. ANSP 83260, Lectotype of Vitrea approxima

W alee and Pilsbry, 1902, width 4.3 mm, photographed by Paul C Slloyinern.

leaves in open areas (Amy Van Devender, W ayne Van

Devender, and Denise Furr, personal communications).

Comparative Remarks: Placement of Vitrea cling-

mani within Pilsbryna is supported by a phylogenetic

analysis of 28 morphological characters from 47 species of

Oxychilidae ( Figure 20). Pilsbryna clingmani has a series

of pariet tal aeratulles that are fused into a parietal lamella

which is resorbed in adults and has a shell of approxi-

mately 5 whorls that expands slowly at first and more

rapidly in the body whorl of adults: these are synapo-

morphies that unite Pilsbryna species within Oxychilidae.

Juvenile Pilsbryna clingmani shells bear basal and parietal

nodules found in Pilsbryna and Paravitrea but not in

other oxychilids.

Pilsbryna clingmani does not share certain characters

found in other cull shelled oxychilid genera. The la-

mellae of Pilsbryna are not grouped axially and separated

by % to “3; whorl as in Paravitrea. Glyphyalinia sensu

stricto have shells textured with widely and regularly

spaced axial indentations, reproductive systems woitin per-

ivaginal and perioviducal glands, and radulae with serrated

marginal teeth and bicuspid lateral teeth; none of these

characters is found in Pilsbryna clingmani. Nesovitrea and

some Glyphyalinia species have penises with subapical

oval pilasters and shells with approximately 4 whorls that

expand rapidly and consistently unlike Pilsbryna which

have apical thorn shaped papillae and shells of 5 whorls only

the last of which expands rapidly.

Both morphological (Figure 20) and molecular data

(Figure 21) support placeme nt of Pilsbryna clingmani

within Pilsbryna. Pilsbryna clingmani has a spiral row of

nodules at basal and palatal positions in addition to the

sinuous parietal lamella that defines all described Pils-

bryna. The basal nodules are not fused into a lamella as

in P. quadrilamellata. Unlike Pilsbryna clingmani, pala-

tal nodules are lacking in P. castanea, P. aurea, and

P. quadrilamellata. Of the species with palatal nodules, P.

nodopalma has flattened blade shaped basal and palatal

nodules unlike the peg-shaped nodules in P. clingmani,

which are similar to those of the much smaller P. vannattai.

DISCUSSION

My survey is the first to report Pilsbryna clingmani from

northern-hardwood forest in the C raggy Mount uins, in-

dicating that the species is not restricted to the Black

Mountains or to spruce-fir forest. However, this species

was only observed at elevations above 1500 m and was not

found in well-sampled cove-forest at lower elevations in

the same mountain ranges, where it is replaced by P.

vanattai (Slapcinsky aunel Calle. 2001). Furthermore, the

species was not found in 41 sites sampled outside of the

Black Mountains and Craggy Mountains and there are no

other records of the species outside of these mountain

ranges (Hubricht, 1985). The majority of the distribution

of P. clingmani occurs in spruce-fir forest which has ex-

perie neal high mortality as a result of 60 years of adelgid

impact (Dull et al., 1988). However, P. clingmani is still

commonly encountered in damaged spruce kr forest near

the summits of the Black Mountains and it appears that

the paucity of observations of this species in the past

century are the result of low sampling effort. eeu

P. clingmani is common among grass and leaf litter i

open areas near the summit of Mount Mitchell (see

above). These exposed habitats are normally too dry for

Pilsbryna species (Slapcinsky and Coles, 2004). Tt is

possible that high elevation mist on Mount Mitchell

maintains soil moisture that is necessary for this Pilsbryna

species even in open habitats. Adc litional sampling is

necessary to determine if P. clingmani also inhabits other

high elevation open habitats such as heath balds. The

occurrence of this species in acidic leaf-litter with high

Page 8

“Ca rolina 50.

R “Rotite 197,

raggy

rope

THE NAUTILUS, Vol. 132, No. 1

\ 1500 m

contour. «

se eee noe tae L

Cattail -

@ Peak

Blue Ridge Parkway

coy

Figure 19. Pilsbryna clingmani (Dall in Pilsbry, 1900). Species distribution in the Great Craggy and Black Mountains, western North

Carolina. Positive sites denoted by solid dots and negative sites by unfilled dots. 1500 m contour indicated.

rates of soil leaching on acid igneous rocks in conifer forest

and open habitats suggests it is an acidophile (Nekola,

2010). Ecological data are needed to determine the

moisture, cover, and microhabitat requirements of this

rare species. This is especially important because global

warming is likely to reduce isolated high elevation spruce-

fir and nomhenie hardwood forest esl Leet F in the southern

Appalachian Mountains. The narrow distribution of P.

clingmani in wet microhabitats above 1500 m on the peaks

of ‘the Great Craggy and Black Mountains make this

species especially vulnerable and a species of special

concern to conservation biologists.

Oxychilidae in eastern North America comprises six

genera: the large shelled Mesomphix and Vitrinizonites

J. Slapcinsky, 2018

Page

Figure 20.

system. Synapomorphies are marked in bold on branches.

that are usually > 20 mm in shell width and the much

smaller Glyphyalinia, Nesovitrea, Paravitrea, and Pils-

bryna, which are usually < 10 mm in shell width. Unlike

most of the other genera, Pilsbryna is morphologically

well defined, composed of morphologically similar species

with similar habitats in reliably moist leaf litter micro-

habitats. The other small bodied genera Glyphyalinia,

Nesovitrea and Paravitrea are morphologically and eco-

logically diverse and probably require taxonomic revision

using additional morphological and molecular characters.

Oxychilus cellarius

Mesomphix capnodes

Mesomphix pilsbryi

Mesomphix cupreus

Mesomphix friabilis

Mesomphix inornatus

Mesomphix latior

Mesomphix perlaevis

Mesomphix rugeli

Mesomphix subplanus

Mesomphix vulgatus

Mesomphix andrewsae

Vitrinizonites latissimus

Glyphyalinia wheatley!

Glyphyalinia cumberlandiana

Glyphyalinia rhoadsi

Glyphyalinia roemeri

Glyphyalinia carolinensis

Glyphyalinia sculptilis

Glyphyalinia paucilirata

Glyphyalinia indentata

Glyphyalinia praecox

Glyphyalinia cryptomphala

Glyphyalinia solida

Glyphyalinia pentadelphia

Glyphyalinia lewisiana

Glyphyalinia burrington!

Nesovitrea electrina

Nesovitrea binneyana

Paravitrea petrophila

Pilsbryna aurea

Pilsbryna quadrilamellata

Pilsbryna castanea

Pilsbryna nodopalma

Pilsbryna clingmani

Pilsbryana vanattal

Paravitrea placentula

Paravitrea pontis

Paravitrea pilsbryana

Paravitrea calcicola

Paravitrea tridens

Paravitrea andrewsae

Paravitrea Clappi

Paravitrea multidentata

Paravitrea variabilis

Paravitrea lamellidens

Paravitrea walker!

ACKNOWLEDGMENTS

I thank the U

Majority rule consensus tree of 100 most parsimonious trees based on 28 characters from shell, radula and reproductive

nited States Forest Service for permission to

work on their lands. North Carolina Wildlife Resources

Commission provided Scientific Wildlife Collection Licen-

ses. Sandy Florence, Pisgah-Nantahala National Forest,

ge ‘nerously shared knowle dias of the lands in her care. Paige

Billin-Frye and Paul @allonnon photographed type material.

Gary Rosenberg and Paul Callomon (ANSP),

Robert

Hershler (USNM) ), Jochen Gerber (FMNH) and Tim Pearce

Page 10

THE NAUTILUS, Vol. 132, No. 1

0.68

0.50

0.75

0.99

0.75

0.58

0.80

0.04

Mesomphix rugeli

Mesomphix subplanus

Nesovitrea electrina

Nesovitrea binneyana

Glyphyalinia indentata

Glyphyalinia carolinensis

Glyphyalinia solida

Glyphyalinia

Glyphyalinia praecox

Glyphyalinia sculptilis

Pil na aurea

0.75 sbryna

1 Pilsbryna quadrilamellata

0.98

Pilsbryna castanea

Pilsb dopal

aes ilsbryna nodopalma

0.50 1

Pilsbryna clingmani

0.76 1

Pilsbryna vanattal

Paravitrea

0.98 :

Paravitrea lamellidens

Paravitrea calcicola

0.64

0.85 Paravitrea multidentata

0.8 Paravitrea clappi

Figure 21. Phylogenetic tree based on maximum likelihood analysis of COI sequence data with node support based on 100 bootstrap

replicates.

(CMNH) lent specimens or facilitated visits to collections in unpublished habitat data and photographs from their surveys

their care. Harry G. Lee and Wayne and Amy Van Devender of Mount Mitchell State Park and provided comments on

loaned material from their private collections. Amy and earlier drafts of this paper. Further improvements were

Wayne Van Devender, and Denise Furr kindly shared suggested by reviewers Robert Cowie and Jeffrey Nekola.

J. Slapcinsky, 2018

age

Page 11

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Notice

THE 2018 R. TUCKER ABBOTT VISITING CURATORSHIP

The Bailey-Matthews National Shell Museum is pleased to invite applications for the 2018 R. Tucker Abbott Visiting

C uratorship. The Curatorship, established originally in accordance with the wishes of the late Dr. R. Tucker Abbott.

Founding Director of the Shell Museum, is awar ded annually to enable malacologists to visit the museum for a period of

one week. Abbott Fellows are expected, by performing collection-based rese arch, to assist with the curation of portions of

the Museum’s collection and to provide one talk for the general public. The Museum collection consists of marine,

freshwater, and terrestrial specimens. The entire collection has been digitally catalogued and the catalogue is available at

the Museum web site at https:/Avww.shellmuseum.org/malacological-collection and via iDigBio at http://Apt.idigbio.org/

resource ?r=bmnsm-shell. The R. Tucker Abbott Visiting Curatorship is accompanied by a stipend of $1,500.

Interested? Send a copy of your curriculum vitae, a letter detailing your areas of taxonomic expertise and research

objectives, a tentative subject for your talk to:

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THE NAUTILUS 132(1):13-18, 2018

Page 13

Two deep-sea Calliotropis species (Gastropoda: Calliotropidae)

from the western Pacific, with the description of a new species

Shugian Zhang

Suping Zhang

Institute of Oceanology

Chinese Acade my of Sadie neces

Qingdao, 266071 CHINA

[email protected]

museum @qdio.ac.cn

ABSTRACT

Examinations of materials deposited in the Marine Biological

Museum, Chinese Academy of Sciences (MBMCAS) revealed

two deep-sea calliotropid species. Calliotropis infundibulum

(Watson, 1879), trawled from an abyssal plain in the South China

Sea (17°15’ N, 111°20' E) ata depth of 1238 m, is anew record of

the species for Chinese waters. Calliotropis yapensis new ae

cies, collected at the Yap Seamount (8°55’ N, 137°48' E),

a depth of 1119 m, is distinguished from its congeners by ee

a globular shell (height/width ratio=0.95), sculpture d with

granular spiral cords an prominent, densely spaced axial ribs.

Phyloge netic analysis using 633 bp of the cytochrome oxidase c

subunit I (COI) gene of Calliotropis yapensis new species

supports its systematic placement within the genus.

Additional Keywords: Mollusca, Vetigastropoda, Seguenzioidea,

South China Sea, Yap Seamount

INTRODUCTION

Based on the distinctive shell morphology and radular

features, Hickman and McLean (1990) established

Calliotropidae (as the tribe Calliotropini) to group some

species previously under the Margaritinae, Angariinae,

Monodontinae, Calliostonmatinae and Solariellinae.

Members of the family are very characteristic among

the Seguenzioidea by having a thin shell with an

unthickened outer lip, aperture without denticles or

lirae, radula with a reduced rachidian and three lateral

teeth per half row. Calliotropis Seguenza, 1903 is the

most speciose genus in this family, with more than

150 valid Recent species predominantly distributed in

the Indo-West Pacific region (e.g. Australia: Jansen,

1994: New Zealand: Marshall, 1979: Solomon Islands:

Vilvens, 2007; Japan: Okutani, 2000; Philippines: Poppe

et al., 2006; China: Vilvens, 2007: Indonesia: Hickman,

2016).

Although many surveys have been carried out during

the last few decades (e. g. Bartsch, 1942; Bouchet and

Warén, 1985; Bouchet and Poppe, 1988; Marshall, 1988;

Okutani and Iwasaki, 2003: Kantor et al., 2013), the

knowledge of deep-water malacofauna of the western

Pacific is still far from sufficient. Almost every survey is

bound to discover new taxa and great numbers of new

distributional records. Over the past decade, more than 50

Calliotropis species have been described as new from the

deep waters of the western Pacific region (Poppe et al.,

2006: Vilvens, 2006; 2007; Hickman, 2016). These find-

ings strongly suggest that many more new deep-sea

species are awaiting description.

In recent years, several deep-sea surveys have been

carried out by Institute of Oceanology, C hinese Academy

of Sciences (IOCAS). During these surveys, two species of

Calliotropis were sampled: one represents a new record

for Chinese water and another one belongs to an unde-

scribed species. In present study, we describe and illus-

trate these two species.

MATERIALS AND METHODS

The studied specimens of Calliotropis infundibulum

(Watson, 1879) were trawled from the South China Sea,

whereas the specimen of Calliotropis yapensis new spe-

cies was collected by the mechanical arm of the ROV

FaxiAn (based on mother-ship R/V Kexuey!) from Yap

Seamount. Specimens are deposited in the Marine Bi-

ological Museum, Chinese Academy of Sciences

(MBMCAS).

Morphological Methods: Shell morphology was ob-

served under a stereomicroscope, and photos taken using

a digital camera. The soft parts of the new species were

retracted too deep inside the shell. Thus, it would be

difficult to extract the soft parts and consequently the

radula, without breaking the shell. Shell measurements

were taken using a caliper with accuracy of 0.1 mm.

Molecular Procedures: One specimen of Calliotropis

yapensis new species was used for molecular analysis.

Inge

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THE NAUTILUS, Vol. 132, No. 1

bes ©

Figures 1-11.

Calliotropis species. 1-5 Calliotropis infundibulum (Watson, 1879), MBM280611, 9.4x9.2 mm, South China Sea.

6-11. Calliotropis yapensis new species, Holotype MBM285093, 25.0 X 26.2 mm, 8°55’ N, 137°48’ E, Yap Seamount. 6-10. Different

views of shell. 11. Operculum, maximum diameter 10.5 mm.

Genomic DNA was extracted with the Column Genomic

DNA Isolation Kit (Beijing TIANGEN, China) according

to the manufacturer instructions. DNA was eluted in

elution buffer and stored at -20°C until use. The COI

region was amplified by polymerase chain reaction

(PCR) using the primers LCO1490 (forward: oe

GGTCA: ACAAATG ATAAAGATATTGG-3) and HCO2198

(reverse: 5/-TTAACTTCAGGGTGACCAAAAAATCA-3)

(Folmer et al., 1994). PCR reactions were carried out in

a total volume of 50 wl, including 1.5 mM MgCls,

0.2 mM of each dNTPs, | wM of both forward and reverse

PCR primers, 10Xbuffer and 2.5 U Taq DNA poly-

merase. Thermal cycling was performed with following

conditions: 95°C for 3 min (initial denaturation), followed

S. Zhang and S. Zhang, 2018

Page 15

by 35 cycles of 95°C for 30s (denaturation), 40°C for 30s

(annealing), 72°C for 60s (extension) and a final extension

at 72°C for 10min. PCR products were verified on

a GelRed-stained 1.5% agarose gel and purified with the

Column PCR Product Purification Kit (Shanghai Sangon,

China). Purified products were sequenced in both directions

using the BigDye Terminator Cycle Sequencing Kit (ver.

BIL Applied Biosystems) and an AB PRISM 3730 (Applied

Biosystems) automatic sequencer. The sequence has been

deposited in GenBank (Accession number: MF499154).

Sequence alignments were generated using Clustal X

(Larkin et al. 2007). Neighbor- -joining (NJ) tree was

performed by MEGA 6.06 (Tamura et al. 2013), using

Kimura 2-parameter (K2P) model (Kimura, 1980).

Bootstrap analyses were performed with 1000 replications.

SYSTEMATICS

Superfamily Seguenzioidea Verrill, 1884

Family Calliotropidae Hickman and McLean, 1990

Genus Calliotropis Seguenza, 1903

Type Species: Trochus ottoi Philippi, 1844. Pliocene-

Pleistocene, Italy, by original designation.

Calliotropis infundibulum (Watson, 1879)

(Figures 1-5)

Trochus infundibulum Watson, 1879: 707-708.

Trochus (Margarita) infundibulum: Watson, 1886: 84, pl.

D, fig: d:

r= go (| 4B365231.1 Ginebis argenteonitens

0.02

96 EU530113.1 Ginebis crumpii

KF644317.1 Cidanna cidans

EU530115. Lischkeia alwinae

KY 426958.1 Lischkeia impenalis

AB365229.1 Calliostropis pagodiformis

MF499154 Calliotropis yapensis new species

AB365230.1 Callotropis sp.

sido.goIjed

HE800595.1 Callotropis antarctica

AB481196.1 Calliotropis chalkeie

KU759008.1 Bayerotrochus delicatus outgroup

Figure 12. Neighbour-joining tree for Calliotropidae based

on available COI sequences from this study and GenBank.

Numbers above branches indicate the bootstrap values.

Solariella infundibulum: Dall, 1889a: 380-381; Abbott,

1974: 41, fig. 287; Cernohorsky, 1977: 105, fig. 1

Calliotropis infundibulum: Marshall, 1979: 531, figs.

4E-G, 9C-F; Okutani, 2000: 59, pl. 29, fig. 25.; Vilvens,

2004: figs. 27-28; Vilvens, 2007, figs. 84-85.

Description: Shell (Figures 1-5) conical (H/W=1.0),

very thin (aperture margin 0.2 mm thick), with 7 whorls,

low-spired, spire 36% of shell height. Shell surface

nacreous, indicative of exfoliation of outer shell layer

that probably occurs during collection and/or preservation.

Suture strongly impressed. Spire whorls angularly convex,

with two be ie d spiral cords separated by wide, smooth

interstice. An additional spiral cord occurring above

periphery of body whorl. Axials obsolete but still visible

near spiral cords. Basal area with 4 spiral cords, separated by

regularly spaced interstices. Interstice region sculptured

with numerous oblique, curved axial isis ts. Umbilicus

deep, wall sculptured with prominent axial ribs.

Material Examined: Two empty shells, MBM280611,

South China Sea, 17°15’ N, 111°20' E, 1238 m deep,

sandy bottom, collected by Agassiz trawl from the abyssal

plain (Mother-ship R/V KEXxuEYI, IOCAS), 13 Se pte »mber

2011.

Distribution: Atlantic North America (Abbott, 1974):

Caribbean (Watson, 1886); Brazil (Dall, 1889); South

Africa (Martens, 1903): Marion Island (Prince Edward

Islands), Atlantic-Indian-Antarctic Basin (Watson, 1879):

north-eastern New Zealand (Marshall, 1879), Japan

(Okutani, 2000) and China (this study). Depth range

230-3259 m (Clarke, 1962).

Remarks: No significant morphological differences

could be found to separate the studied specimens from

the photographs of the holotype (Cernohorsky , 1977)

and syntypes (Vilvens and Swinnen, 2008).

Calliotropis infundibulum (Watson, 1879) (Type lo-

cality: Prince Edward Island, Indian-Atlantic Ridge area,

46°46 S, 45°31’ E, 2514 m) is regarded as broadly dis-

tributed throughout the Indo-Pacific and tropical western

Atlantic. However, the eastern Pacific barrier and the

Mid-Atlantic barrier, and the presence of cold waters

around polar or temperate margins of continents, have

played important vicariant roles in the evolutionary history

of tropical marine taxa (Duda and Kohn, 2004). These

vicariant factors suggest that the spe cimens identified as

Calliotropis infundibulum from Indo-West Pacific are

probably not conspecific with those from the Atlantic.

Previous authors compared the specimens from the two

localities, and found no reason for further separation

(Marshall, 1979: Vilvens and Swinnen, 2008). These

studies, however, have mainly focused on shell mor-

phology and lack detailed information on anatomical and

molecular characters, although a radula from New Zea-

land was described by Mar Shall (1979) and a soft part from

America described by Dall (1889b). Consequently, iden-

tification of Indo-Pacific specimens must be tentative until

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THE NAUTILUS, Vol. 132, No. 1

supporting antomical and molecular data are available for

comparison.

At a morphological level, although some variations exist

among the shells from different localities e.g., shell size

varies from 15.3 mm to 23.5 mm, spire depressed to high,

suture shallowly to deeply compressed, etc., serial in-

termediate forms may present (Vilvens and Swinnen,

2008). The main sculpture characters of this species are

relatively consistent, e.g., spire whorl with 2 granular

spiral cords and body whorl with 3 ones, with the adapical

one strongest, axial ribs obsolete but still visible near the

granules; basal area with 4 thin g granular spiral cords, the

innermost one with strong nodules; umbilicus broad,

deep, with axial ribs, no spiral cords.

To date, about five species of the genus have been

reported from the South China Sea including Calliotropis

annonaformis Lee and Wu, 2001, Calliotropis scalaris

Lee and Wu, 2001 and Calliotropis chunfuleei Chino,

2014 from Pratas Island at depths of 400-500 m,

Calliotropis philippei Poppe, Tagaro, and Dekker, 2006

from off the Philippines at depths of 550-884 m, and

Calliotropis helix Vilvens, 2007 from Taiwan at depths

of 790-904 m. Among them, Calliotropis helix most

resembles the present studied specimens in general

shape, but differs in having distinct sculpture and a much

more depressed body whorl. Calliotropis infundibulum

(Watson, 1879) is recorded for the first time from the

coast of China.

Calliotropis yapensis new species

(Figures 6-11)

Description: Shell (Figures 6-10) solid (aperture

margin ~1.0 mm thick), large for the genus, 25.0 mm high

and 26.2 mm wide (height/width=0.95), umbilicate; spire

low, broadly conical, height 66% of width; suture mod-

erately impressed. Body egal globular, occupying 2/3 of

shell Jer igth. Exterior shell layer very thin and translucent,

underlying nacre visible. A very thin, brownish intritacalx

well pre eRe on body whorl. Protoconch unknown.

Teleoconch of 6 regularly expanding convex whorls.

Subsutural ramp broad and ornamented by axial ribs.

Sculpture of granular spiral cords with prominent,

densely packed prosoeline axial ribs between interstices.

Intervals between ribs twice as broad as ribs. In addition to

three spiral cords on spire whorl, a fourth finely granular

spiral present, concealed by suture throughout ¢ ‘aowiah

becoming exposed on periphery of body whorl. Nodnles

very pronounced, rounded or slightly sharp. Number of

nodules on penultimate whorl: 24 on first cord, 25 on

second cord, 30 on third cord; number of nodules on body

whorl: 33 on first cord, 38 on second cord, 45 on third

cord, 75 on fourth cord. Basal area with 6 finely beaded

spiral cords, innermost one strongest, bordering the

umbilicus, interstices between cords with secondary

axial riblets. Umbilicus wide, fully covered by reflected

projection of parietal and columellar callus, wall sculptured

with thin axial riblets. Aperture subquadrate, 1.2 times

higher than spire, backward-sloping, inside yellowish with

iridescence. Parietal lip with a very thin, translucent callus.

Operculum (Figure 11) paucispiral, sub-circular in shape,

with central mmeleuse

Type Locality: Yap Seamount, Say IN, IBA" 18,

foraminiferal ooze, 1119 m depth.

Type Material: Holotype. MBM285092, collection

number: Y30057, station: FX-DIVE 18, Yap Seamount,

8°55’ N, 137°48’ E, foraminiferal ooze, 1119 m deep, 18

December 2014.

Distribution and Habitat: Only known from the type

locality, living on foraminiferal ooze bottom.

Etymology: The name of new species refers to its type

locality.

Remarks: Calliotropis yapensis new species is char-

acterized by a large shell with convex teleoconch whorls,

shell surface sculptured with granular spiral cords and

fine, dense, regularly spaced axial ribs. These features can

easily separate new species from most species of the

genus. Calliotropis yapensis new species is in general shell

shape most similar to Calliotropis ammos Valvene! 2012

from French Polynesia, which also has low spire and

rounded body whorl. Calliotropis ammos Vilvens, 2012,

however, differs from the new species in having a smaller

shell (height 294-945 mm, width 21.4-23.8 mm) with

a larger number of teleoconch whorls (6.9-7.1), and a

lower aperture (height/height of the aperture 2.55—3.03

vs. 2.27 in new species). In addition, Calliotropis yapensis

new species can be separated from Calliotropis ammos

Vilvens, 2012 in having a very thin exterior shell layer

through which the underlying nacre can be seen, 6

cmnteaG of 5 spirals on the base, prominent axial ribs

present between the spirals, and an aperture that

not transversely elongated (Vilvens, pers. comm.).

Calliotropis blacki Mleneslheall, 1979 and Calliotropis

derbiosa Vilvens, 2004 may be also confused with

Calliotropis yapensis new species, but differ in

Table 1. Pairwise distances among species of Calliotropis based on Kimura 2-parameter model.

1 2 3 4

l MF499154 Calliotropis yapensis new species

2 AB365229.1 Calliotropis pagodiformis 0.092

3 AB365230. 1 Calliotropis sp. 0.117 0.117

4 AB481196.1 Calliotropis chalkeie 0.192 0.197 0.199

5 HE800595.1 Calliotropis antarctica 0.100 0.120 0.105 0.197

S. Zhang and S. Zhang, 2018

Page 17

having much more angulate teleoconch whorl and

depressed body whorl.

Molecular Analyses: One sequence for the COI re-

gion of Calliotropis yapensis new species was obtained.

After manual adjustment, the length of the partial COI

sequence of new species is 633 bp. “The Neighbor-joining

(N]) tree (Figure 12) was reconstructed using available COI

sequences feeorta this study and GenBank. The alignment

of COI had a total 594 bp, without insertions and alate tions.

The NJ tree shows that the new species fall into the genus

Calliotropis in which Calliotropis yapensis new species

together with other Calliotropis spp. form a well-supported

clade, supporting its placement within Calliotropis. The

phy logenetic tree also supports monophyly of a group of

5 Calliotropis species within the family. Within the available

sequences from GenBank, Calliotropis yapensis new species

is closest related to Calliotropis pagodiformis, with 9%

pairwise distance, whereas the pairwise distances among

Calliotropis yapensis new species and other species of

Calliotropis are 10-12% (see Table 1).

In addition to the present new species, other two basal

(Vetigastropoda) species were collected together during

the survey, including a pleurotomariid species, Bayero-

trochus delicatus Zhang, Zhang and Wei, 2006, and

a undescribed colloniid species, Homalopoma sp. The

three species had never yet been found elsewhere, and

thus were considered potential Yap Seamount endemics.

ACKNOWLEDGMENTS

We would like to express our sincere thanks to Claude

Vilvens (Belgium) for his enthusiastic support and in-

formation. Thanks also to the crews of R/V KExuEy! for

their cooperation during the survey. This research was

supported by the Strategic Priority Research Pr ogram of

the Chinese Academy ae Sciences (XDA11030201). the

National Natural Science Foundation of China (No.

41606162) and the Senior User Project of R/V KEXUE

(KEXUE2017G05)

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THE NAUTILUS 132(1):19-29, 2018

wey IC

Page 19

Redescription of the jumping snail Ovachlamys fulgens

(Gude, 1900) (Gastropoda: Helicarionoidea: Helicarionidae):

An anatomical and conchological approach

Anna C. A. Salles

Cléo D. C. Oliveira and

Ricardo S. Absalao

Laboratorio de Malacologia, Instituto de Biologia

Departamento de Zoologia, Universidade Federal do Rio de Janeiro

21941-902 Rio de Janeiro, BRAZIL

[email protected]

ABSTRACT

The geographical distribution of the jumping snail Ovachl-

amys fulgens Gude, 1900, originally described from Loo-Choo

Island, is expanding rapidly throughout the tropics. The full

extent of the environmental damage caused by the in-

troduction of this exotic species is ill unknown. So far, it has

been implicated in damage to orchids and horticultural plants.

Ovachlamys fulgens is only known by a few characteristics of

the shell and information on its anatomy is scant. The present

study aims to redescribe this species based on specimens from

Brazil, which is here characterized by the a globose shell, with

wide aperture, externally micro- sculptured with undulating

spiral groves, umbilicus partially obliterated by a thin plate,

pedal Role tripartite, cephalopedal mass consisting of a ser-

ies of oblique muscles, mantle with two flattened lobes, large

caudal hom, pallial cavity small, vascular system evident,

auricle fibrous, ventricle highly muscular, aorta subdivided in

cephalic and gastric veins, a large kidney internally folded, jaw

seers ail@xeaTonte, acne. with 127 teeth per row (55-(8)-

1-(8)-55), each row with about 25 ym, crop absent, salivary

glands eae stomach large with thin walls, ovotestis with at

least three distinct lobes, talon totally immersed in the al-

bumen gland, uterus with two regions, capsule gland hardly

visible, bursa copulatrix small arnal sacculiform, penial sheath

present, epiphallus small and narrow, nervous ring asym-

metrical, visceral ganglion on left side only, five and six nerves

running from each cerebral and pedal ganglia, respectively,

one statocyst immersed in each pedal ganglion, and two pairs

of ganglia (mandibular and buccopharyngeal ganglia) extra to

theinenons ring.

Additional Keywords: Atlantic Forest, exotic species, shell

morphometrics, PARNASO

INTRODUCTION

The Helicarionoidea Bourguignat, 1877 is a diverse

group of pulmonate gastropods usually found in Africa,

Asia, and Australasia (Smith et al. 2002). Anatomical data

for many lineages in the Helicarionoidea is unknown, the

phylogenetic relationships of those lineages are poorly

understood, and the taxonomic concept fan many families

is unclear (Hyman and Ponder, 2010). The key characters

considered to discriminate families are anatomical (e.g.,

stimulator, epiphallic caecum, flagellum) which seem to

have been reduced or lost several times in many groups

(Hausdorf, 1998; Schileyko, 2002a, b; Hyman, 2007; Hyman

and Ponder, 2010).

Among the Helicarionoidea, Ovachlamys fulgens

(Gude, 1900) is known from shell characters and poor

anatomical data from reproductive system (Schileyko,

2002a, b). This species was originally described from

Loo-Choo Island, Japan. So far, this is considered the

place of origin for this species, but its distribution is

expanding rapidly throughout the tropics, most likely

unintentionally, as a neat of the horticultural trade

(Robinson and Slapcinsky, 2005). Ovachlamys fulgens

has already been recorded in some Pacific islands

(O’ahu and Hawai'i islands, in the Hawaiian Archi-

pelago and Tutuila and Olosega, in American Samoa), in

Brazil (Sao Paulo State), in Trinidad and Tobago, Costa

Rica, and Florida (Barrientos, 1998, 2000; Cowie, 2001:

Robinson and Slapcinsky, 2005; Cowie et al., 2009;

Teixeira et al., 2017). It has also been reported i

horticultural shipments from Thailand, Singapore, er

Colombia, but identification still lacks proper confir-

mation (Robinson, 2003; Robinson and Slapcinsky,

2005).

Ovachlamys fulgens is considered an important pest to

orchids and it is able to attack horticultural plants (Stange,

2004). The environmental damage caused by the in-

troduction of this exotic species is still unknown. The

main objective of this paper is to redescribe and il-

lustrate shells of type material and anatomy of recently

collected material of O. fulgens in a Brazilian national

park, in Rio de Janeiro State. With this redescription,

we hope to provide a baseline for future comparative

Page 20

THE NAUTILUS, Vol. 132, No. 1

_ Rio de Janeiro —

Figure 1. Ovachlamys fulgens. Locality of PARNASO (red

star), where the field work was conducted in Rio de Janeiro

State, Brazil.

studies among Helicarionoidea and help understand

the biology oe this exotic species, which has potential

to Keone distributed worldwide in the tropics and

subtropics and cause serious damage to environment

and horticultural activities.

MATERIALS AND METHODS

The specimens from Brazil were collected in Parque

Nacional da Serra dos Orgaos (PARNASO), a national

park with 20,024 ha of Nelemtre Forest biome. This

conservation area is located in Rio de Janeiro State

quantitative features. Syntypes in blue.

3.00

2.25;

-1.50;

-3.00

(Figure 1) and encompasses part of the municipali-

ties of Guapimirim, Teresépolis, Petropolis, and Magé.

PARNASO presents an altitudinal gradient ranging

from 200 to 2,263 m above sea level, with average annual

temperature of 13-23° C, and average annual pr ecipitation

reaching about 2,800 mm/year, with the highest values

between November and March (~ 360 mm/month)

(ICMBIO, 2008).

Specimens were hand-collected, mainly around the

park headquarters in Teres6polis and Guapimiri im at low

elevations (400-950 m a.s.l.) from October 2015 to

August 2017, totaling 13 field trips (2-3 days each).

Specimens were drowned in distilled water for 24h and

then preserved in 70% ethanol. Voucher specimens are

deposited at the Biological Institute of Universidade

Federal do Rio de Janeiro (IBUFRJ) according to au-

thorization provided by Chico Mendes Institute for

Biodiversity Conservation (ICMBIO).

Taxonomic identification was based on conchologi-

cal comparisons with type material and the original

description and illustration. Twenty-two undamaged

shells (2 shells of syntypes and 20 additional recently

collected shells deposited at IBUFR] 21236) were se-

lected for morphometric analysis, based on size, grading

from the smallest juvenile to the largest adult shell. Six

measurements were taken as descriptors of morpho-

logical variation (Figure 2): shell height (H), shell width

(W), height of spire (Hs), height of last whorl (H1),

height of aperture (Ha), width of aperture (Wa). Also,

PCL

Figure 2. Ovachlamys fulgens. Shell descriptors and the scatter plot of 22 studied specimens along PC1 and PC2 based on 11

A.C.A. Salles et al., 2018

age 2

Page 2]

Figures 3-6.

Brazil (IBUFRJ 21236). Scale bar =1 mm.

the ratios of: shell height to shell width (H/W), height of

aperture to width of aperture (Ha/Wa), shell width to

width of aperture (W/Wa), shell height to height of spire

(H/Hs), and shell height to height of last whorl (H/H1).

Original data were standardized and size: adjusted using

the Burnaby (1966) method. A principal component

analysis was conducted. The curve of expected eigen-

values of a random model (Broken Stick) was use a to

select significant components. Eigenvalues for components

under This curve were considered non-significant and

then discarded (Jackson, 1993). All stetaciical pro-

cedures were performed with PAST software (Hammer

et al. 2001).

Detailed external morphology and gross anatomy were

examined in laboratory using a stereo- microscope Zeiss

Stemi SV 11 with attached camera lucida. Dissections were

performed using standard techniques, with specimens im-

mersed in fixative under the stereo-microscope. Organs of

the pallial cavity, reproductive, digestive, circulatory, excre-

tory, and nervous systems were investigate »d. Radulae and jaw

were manually extracted and prepared by immersion in

a 10% solution of sodium hypochlorite for 24h, and later

rinsed in distilled water. Nineteen specimens were dissected,

from which six had the radulae and jaw removed for imaging.

Ovachlamys fulgens. Shells in apical and apertural view. 3-4. Syntype (NHMUK 1922.8.29.12). 5-6. Specimens from

Images were taken using a Canon SX170 IS digital

camera and SEM photography using a JEOL JSM-6510 at

‘Laboratorio de Imagens em Microscopia Optica e de

Varredura (LABIM-UFR]J)”, at Department of Zoology,

UFR].

Anatomical comparisons of the reproductive system

were performed against taxa illustrated in Schileyko

(2002a, b), Sutcharit and Panha (2008), and Hyman

and Ponder (2010). Only species ah similar anatomy

and detailed information available were included on

Discussion.

Anatomical abbreviations used are: A, anterior; ag,

albumen gland; aga, parietal ganglia; ao, aorta; au, au-

ricle; be, bursa copulatrix; bga, buccopharyngeal ganglia;

bm, buccal mass; br, retractor muscle of buccal mass; ea,

caudal apparatus; car, carrefour; eg, capsule gland; ega,

cerebral ganglia; dg, digestive gland; ep, epiphal llus; ga,

genital atrium; gv, gastric vein; her, hermaphroditic duct;

il, intestinal loop; ja, jaw; ki, kidney; Iga, Sua ganglia;

me, mantle edge; mga, mandibular ganglia; ml, “anes

lobes; ms, moni slit; nel-ne5, nerves of the cerebral

ganglia; np1—np6, nerves of the pedal ganglia; oc, ocular

tentacle; oe, oesophagus; or, oral tentacle; ov, oviduct;

ovt, ovotestis; pe, pericardium; pe, penis; pec, epiphallic

Page 22

THE NAUTILUS, Vol. 132, No. 1

Figures 7-14.

caecum; pga, pedal ganglia; pn, pneumostome; po,

protractor muscle of the buccal mass; pr, prostate; prm,

penial retractor muscle; ps, penial sheath; pu, primary

ureter; pv, pulmonary vein; re, rectum; ro, retractor

muscle of the odontophore; rs, radular sac; sd, ducts of

salivary gland; sg, salivary gland; st, statocyst; sto,

stomach: su, secondary ureter; ta, talon; ut, ute rus: ut,

second region of the uterus: vd, vas deferens: ve, ve anata

vg, vagina; vga, visceral ganglia; vs, vascular system.

RESULTS

Systematics

Helicarionoidea Bourguignat, 1877

Helicarionidae Bourguignat, 1877

Ovachlamys Habe

Ovachlamys fulgens (Gude, 1900)

(Figures 2-30)

Macrochlamys fulgens Gude, 1900: 75, pl. 8, figs 24-26.

Description: SHELL (Figures 3-4) globose, dextral,

very fragile, translucent, reddish to pale g golden. Shell

surface micro-scul ptured with undulating spiral grooves.

Ovachlamys fulgens. Shells. 12-14. IBUFRJ 21217. 7-9. Growth series, apical view. 7. IBUFR]J 21218. 8. IBUFR]J

21219. 9-11. IBUFRJ 21221. 10-11. Details of protoconch and extemal microsculpture. 12. Adapical view. 13-14. Details of external

microsculpture and umbilical perforation. Scale bar for 7-9, 12=1 mm; 10, 14=200 pm; 11, 13=50 pm.

Whorls 2-4, regularly increasing in diameter. Last whorl

rounded or somewhat SHouloreadl Umbilicus deep,

partially filled by a small reflection of the peristome.

Aperture crescent- shaped, with simple lip.

EXTERNAL MorpHoLocy: Soft parts (Figures 15-17):

External surface grades from light- brown to dark- gray

during ontogeny. Pedal sole tripartite. Cephalopedal

mass Taaetally built from a series of oblique muscles.

Two pairs of cephalic tentacles. Mantle edge extended

giving rise to two flattened lobes, which cover part of

the shell when the animal is active. Caudal apparatus

consisting of curled-up posterior part of pedal sole,

caudal Theva large, nearly heart-shaped. Caudal groove

present.

PALLIAL Cavity, CIRCULATORY AND EXCRETORY SYSTEMS

(Figures 18-20): Pallial cavity small, short, nearly rect-

angular in shape. A secondary ureter crosses pallial cavity

eral opens to exterior close to pheumostome. Rectum

large, running close to but not attached to secondary

ureter, opening outside pallial cavity. Vascular system with

large and well defined veins. Pericardium small, dis-

cernible on fresh specimens. Auricle fibrous, translucent,

connected to a pulmonary vein that enters pallial cavity.

Ventricle highly muscular, opaque, small, half size of

auricle, bonneeted to short aorta. Aorta subdivided into

cephalic and gastric veins, former hardly visible, latter

evident, with Wasollan ramifications on digestive gland.

A.C.A. Salles et al., 2018

Figures 15-17. Ovachlamys fulgens. Living specimens. 15.

General external morphology. 16. Detail of caudal apparatus. 17.

Detail of mantle lobe lining the shell; pulmonary vein and rectum

are visible through the shell. Empty and filled arrows indicate the

caudal horn and mantle lobe, respectively. Scale bar=2 mm.

Kidney rectangular to trapezoidal, corresponding to

about 40% of pallial cavity size. Primary ureter large, nar-

rowing toward secondary ureter. Nephridiopore opening to

ete close to pneumostome. Kidney and ureters in-

ternally folded.

DIGESTIVE SysTEM (Figures 21-26): Mouth a vertical slit

in antero-ventral margin of snout. Buccal mass rectangular.

Odontophore about half of buccal mass volume,

subdivided into two elliptical bulbs. Two pairs of

muscles insert at each odontophore bulb: one strong

and large retractor muscle inserted at posterior region

runs backward, and one thinner protractor muscle

inserted at posterior region of bulb runs obliquely

toward lateral body wall. Thin retractor muscle,

inserted ventrally, near center of buccal mass, runs

backward between odontophore bulbs. Radular sac

projected posteriorly, with about half size of buccal

mass. Oesophagus long, medium-dorsally inserted at

buccal mass. Crop absent. One pair of salivary glands

fused, overlaying oesophagus latero-dorsally. One pair

of narrow and thin walled ducts connects salivary

glands to buccal mass near buccopharyngeal ganglia.

Stomach large and elongated, with thin walls, dorsal to

visceral mass. Stomach connects to ventral, S-shaped,

intestine. Both intestine and stomach partially em-

bedded in granular digestive gland, which occupies the

remaining valine of ne spire. Anus opening close to

pneumostome.

RADULA AND JAw: Radulae with 127 teeth per row

(55-(8)-1-(8)-55), each row measuring about 25 pm.

Central tooth symmetric tricuspid, rectangular, mes-

ocone large, lanceolate, ectocones pointed, located on

about 2/3. of tooth length. Lateral teeth asymmetric,

tricuspid; mesocone large; lanceolate: endocone small,

blunt, placed near foot tip; ectocone, triangular,

pointed, located on middle of tooth. Marginal teeth

sinoidal and elongated, bicuspid, endocone slightly

longer than ectocone. Jaw smooth and crescentic,

without central cusp.

REPRODUCTIVE SYSTEM (Figure 27): Ovotestis fragile,

with at least three distinct lobes; hermaphroditic duct

almost straight, convoluted towards small talon, which

is embedded in albumen gland. Carrefour rounded to

ovate, close to talon, partially immersed in albumen

gland. Albumen gland bean-shaped, with corrugated

Sell touching sioner. Prostate and uterus running

alongside. Uterus spacious, with large lobules, and

anindlividled! in two regions, one with prostate partially

embedded and other “expanded below albumen gland,

grading in size during ontogeny. Capsule gland Fenalhy

“Agile, located close to Cancefour Benncont uterus and

albumen gland, totally immersed in latter. Bursa

copulatrix small and sacculiform, inserted on vagina by

short duct. Penial sheath present. Penis long, cylin-

drical and slightly compressed in transversal section.

Epiphallus Sinlll and narrow, about 1/5 of penial

length. Penial retractor muscle attached to tip of

epiphallic caecum. Vas deferens emerging from prostate

gland, connected at junction of penis and epiphallus.

Penis and vagina inserting side-by-side in long genital

atrium.

Nervous SysTEM (Figures 28-30): Pairs of cerebral,

pleural, parietal, and pedal ganglia connected and

partially fused, forming agmmnenaical nervous ring.

Visceral ganglion on lefticide only. Five nerves running

from each cerebral ganglion (ncl- ned), two ae

ventro-lateral region (ncl—nc2), one connects to tip

(nel) and the ane to the base (nc2) of ocular ten-

tacles. The remaining three cerebral ganglion nerves

(nc3-ne5) run from anterior region and connect,

respectively, to oral tentacle, to mandibular ganglion,

and to buccopharyngeal ganglion. Pair of pedal gan-

glia gives rise to Barer al diclhente nerves running

ventrally towards pedal sole (np1—np6), but exact e nl

target of those nerves in pedal sole not determined.

One statocyst immersed in each pedal ganglion. Two

additional pairs of ganglia are connected to nervous

ring: one pair of buccopharyngeal ganglia close to

pharynx, on dorso-lateral surface of areca mass; and

one pair of mandibular ganglia near distal end of

buccal mass.

Page 24 THE NAUTILUS, Vol. 132, No. 1

Figures 18-26. Ovachlamys fulgens. 18-19. External morphology. 18. Apical view, shell removed. 19. Lateral view, digestive gland

slightly turned up to expose the gastric venation and kidney partially removed to expose the heart. 20. Pallial cavity and associated organs.

21. Digestive system. 22. Buccal mass, ventral view. 23. Jaw. 24. Radular teeth. 25. Close-up to show the central, lateral and (26)

marginal teeth. Scale bar for 18-22 =1 mm; 23=100 pm; 24=50pm; 25-26=20 pm.

A.C.A. Salles et al., 2018

Page 25

her

Figure 27.

Morphometric Analysis (Figure 2, Tables 1-3): Only

the first two components were considered significant. No

prm

segregation in clusters were observed in the scatter plot of

voucher specimens collected in PARNASO and syntypes

of O. fulgens ordinated along the PC] and PC2. The

Component | was most meueneed by the ratios of the

shell height to height of spire (H/Hs) and the shell height

to height of last whorl (H/H1). The Component 2 was

most influenced by the ratio of the shell width to width of

aperture (W/Wa).

Type Material: Field Museum of Natural History,

FMNH 43045, syntype, 1 dry shell; Natural History

Museum, ondon U.K.., NHMUK 1922.8.29.12, syntype,

1 dry shell.

Type Locality: Loo Choo Islands, Japan.

Other Material Examined: Brazil, Rio de Janeiro,

PARNASO: Guapimirim: IBUFR] 21217 (1 shell), IBUFR]

21218 (1 shell), IBUFRJ 21219 (1 shell), IBUFR] 21220

(1 shell), IBUFRJ 21221 (1 shell), IBUFRJ 21222 (1 shell),

IBUFR] 21225* (93 specimens), IBUFR] 21229 (4 speci-

mens). Teres6polis: IBUFR] 21226 (1 specimen), IBUFR]

21236 (20 shells). Lot with ne, specimens marked by

an asterisk.

Behavior: Activity of O. fulgens in PARNASO seems

to be correlated with high humidity and all live speci-

mens were collected at night or after rainfall, always

near the trails edge, inside the park. Most specimens

were found on herbaceous vegetation (e.g. Hibiscus sp.,

Quesnelia sp., and Heliconia sp.) about 100-150 cm

above ground. Some live specimens were also found in

Ovachlamys fulgens. Reproductive system, view from two sides. Scale bar=1 mm.

leaf litter, on exposed rocks, on green moss, and on

fallen trunks.

When disturbed, the specimens started the typical

behavior under stress conditions exhibited by this

species, for which it is popularly known as the jumping

snail. In such behavior, the animal retracts the head

inside the shell and turns the pedal sole, including the

caudal horn, in an abrupt and repeatedly circular movement

(usually accompanied by an increase in mucous secretion).

Most likely this activity is intended to discourage pred-

ators and often results in the snail falling from Teave 5 to

the ground.

Such behavior represents a peak of high metabolic

rates and comes at a cost. After a short period of intense

activity with usually two or three cycles of intermittent

jumps (each cycle takes about 2 seconds long) the

animal often becomes sluggish and appears exhausted.

The presence of a powerful vascular system with large

vessels certainly assists in promptly supplying the Beale

needs of energy and oxygen during the jumps, but the

small sized pallial cavity seems counterintuitive in

providing the necessary amount of oxygen. We suppose

the presence of body extensions may assist this need.

Such extensions are most often observed in marine

gastropods (e.g. Cypraeidae and Ovulidae) as projec-

tions of the foot and are associated with camouflage,

shell repair, anti-predatory activities, etc. (Vermeij,

1978). In O. fulgens the mantle edge extends forming

two thin, wide lobes, each one lies externally on each

side of the shell. These lobes presumably assume

a gas exchange function. Although further studies are

required, structures with increased surface area to

volume ratio are usually associated to exchange with

Page 26

THE NAUTILUS, Vol. 132, No. 1

28 see

NESrnGe4

nc3

Figures 28-30. Ovachlamys fulgens. Nervous system. 28. Nervous ring, cerebral commissure cut; 29. Pair of pedal ganglia, ventral

view. 30. Lateral view, right and left side, respectively; nerves from the cerebral ganglia (nc] and nc2) and nerves nerves from the pedal

ganglia (np2—np5) not illustrated. Scale bar=1 mm.

environment, mainly to gas exchange in terrestrial organ-

isms (Schmidt-Rhaesa, 2007). Since the presence of such

lobes increases the water loss by the animal, the foraging

activity is limited to high humidity periods and spots.

DISCUSSION

Shell Morphometrics: Anatomical data were absent

in the original description of Ovachlamys fulgens.

Based solely on shell attributes, no significant differ-

ences were found among between the syntypes

(photos) of O. fulgens and the voucher specimens from

PARNASO. It reinforces the conclusion that the re-

cord of this species to PARNASO is not based on

a misidentification. The most discriminative features

among the quantitative descriptors used (Table 3) are

the globose shell and the wide aperture. Those fea-

tures emerge as important conchological characteris-

tics for the taxonomy of O. fulgens.

Morphology: REPRODUCTIVE SysTEM: One remark-

able feature of O. fulgens is the uterus subdivided in two

distinct regions. The region close to the albumen gland is

reduced or even inconspicuous in young specimens, be-

coming as large as the albumen gland in adults.

Among Ariophantidae, Macrochlamys petrosa and

Sarika asamurai differ from O. fulgens by the very

convoluted hermaphroditic duct, the short genital atria,

and the presence of flagellum and stimulator. Addi-

tionally, in M. petrosa the bursa copulatrix is large and

A.C.A. Salles et al., 2018

Page 27

Table 1

Ovachlamys fulgens. Morphometrics. Unprocessed values: minima, maxima, means, and standard deviations. Values in

millimeters. Total of 20 analyzed specimens from PARNASO and two syntypes. Measured variables: shell height (H), shell width (W),

height of spire (Hs) height of last whorl (H1), height of aperture (Ha), width of aperture (Wa), shell height to she sll width ( H/W), height of

aperture to width of aperture (Ha/Wa), shell width to width of aperture (W/Wa), shell height to height of spire (H/Hs), and shell he ight to

height of last whorl (H/H1).

Minimum Maximum Means

H 2.73 4.86 3.80

Ww 3.99 6.81 5.47

Hs 0.36 0.75 0.50

HI 2.36 4.26 3.30

Ha 2.16 3.28 2.79

Wa B18} 3.56 2.93

H/W 0.65 0.74 0.69

Ha/Wa 0.87 1.05 0.95

W/Wa 1.82 1.92 1.87

H/Hs 5.39 10.51 7.69

H/Hl LIT 1E22 1.15

elongated, with a long duct, and the epiphallus is of

almost equal size to that of the penis. In S. asamurai,

the bursa copulatrix is sacculiform and elongated,

with a short duct, and the epiphallus is about three to

four times longer than the penis. Among the Heli-

carionidae, Helicarion cuvieri and Nitor circumcincta

differ from O. fulgens by the presence of a flagellum.

Additionally, in H. cuvieri the bursa copulatrix is

spherical and large, with a duct as long as the bursa,

the epiphallic caecum is absent, and the epiphallus is

slender and about two times longer than the penis. In

N. circumcincta, the talon is free (not immersed in

albumen gland), the hermaphroditic duct is very

convoluted, the bursa copulatrix is elongated and

indistinct from the duct, the penis is enlarged close to

the penial sheath, and the epiphallus is of same size as

the penis, narrowing towards the flagellum.

Nervous System: The mandibular and the bucco-

pharyngeal pairs of ganglia were not previously re-

ported for the family. Both are extra to the nervous

ring and directly connected to the cerebral ganglia.

The asymmetrical structure of the nervous ring is

determined by the visceral ganglion present on the left

side only. The main differences in nervous ring between

Table 2... Ovachlamys fulgens. Summary table of PCA analysis.

PCs Eigenvalue %o variance

1 3.09004 58.177

2 1.2235 23.035

3 0.656144 12.353

4 0.297415 5.5995

5 0.031913 0.60083

6 0.0102383 0.19276

7 0.0016076 0.030266

8 0.0005785 0.010892

9 3.6403E-05 6.8537E-04

10 1.09737E-05 2.066E-04

11 7.03218E-16 1.324E-14

S.D. Syntype FMNH Syntype NHMUK

0.52 4.85 4.86

0.68 6.57 6.81

0.10 0.59 0.70

0.46 4.26 4.17

0.28 3.28 3.25

0.35 3.44 3.56

0.02 0.74 0.71

0.05 0.95 0.91

0.03 1.91 1.92

1.18 8.23 6.92

0.03 1.14 7

O. fulgens and the close-related species Macrochlamys

petrosa (Hutton, 1834) (illustrated in Hyman and

Ponder, 2010: 32) are the larger thickness of the

commissures among the cerebral, pleural and pedal

ganglia in M. petrosa; and the pair of pedal ganglia,

which is partially fused in O. fulgens and separated in

M. petrosa.

Hausdorf (1998) noticed the absence of direct in-

nervation from cerebral ganglion to the penis as a non-

convergent feature and a possible synapomorphy for

Ariophantidae. To recognize the boundaries among the

families of Bicliteardloraaticlon is beyond the scope at the

present study. But it is noteworthy that the cerebral

innervation of the penis was not found in any specimens

of O. fulgens studied here, which triggers the need for

further studies and questions che validity of the

placement of O. fulgens among Helicarionoidea.

ACKNOWLEDGMENTS

We are grateful to Isabel Hyman (University of Sydney)

and Rei eshte (The University of Tokyo) for valuable

exchange of information; Jonathan Ablett (Natural

History Museum, London), Jochen Gerber and Martin

Pryzdia (both from Field Museum of Natural History,

Chicago) for providing type photos of O. fulgens. From

Universidade Federal do Rio de Janeiro, we are in-

debted to Inacio Domingos and Marcelo de Oliveira

Sales for the support with scanning electron micros-

copy; to Manuella Folly for assistance with mapping; to

Luis Vitor, Raquel Figueira, and Tarcilla Carvalho for

helping during field work. To Suzete Gomes (Oswaldo

Cruz Foundation) for providing literature and critics to

the first draft of this manuscript; to Tim Pearce (Car-

negie Museum of Natural History) ) and one anonymous

reviewer for critics and sugge stions. To Chico Mendes

Institute for Biodiversity Cones rvation (ICMBIO) for

providing authorization (License No. 51246-2) to the

present study and to PARNASO for all logistics assis-

tance during field work.

Page

THE NAUTILUS, Vol. 132, No. 1

Ovachlamys fulgens. Component loadings for each variable.

Table 3.

AG ul

0.41264

0.41406

PC 10

-().742,

Ine Y)

0.14198

-0.76835

7S)

0.113

RC 7

().4606

-0.11013

-0).12938

0.014812

-0.021614

-0.04663

0.070687

-O.11811

0.0

eC Ih

1D)

0.0657

(0.0482

(0.44066

-0.18812

-0.17176

-().60639

0.0019449

0.806

-),.22881

-0.20487

32733

0.0

Bye

-0, 10904

0.18106

0.091972

331

0.01

-0.339

0.31784

3453

0.081207

-0.05419

0.60843

-0.058

0.00

0.11285

0.56131

-0.38764

7074

-0.025868

-0.031684

-0.04283

0.057443

0.04

0.15458

-0.081263

0.15364

0.054471

-0.058934

-0.034111

().39669

0.37479

0.4077

0.62971

-0.11444

-0.0077193

-).087774

-0,.20558

0.48139

0.17965

0.196

().48253

0.15926

0.019415

0.059089

0.00

-0.0089564

0.49116

38112

-0.

(

0.03

-0.056812

0.071171

-0.25606

0.002147

0.31489

0). 10982

-0).2295

0.156

0.037357

0.21659

-().12573

-0).299

0.70018

-0.01

0.030757

33019

-0.046

().

0.34911

31889

-0.0004816

0.010777

0.0

389

0.0066025

().29406

-0.376

0.073018

0.72597

0.67528

0.65692

-().2899

0.088697

W/Wa

3445

0.046207

-0.04

-0.029504

7254

0.006

-). 10714

-0.1011

().055362

0.0035608 0.075902

(

().22801

O.1011

0.18826

-).54157

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THE NAUTILUS 132(1):30-32, 2018

Research Note

Page 30

First occurrence of a novel Corbicula

(Bivalvia: Corbiculidae) Form D lineage

in the Ohio River, USA

Additional Keywords: Asian Clam, invasive species, invasion,

exotic species, non-native species

Asian Clams (Genus Corbicula) are moderately-sized

(typically <50 mm) freshwater bivalves native to the

temperate/tropical regions of Africa, southern Asia, and

eastern Australia (Britton and Morton, 1986). Corbicula

was first recorded in North America in 1924 in British

Columbia, Canada, and breached the Continental Divide

in the late 1950s (Bates, 1962: Fechtner, 1962: Stein,

1962; Counts, 1986). Since then, it has spread throughout

the United States and into South America (Lee et al.,

2005).

Literature reports vary from an invasion of only a single

Corbicula species to invasions of multiple taxa (C ounts,

1986; Hoagland, 1986; Lee et al., 2005). However,

Corbicula taxonomy is muddled and unclear, as is the

number of species that have become established in North

America (Hillis and Patton, 1982; Hoagland, 1986).

Corbicula species appear to switch reproductive strategies

and become androgenic clones when they invade new

ecosystems outside of their native range (see Lee et al.

[2005] and references therein; Hedtke et al., 2011;

Pigneur et al., 2011; 2014). This complex reproductive

behavior can include not only hermaphroditism but also

capturing eggs of other Corbicula lineages and potentially

replacing Gteiaal nuclear DNA. Separate Corbicula

lineages can differ in their nuclear 28S ribosomal geno-

type yet share identical mitochondrial mt DNA genotypes

(Lee et al., 2005; Tiemann et al., 2017). This phenomenon

contorts our under standing of traditional species concepts

and makes it difficult to detemnine if Corbicula taxa are

either separate species or individuals of distinct pheno-

types belonging to a single polymorphic species (Britton

and Morton, 1986: Lee et al., 2005).

Hillis and Patton (1982), based on electrophoretical and

morphological data, suggested two Corbicula “forms”

occur in North America. Shortly thereafter, Britton and

Morton (1986) proposed the Form morphotype classifi-

cation and recognized two Corbicula forms in North

America—Form A and Form B. Later, Lee et al. (2005)

expanded on the Form concept and reported three forms

in the New World—Forms A and B in North America and

Mis Sour Rive’

Legend

c=) Ohio River specimen

* Corbicula Form D

Figure 1.

near Newburgh, Indiana (hexagon).

Ohio we

@® 2D Sy 100 Kilometers

non Ln}

Location of where Tiemann et al. (2017) reported Corbicula Form D (star) and where it was collected in the Ohio River,

J. Tiemann et al., 2018

Form C in South America. Morphometrically, small

(<20 mm) Forms A and B specimens can be distinguished

via nacre color—Form B has a deep purple nacre whereas

Form A has a white nacre with purple highlights (Hillis

and Patton, 1982: Britton and Morton, 1986). However,

these morphological differences become unreliable on

larger specimens (Britton and Morton, 1986).

mb Midwest had long been recognized as having only

one Corbicula taxon, Corbicula fluminea (= Form A), until

recently (Cummings and Mayer, 1992). Tiemann et al.

(2017) reported hee distinct Corbicula taxa occurring in

Illinois, including one that had never been documented

in the New World. The newest taxon, referred to as

Corbicula Form D, has fine rust colored rays on the

periostracum, white nacre with purple teeth, and external

ridges that are not as pronounced as other Corbicula taxa.

Corbicula Form D was known from only a small stretch of

the Illinois River in northern Illinois, but data were lacking

as to other locations. Herein, we report on an additional

location of Corbicula Form D.

On 24 October 2017, one live individual of Corbicula

Form D (Figure 1) was collected in the Ohio River in the

Page 3]

Newburgh Pool upstream of Rockport, Indiana at river

mile 744.4 (Latitude 37.9111° N, Longitude 87.0123° W).

This specimen (Figure 2) was deposited for accession in

the Illinois Nana History Survey Mollusk Collection,

Champaign (catalogue number INHS 87523). It should

be noted that the primary objective of this field outing to

the Ohio River was to conduct unionid sampling and

Corbicula were not the primary focus. Therefore,

we cannot assess density or relative abundance of the

Corbicula here, and are only reporting presence via

incidental encounter.

Accurate species identification and reporting is critical

in monitoring the spread and, ultimately, minimizing the

ecological and economic effects of this novel Corbicula

form (Tiemann et al., 2017). We are hesitant to speculate

as to when and how Form D arrived in the Ohio River.

Because of logistical constraints (e.g., time, effort, and

monetary ORS). Corbicula from navigable rivers, like the

Ohio River, are likely to be underrepresented during field

surveys and in museum collections. If not overlooked,

Corbicula are often simply noted as “present” with no

discernable differentiation among taxa when conducting

Figure 2. Corbicula Form D, intemal view of right valve (top left) and external view left valve (top right) and hinge of left valve

(bottom). Collected from the Ohio River, near Newburgh, Indiana (INHS 87523). This specimen is 18 mm in length. Distinguishing

characteristics for Corbicula Form D include fine rust colored rays on the periostracum and extemal ridges that are not as pronounced as

other Corbicula taxa.

Page 32

THE NAUTILUS, Vol. 132, No. 1

field surveys. Natural history museums (e.g., INHS

Mollusk Collection) do have Corbicula specimens, but

these are often larger specimens, which can be unreliable

for morphotype discrimination. Regardless, Corbicula

Form D is known from a larger area than reported by

Tiemann et al. (2017). It seems likely that it has gone

unnoticed in other areas, including the Mississippi River,

which is a likely dispersal pathway een the specimen

reported by Tiemann et al. (2017) and the Ohio River

specimen we are reporting. With the aid of external

funding, additional field surveys will help determine the

validity of this claim.

ACKNOWLEDGMENTS

K.S. Cummings offered constructive criticism and pro-

vided the shell photographs in Figure 2. R. Vinsel pro-

vided the hinge photo.

EME RARURE CIE D

Bates, J.M. 1962. Extension of the range of Corbicula fluminea

within the Ohio Drainage. The Nautilus 76: 35-36.

Britton, J.C. and B. Morton. 1986. Polymorphism in Corbicula

fluminea (Bivalvia: Corbiculoidea) from North America.

Malacological Review 19: 1-43.

Counts, III, C.L. 1986. The zoogeography and history of the

invasion of the United States by Corbicula fluminea

(Bivalvia: Corbiculidae). American Malacological Bulletin,

Special Edition 2: 7-39.

Cummings, K.S. and C.A. Mayer. 1992. Field guide to fresh-

water mussels of the Midwest. Illinois Natur: al History

Survey, Manual 5. 194 pp.

Fechtner, F.R. 1962. Corbicula fluminea (Miiller), from the

Ohio River. The Nautilus 75: 126.

Hedtke, S.M., M. Glaubrecht, and D.M. Hillis. 2011. Rare gene

capture in predominantly androgenetic species. Pro-

ceedings from the National Academy of Sciences 108:

9520- 9524.

Hillis, D.M. and J.C. Patton. 1982. Morphological and elec-

trophoretic evidence for two species of Corbicula (Bivalvia:

Corbiculidae) in North America. American Midland Nat-

uralist 108: 74—80.

Hoagland, K. E. 1986. Unsolved problems and promising ap-

‘proaches i in the study of Corbicula. American Malacological

Bulletin, Special Edition 2: 203-209.

Lee, T., S. Siripattrawan, C. Ituarte, and D. O Foighil. 2005.

Hee of the clonal clams: Corbicula lineages in the New

World. American Malacological Bulletin 20: 113-122.

Pigneur, L.M., J. Marescaux, K. Roland, E. Etoundi, J. P. Descy,

and K. Van Doninck. 2011. Phylogeny and androgenesis in

the invasive Corbicula clams (Bivelkia. Corbiculidae) in

western Europe. BMC Evolutionary Biology 11: 174.

Pigneur, L.M., E. Etoundi, D.C. Aldridge, J. Marescaux,

N. Yasuda, and K. Van Doninck. 2014. Gencue uniformity

and long-distance clonal dispersal in the invasive androge-

netic Osriritenle clams. Molecular Ecology 23: 5102— 5116.

Stein, C.B. 1962. An extension of the known range of the Asiatic

clam Corbicula fluminea (Miiller) in the ‘Ohio and Mis-

sissippi Rivers. Ohio oor: a Science 62: 326-327.

Tiemann, J.S., A.E. Haponski, S.A. Douglass, T. Lee, K-.S.

Cummings, M.A. Davis, anal + O Foighil. 2017. First

record of a putative novel invasive Corbicula lineage dis-

covered in the Illinois River, Illinois, USA. BioInvasions

Records 6: 159-166.

Jeremy Tiemann

Illinois Natural History Survey

Prairie Research Institute

University of Illinois

1816 South Oak Street

Champaign, IL 61820 USA

[email protected]

Clarissa Lawlis

Lewis Environmental Consulting, LLC

223 Rocky Lane

Farmington, KY 42040 USA

Sarah Douglass

Illinois Natural History Survey

Prairie Research Institute

University of Illinois

1816 South Oak Street

Champaign, IL 61820 USA

THE NAUTILUS 132(1):33-34, 2018

Page 33

Research Note

Flabellina McMurtrie, 1831: The earliest

valid Latinization?

Authorship and dating of the nudibranch genus-level

name Flabellina is dependent upon the first Latiniza-

tion of the French vernacular name ‘Les Flabellines’

introduced and used by Cuvier (1830) in the second

edition of his Le regne animal... Lemche (1964) traced

a Latinization to Voigt (1634: 124) in the German edition

of Cuvier’s work and, in due course, Flabellina Voigt, 1834

was added to the Official List of Generic Names in Zo-

ology, as Name 1721 (ICZN, 1966).

However, following Winckworth (1941) and Cowan

(1969a; 1969b), Burn (2006) attributed Flabellina to

Griffith and Pidgeon, 1833, certainly to December of that

year, but possibly as early as September (Cowan, 1971).

The online World Register of Marine Species (Caballer

et al., 2014) and Miollinrenitnes (2018) as well as the recent

taxonomic re-assessment of the family Flabellinidae

(Korshunova et al., 2017) list J.E. Gray, 1833 as the author

of Flabellina, an incorrect attribution, as Gray had no

responsibility for the English edition of Cuvier’s text (Petit

and Coan, 2008: Petit, 2012). ). J.C. Smith (1993: 188) noted

a translation into English by McMurtrie (1831) as “the

basis of most of the English translations” of Cuvier.

Reference to McMurtrie (1831) reveals that the latinized

genus name Flabellina occurs on page 343 of volume 2

(Figure | ), the title page of which is dated 1831. This year

1S eonltianeal in the publication registration statement

printed on the reverse of the title page: “Entered

according to the act of congress, in the year one thousand

eight hundred and // thirty one, by G. + C. + H.Camill, in

the clerk’s office of the southern district of // New York.”

Despite his Latinization of the vernacular name,

McMurtrie still considered Cuvier as the author of Flabellina.

But in fact that name should be credited to McMurtrie

alone as he was the first to adopt it. “Adopt”, as defined

in the glossary of the International Code of Zoological

Nomenclature means “To use an unavailable name as

GASTEROPODA NUDIBRANCHIATA. 843

SPLABELLINA, Cuv.

The tentacula of the Eolidx, with radiating rectiform branchiz,

supported by five or six pedicles on each side; they are closely allied

to the Glauci, and in fact to all the Nudibranchiata, whose branchiz

are situated on the sides of the back(1).

(1) Doris affinis, Gm., Cavol., Polyp. Mar, VI, 4

Figure 1. McMurtrie’s (1831) Latinization, genus description

(at top of page 343) and type designation of Flabellina (at bottom

of page 343). The intervening text, containing translation of

other genera and their footnotes, has been oxaindle d.

a valid name of a taxon in a way which established it as

a new name with its own authorship and date” (ICZN,

1999: 99).

The short timeframe between publication of Cuvier

(1830) and McMurtrie’s English edition (1831) makes it

highly unlikely that an rien valid Latinization will

he. found (but see note in Acknowle dgments below).

Flabellina McMurtrie, 1831 thus appears to encompass

the earliest Latinization, author, and date. Doris affinis

Gmelin, 1791, the only species included by Cuvier (1830)

in his “Les Blabellness , was faithfully translated as the

only species in Flabellina by McMurtrie (1831). Doris

affinis Gmelin, 1791 is therefore type species, by mon-

otypy, of Flabellina McMurtrie, 1831.

Accordingly, corrections now need to be made to the

entry on ane Official List of Generic Names in Zoology,

and on the MolluscaBase/World Register of Marine

Species website.

ACKNOWLEDGMENTS

Correspondence with Charles Cowan and Richard Petit,

both regrettably deceased, concerning the Griffith and

Pidge -on translation of Cuvier (1830) was informative and

greatly appreciated. Gemma Steele, Librarian, Museums

Vitctorta. and other library staff, exceeded all expectations

by securing the McMurtrie translation, and those refer-

ences irewards my personal library. Gary Rosenberg (in

litt.) provided a reference to an earlier latinization of ‘Les

Flabellines’ (Van der Hoeven 1830), but this was un-

accompanied by a description or bibliographic reference,

and is therefore a nomen nudum not altering the in-

formation presented above. He also noted that no other

names of mollusks appear to have been introduced

in McMurtrie’s work. I am also grateful for the help-

ful comments of an anonymous reviewer. Richard

Willan graciously prepared and presented this note for

publication.

LITERATURE CITED

Burn, R. 2006. A checklist and bibliography of the Opistho-

branchia (Mollusca: Gastropoda) of Victoria and the Bass

Strait area, south-eastern Australia. Museum Victoria Sci-

ence Reports 10: 1-42.

Caballer, M., P. Bouchet, G. Rosenberg, and S. Gofas. 2014.

Flabellina Gray, 1833. In: MolluscaBase (2017). Accessed

through: World Register of Marine Species at http:/howw.

marinespecies.org/aphia. php ?p=taxdetails bid = 138019

[accessed 27 November 2017].

Cowan, C.F. 1969a. Notes on Griffith's Animal Kingdom of

Cuvier (1824-1835). Journal of the Society for the Bibli-

ography of Natural History 5(2): 137-140.

Cowan, C.F. 1969b. Griffith’s Animal Kingdom. Coe o the

Society for the Bibliography of Natural History 5(3): 249.

Page 34

Cowan, C.F. 1971. Griffith’s Cuvier and Guerin’s Iconographie.

Journal of the Society for the Bibliography of Natural

History 5(6): 498.

Cuvier, G. 1830. Le régne animal distribué d’apres son orga-

nisation, pour server de base a |histoire naturelle des

animaux et introduction a l’anatomie compare. Nouvelle

edition, revue et augmentée. Vol. 3. Deterville, Paris. xvi +

504 pp.

Griffith, E. and E. Pidgeon. 1833. The Mollusca and Radiata. In:

sriffith, E. (ed.), 1824-1835. The animal kingdom arranged

in conformity with its organization, by the Baron Cantar

member of the Institute of France, +C. +C. +C. with

supplementary additions to each order, by Edward Griffith,

F.L.S., A.S., corresponding member of the Academy of

Natural Sciences of Philadelphia, +C. and others. Vol. 12

[((1) = Part 38]. Whittaker and Co., London. 192 pp.

I.C.Z.N. {International Commission on Zoological Nomencla-

ture]. 1966. Flabellina Voigt, 1834 (Gastropoda): placed on

the Official List of generic names. Bulletin of Zoological

Nomenclature 23(2—3): 104-105.

I.C.Z.N. [International Commission on Zoological Nomencla-

ture]. 1999. International Code of Zoological Nomenclature,

Fourth Edition. The International Tiatet for Zoological

Nomenclature, London, 306 bp

Korshunova, T., A. Martynov, Bakken, J. Evertsen, kK.

Fletcher, LW. Mundianta, H. Laas K. Lundin, M. Schrédl,

and B. Picton. 2017. Polyphyly of the traditional family

Flabellinidae affects a major group of Nudibranchia:

aeolidacean taxonomic reassessment with descriptions

of several new families, genera, and species (Mollusca,

Gastropoda). ZooKeys Mie 1-139),

Lemche, H. 1964. Flabellina Voigt, 1834 (Gastropoda): pro-

posed addition to the Official List of Generic Names.

Bulletin of Zoological Nomenclature 21: 120-122.

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McMurtrie, H. 1831. The animal kingdom arranged in con-

formity with its organization by the Baron Cuvier. The

Crustacea, Arealnitictes and Insecta by P. A. Latreille.

Translated from the French, with notes and additions. Vol.

2. G. +C. +H. Carvill, New York, xv + 475 pp + 10 pls.

MolluscaBase. 2018. Flabellina Gray, 1833. Accessed through:

World Register of Marine Species at: http./hvww.marinespecies.

org/aphia.php?p=taxdetailstid=138019 [accessed 14

February 2018].

Petit, R.E. 2012. John Edward Gray (1800-1875): his malaco-

logical publications and molluscan taxa. Zootaxa 3214:

=),

Petit, R.E. and E.V. Coan. 2008. The molluscan taxa made

available in the Griffith + Pidgeon (1833-1834) edition of

Cuvier, with notes on the editions of Cuvier and On Wood’s

Index Testaceologicus. Malacologia 59: 219-264.

Smith, J.C. 1993. Georges Cuvier. An annotated bibliography of

his published works Smithsonian Institution, Washington,

D.C. + London. xx + 251 pp.

Van der Hoeven, J. 1830. Handboek der dierkunde, of grond-

beginsels der natuurlijke geschiedenis van het Dierennijk,

Vol. 2. Volledige Weergave.

Voigt, F.S. 1834. Das Thiol ht, geordnet nack seiner Orga-

nization. Vol. 3. Die Nrolluelent enthaltend, xviii + 621 pp.

F. A. Brockhanss Leipzig.

Winckworth, R. 1941. The name Cratena. Proceedings of the

Malacological Society of London 24: 146-149.

Robert Burn

Marine Invertebrates

Museums Victoria

GPO Box 666 Melbourne

Victoria 3001, AUSTRALIA

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