1VIBL/WH01

THE NAUTILUS

Volume 108

1994-1995

AUTHOR INDEX

Allmon, W. D 15 Lanfair, G. A 25

Dayan, N. S 49 Marshall, B. A 1, 27, 83

Dillon, R. T. Jr 49 McLean, J. H 39, 80

Fell, P. E. 42 Morris, P. J 15

Filippova, Yu. a 67 Nesis, K, N 67

Gordon, ME 55 Nikitina, I. V 67

Harasewych. M G 61 Pierce, H G 23

Helvenston, L. L 42 Smith, ME 25

HouART, R 27 Spelke, J A 42

Kabat, a. R 61 TippETT, D. L 37

Khromov, D. N 67 Tucker, J. K 37

KiLBURN, R. N 34 Yokes, HE 9

Kohl, B 9

NEW TAXA proposed IN VOLUME 108 (1994-1995)

gastropoda

Osteopeltidae

Osteopelta praeccps Marshall, 1994, new species 3

Skeneidae

Bruceiella laevigata Marshall, 1994, new species 5

Bruceiella pruinosa Marshall, 1994. new species 5

Trochidae

Costomargarites McLean, 1995, new subgenus 80

Margarites (Costomargarites) haxteri McLean, 1995, new name 81

Calliostomatidae

CaUiostoma (Otukaia) alcrtae Marshall, 1995, new name 117

Calliostoma (Maurea) antipodense Marshall, 1995, new species 105

CaUiostoma (Maurea) aupourianum Marshall, 1995, new species 97

Calliostoma (Maurea) eminens Marshall, 1995, new species 100

Calliostoma (Maurea) gibbsorum Marshall, 1995, new species 93 '

Calliostoma (Maurea) jamiesoni Marshall, 1995, new species 93

Calliostoma (sensu lata) hopua Marshall, 1995, new species 123

Calliostoma (serisu lata) limatulum Marshall, 1995, new species 122

Calliostoma (Maurea) maui Marshall, 1995, new species 106

Calliostoma (Maurea) penniketi Marshall, 1995, new species 115

Calliostoma (Maurea) regale Marshall, 1995, new species 96

Calyptraeidae

Grandicrepidula McLean, 1995, new subgenus 80

Naticidae

Bentliobulbus McLean, 1995, new genus 39

Euiimidae

Pseudosabinella McLean, 1995, new genus 39

Muricidae

Poirieria syrinx Marshall & I louavt, 1995, new species 30

Ocenotrophon McLean, 1995, new genus 40

Buccinidae

Retimohnia McLean, 1995, new geiuis 40

Mitridae

Charitodoron rosadoi Kilburn, 1995, new species 34

Turridae

Pseudotaranis McLean, 1995. new genus 81

Leucosijrinx kantori McLean, 1995, new name 81

Xylodisculidae

Xylodiscula osteophila Marshall, 1994. new species 7

THE NAUTILUS

Volume 108, Number 1

June 17. 1994

ISSN 0028-1344

A quarterly devoted

to malacology.

Marine Biologicai Laboratoiy/

Woods Hole Oieanographic Institution

Library

JUN 3 0 1994

Woods Hots. MA 0;i543

EDITOR-IN-CHIEF

Dr. M. G. Harasevvych

Division of Mollusks

National Museum of

Natural History

Sniitlisonian Institution

Wasliington, DC 20560

ASSOCIATE EDITOR

Dr. R. Tucker Abbott

American Malacologists, Inc.

P.O. Box 2255

Melbourne, FL 32902

CONSULTING EDITORS

Dr. Riidiger Bieler

Department of Invertebrates

Field Museum of

Natural History

C;hicago, IL 60605

Dr. Robert T. Dillon, Jr.

Department of Biology

(lollege of Charleston

Charleston, SC 29424

Dr. William K. Emerson

Department of Living Invertebrates

The American Museum of Natural

History

New York, NY 10024

Dr Robert Hershler

Division ol Mollusks

National Museum of

Natural History

Smithsonian Institution

Washington, DC 20560

Mr. Richard 1. Johnson

Department of Mollusks

Museum of Comparative Zoolog\

Harvard University

Cambridge, MA 02138

Dr. .\urele La Rocque

Department of Geology

The Ohio State University

Columbus, OH 43210

Dr. James H. McLean

Department of Malacology

Los Angeles County Museum of

Natural History

900 Exposition Boulevard

Los Angeles, CA 90007

Dr. Arthur S. Merrill

% Department of Mollusks

Museum of Comparative Zoology

Harvard University

Cambridge, MA 02138

Ms. Paula M. Mikkelsen

Harbor Branch Oceanographic

Institution, Inc.

Ft. Pierce, FL 33450

Dr. Donald R. Moore

Division of Marine Geology

and Geophysics

Rosenstiel School of Marine and

Atmospheric Science

University of Miami

4600 Rickenbacker Causeway

Miami, FL 33149

Dr. Gustav Paulay

Marine Laboratory

University of Guam

Mangilao, Guam 96923

Mr. Richard E. Petit

P.O. Box 30

North Myrtle Beach, SC 29582

Dr. P'dward J. Petuch

Department of CJeology

Florida Atlantic University

Boca Raton, FL 33431

Dr David H. Stansbery

Museum of Zoology

The Ohio State Universit>

Columbus, OH 43210

Dr. Ruth D. Turner

Department of Mollusks

Museum of Comparative Zoology

Harvard University

C:ambridge, MA 02138

Dr. Geerat J. Vermeij

Department of Geology

University of California at Davis

Davis, CA 95616

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T H E t^N AU T I L U S

CONTENTS

Volume 108, Number 1

Jiiiw 17. 1994

ISSN 0028-1344

Bruce A. IHarshall

Deep-sea Gastropods from the New Zealand Region

Associated with Recent Whale Bones and an Eocene Turtle .

Barry kohl

Harold E. Yokes

On the Living Habits of Aceita buUiai (\'okes) in

Cheniosv nthetic Bottom Communities, Gulf of Mexico

Paul J. Morris

^ arren D. Allnion

Shell Alignment for the Morphometric Analysis of High-

spired Gastropods

15

Harold G. Pierce

On Planorbula nebraskensis Leonard, 1948, and its

position in the evolutionar\ sequence of North American

planorbulids

23

Michael E. Smith

Gerrv A. Lanfair

Effects of Preservati\es on Wet-weight Biomass of the

Asiatic Clam, Corbicula fluminca

25

Marine Blolooical Laboratory/

Wood! Hole Oteanographic IrtttitutJon

Liiirary

JUN 3 0 1994

Woods Hot*. MA 02543

THE NAUTILUS 108(1):1-S, 1994

Page 1

Deep-sea Gastropods from the New Zealand

Region Associated with Recent Whale

Bones and an Eocene Turtle

Bruce A. Marshall

Museum of New Zealand Te Papa

Tongarevva

P.O. Box 467

Wellington, New Zealand

ABSTRACT

Five species of gastropods are new iy recorded from deca> ing

whale bone from the deep-sea floor off New Zealand: Para-

cocculina cervae (Fleming, 1948). and new species of Osteo-

pelta. Marshall, 1987 (Cocculiniformia), Brucciclla Waren &

Bouchet, 1993 (Vetigastropoda), and Xylodiscula Marshall, 1988

(Heterobranchia). RecentK discovered material of Osteopelta

mirabilis is recorded, and this or a closely similar species is

recorded associated with turtle bones from the Middle Eocene

of New Zealand. The new Bruceiella and Xyhidiscula species

are the first records of skeneimorph gastropotis from bone.

.\part from w hale bone, Paracocculina cervae is associated with

sunken wood and algal holdfasts (nev\ record), so it has the

most generalised habitat of any know n cocculiniform limpet.

Key words: Recent, fossil, Mollusca, Gastropoda, deep-sea,

w hale bone, turtle bone. New Zealand.

INTRODUCTION

Since discovery of the limpet Osteopelta mirabilis Mar-

shall, 1987, the first record of a gastropod living in as-

sociation with decaying whale bones, a second species of

Osteopelta has been recorded from bone off Iceland

(Waren, 1989). More recently McLean (1992) has re-

corded a nev\ species of Coeeiilina Dall, 1882 (Coccu-

linidae) and three species of Pywpelta McLean & Hasz-

prunar, 1987 (Pyropeltidae) from whale bone from the

eastern Pacific. One of these pyropeltiii limpets was de-

scribed as new, w hile the other two were evidently con-

specific w ith specimens originally obtained from hydro-

thermal vents (McLean & Haszprunar, 1987). Dell (1987)

has reviewed my tilid bivalves associated with whale re-

mains. Additional ta.xa have been recorded by Smith et

al. (1989), who drew attention to similarities between

faunas associated with decaying whale bones, hydro-

thermal vents and hydrocarbon seeps, and suggested that

whale skeletons may provide stepping stones for wide

dispersal of deep-sea chemosynthetic communities (see

also Smith, 1992).

In this paper I introduce a third species of Osteopelta.

another cocculinid, and species of Bruceiella \\'aren &

Bouchet, 1993 and Xylodiscula Marshall, 1988 new to

the fauna of this unusual habitat Bruceiella was based

on a species from a hydrothermal vent, while previously

known Xylodiscula species occurred in association with

sunken wood, old sea grass (Posidonia) fibres, and a hy-

drothermal vent.

ABBREVIATIONS

.\MS .Australian Museum, S\dne\

BMNH The Natural History Museum, London

L.\CM Los ,\ngeles Count>- Museum of Natural His-

tor\'

MNHN Museimi National d'Histoire Naturelle, Paris

MNZ Museum of New Zealand, Wellington

NMP Natal Museum, Pietermaritzburg

OL' Geology Department, Uni\ersit\ of Otago,

Dunedin

USNM National Museum of Natural History, Wash-

ington, DC

Order ARCHAEOGASTROPODA Thiele, 1925

Suborder COCCULINIFORMIA Haszprunar, 1987

SuperfamiK COCCULINOIDEA Dall. 1882

FamiK COCCULINIDAE Dall, 1882

Genus Paracocculina Haszprunar, 1987

Paracocculina Haszprunar, 1987:321. Type species (by original

designation!: Cocculina laevis Thiele, 1903: Recent, off

Nias Island, Sumatra.

Remarks: The genus Paracocculina was introduced by

Haszprunar (1987) for species that differ from Cocculina

Dall, 1882 primarily in having a prominent subpallial

gland, a pedalK innervated copulator\ organ on the right

side of the foot, and a hypoathroid nervous system with

the osphradial ganglion situated at the far left side.

Page 2

THE NAUTILUS, Vol. 108, No. 1

Figure 1. Map of New Zealand region showing localities for

Osteopelta praeceps n. sp. and O. mirahilis (stars), BruceieUa

laevigata n. sp. (solid circle), and BruceieUa pruinosa n. sp.

(open circle), 200 and 1,000 meter contours indicated.

Paracocculina eervae (Fleming, 1948)

Cocculina eervae Fleming, 19-48:88, text fig. la-d; Marshall,

1986:508, figs. 2A, 3A-C, 12AB.

Tecticrater eervae. Dell, 19.56:60; Powell, 1979:81, fig. 10/9

Paraeoeeulina eervae. Haszprunar, 1987:321.

IVlaterial examined: (Additional to that recorded bv

Marshall, 1986); 39°53.2'S, 168°01.2'E, Challenger Pla-

teau, New Zealand, alive on whale bone, 908-912 m, 3

July 1989, f.v. Amaltal Explorer (3 MNZ); 44°40.5'S,

174°01.0'E, off Banks Peninsula, alive on whale skull, 844

m, 11 May 1987, f.v. Oijcing 7 (5 MNZ); 38°38.9'S,

178°38.3'E, off Gable End Foreland, New Zealand, alive

on algal holdfast, 529-565 ni, 23 September 1985, f.v.

Wanaka stn WK2/68/85 ( 1 MNZ); 43°02.5'S, 174°09.0'E,

off Kaikoura, New Zealand, alive on algal holdfast, 848-

877 m, 24 September 1988, f.r.v. James Cuok stn J12/

10/88 (7 MNZ); ,50°02.6'S, 169°31 'F, east of the Auckland

Is, New Zealand, alive on algal holdfast, 614-620 m, 4

December 1992, f.r.v. Tangaroa stn 92011/97 (many

MNZ).

Distribution: North ('ape to the .Auckl;

ds, Nt

Zealand, 18-891 ni, on sunken wood, algal holdfasts and

whale bone.

Remarks: Specimens from whale bone and deep-sunken

algal holdlasts and wood proved to be indistinguishable

in shell and radular morphology and in e.xternal anatom\ .

Superfamilv LEPETELLOIDEA Thiele, 1908

Family OSTEOPELTIDAE Marshall, 1987

Genus Osteopelta Marshall. 1987

Osteopelta Marshall, 1987:121. T\pe species (by original des-

ignation): Osteopelta mirabilis Marshall, 1987; Recent,

New Zealand.

Osteopelta mirabilis Marshall, 19S7

Osteopelta mirabilis Marshall, 1987122, figs. lA. B-D, F-J,

2A, 2B, 3C; Haszprunar, 1988:6, figs. 1.5-24 (anatomy).

Material examined: (.ALilditional to that recorded by

Marshall, 1987); .39°.53.2'S, 168°01.2'E, Challenger Pla-

teau, New Zealand, alive on whale bone, 908-912 m, 3

July 1989, f.v. Amaltal Explorer (18 MNZ); off Mernoo

Bank, Chatham Rise, New Zealand, alive on large whale

skull, ca. 900 m, October 1988, f.v. Amaltal Explorer (5

MNZ).

Distribution (figure 1): Challenger Plateau and Chat-

ham Rise, New Zealand, on whale bone, 800-955 m.

Remarks: The Challenger Plateau specimens are in-

distinguishable from type and other material from the

Chatham Rise. The protoconch is retained in a specimen

5,00 mm long, and is bilateralK symmetrical and 200

/um long, with a long tapered apical fold tip on each side.

Regrettably the sculpture is unknown, as the outer shell

layer has been etched away. The Icelandic species Os-

teopelta ceticola Waren, 1989, also from whale bone,

differs in minor details of radular morphology (Waren,

1989).

Osteopelta sp cf. mirabilis Marshall. 1987

IVlaterial examined: Isolated concretion of Waihao

Greensund on right bank of South Braiiich of Waihao

River, north of Waihao Dow ns (map ref. J40/45000030),

closely associated with bones of fossil leatherback turtle

(Dermochelys sp. — Reptilia, (-helonia. Dermochelyi-

dae), coll. R. E. Fordyce, A. Grebneff, C. M. Jones and

P. A. Maxwell, .^ge: Bortonian (Middle Eocene) (1 spec-

imen Ol'41112).

Remarks: The single specimen is enibeddetl api.'.\ (low n

in a block of gritt\ niatri.x with the full\ exposetl interior

surface showing clearK tlefineil muscle scars. .\ mould

of the exterior where a small piece of the shell margin

has liroken away shows impressions of weak concentric

sculpture. The specimen is indistinguishable from the

Recent specimens of Osteopelta mirahilis in shape, shell

thickness, teleoconch sculpture, muscle scar outline, and

size (length est. 7.50 nun, width (i,05 mm). While it is

B. A. Marshall, 1994

Pages

impossible to be absoluteh certain of the relationships

of this limpet from the teleoconch alone, particularK the

interior, the association with bone suggests that it is prob-

alily an osteopeltid.

The W'aihao Green sand beds w ere considered by Max-

well (1992) to ha\e been deposited at 150-250 ni depth,

which is substantialK shallower than the batlnmetric

range of li\ing O. mirabilif; (800-955 m),

Osteopella praeceps n. sp.

(figures 2-6. 18, 19)

Description: Shell up to 6,25 mm long, translucent white,

thin and brittle, high])- arched, anterior end occup\ing

48.8-62.5% of shell length in specimens over 4 mm in

length. .Aperture elliptical, anterior end more narrowly

rounded than posterior; apertural plane shallowly con-

cave at ends, weakly convex at sides; broadest slightK'

behind midlength. .\pex tightK- rounded, anterior slope

flat or weakK conca\e, posterior slope broadly convex,

lateral slopes weakK con\ex. Periostracuni transparent,

ver> thin, smooth.

Protoconch 200 urn long, bilaterally symmetrical, api-

cal fold producing a small, shallow dimple on each side,

sculptured throughout with densely crowded minute

punctations arranged in spiral lines,

Teleoconch sculptured with fine collabral grciw tli lines,

obscure radial lines, and many minute, randomly dis-

tributed, shallow pits.

Animal white. Foot broad and thick. Mantle edge

thickened, with weak left anterolateral fold. Snout very

large, tapered, longer than broad, deeply concentrically

wrinkled, mouth a vertical slit in small, circular, flattened

tip. No oral lappets. Cephalic tentacles slender, tapered,

tips rounded, minute black eyes sunken in swellings at

outer bases. A large secondary gill extending along right

mantle groove to above right of head, comprising about

30 anteriorly enlarging leaflets. Seminal groove not de-

tected. Two slender blunt-tipped, dorsoventrally flat-

tened epipodial tentacles at posterior end.

Radula (figures 18, 19) with the formula oo -I- 6 + 1

-H 6 + oo, indistinguishable from that of O. mirahilis.

Type data: Holot\'pe MXZ Ml 16970 (length 5.50 mm,

width 4.20 mm, height 3.40 mm) and 57 paratypes (51

MNZ; others AMS, BMNH, LACM, MNHN, NMP,

USNM): 43°34.14'S, 176°18.69'E, E of Mernoo Bank,

Chatham Rise, New Zealand, 372-379 m, 29 January

1992, f.r.v. Tangaroa stn 9106 168, alive on whale ver-

tebra among crowded indi\iduals of a species of Idas

Jeffreys, 1876 (Mytilidae).

Other material examined: (Several hundred specimens

MNZ): Topotypes (several hundred juveniles, MNZ

M. 117279); 39°53.2'S, 168°01.2'E, Challenger Plateau,

New Zealand, 908-912 m, 3 July 1989, f.v. Amaltal Ex-

plorer stn 34S ' 1 29, alive on a piece of whale bone among

crowded mytilids (Idas sp. ) (8 subadult specimens, MNZ

M.92453).

Distribution (figure 1): C^hatham Rise and Challenger

Plateau, New Zealand, on whale bone, 372-912 m.

Remarks: Ostcopclta pracccp.s differs from O. mirahilis

in attaining smaller size and in ha\ing a narrower, taller

shell w ith a longer anterior end. Their external anatomies

and radulae are extremely similar. O. praeceps and O.

mirahilis occurred li\ing together on the same piece of

whale bone from the Challenger Plateau (M.92453 and

M. 92451, respectively).

Immature specimens of O. praeceps are strikingly sim-

ilar to young of the cocculinid Paracocculina cervae that

also occurred on the C^hallenger Plateau bone. P. cervae,

however, is distinguishable by its larger adult size, lack

of shell micropunctations, in having a prominent copu-

latory organ behind the right cephalic lappet, and in

having a snout that is short and broad instead of pro-

boscis-like. Moreover, their radulae and anatomies are

entirely different

The highK arched shell and narrow aperture of O.

praeceps enable it to live deeply (and inconspicuously)

among the associated mstilids, which form densely

crowded aggregations at nutrient-rich sites on the bones.

By contrast, the larger and broader-shelled species O.

mirahilis has been obser\ed adjacent to m\ti!id aggre-

gates (Marshall 1987: fig. L\), and is probably unable to

live as deeply among them.

Etymology: Latin praeceps (steep).

Suborder VETIGASTROPODA Salvini-Plawen, 1980

Superfamilv TROCHOIDEA Rafinesque. 1815

.^FamiK SKENEIDAE Clark, 1851

Genus Bruceiella Waren & Bouchet, 1993

Brucciclla Waren & Bniichet, 1993:26. Type species (by orig-

inal designation): Bruceiella globulus Waren & Bouchet,

1993; Recent, North Fiji Basin (hydrothermal vent).

Remarks: In ha\ing epipodial tentacles and papillate

cephalic tentacles, the animal of Bruceiella bears a gen-

eral resemblance to those of members of the family Ske-

neidae, the limits of which have been tightly restricted

by Waren (1992) and Waren and Bouchet (1993). The

lack of a propodial penis and the distally expanded snout,

however, led Waren and Bouchet (1993) to suggest that

the genus probably does not belong in Skeneidae, but

by implication perhaps in a family of its ow n. The radula

bears a striking resemblance to those of a variety of

archaeogastropods associated with hydrothermal vents

and seeps, especially neomphalids and peltospirids, a sim-

ilarity that Waren and Bouchet (1993) consider to be the

result of convergence.

The otherwise simple shell of Bruceiella species is dis-

tinctive in having a protoconch sculpture of minute den-

dritic threads and granules arranged in fine spiral lines,

and a pronounced varix almost immediately after the

protoconch-teleoconch boundary. Unlike Waren and

Bouchet (1993), who interpret the varix as the termi-

nation of the protoconch, I consider that the \arix is on

the teleoconch. and that the actual protoconch teleo-

Page 4

THE NAUTILUS, Vol. 108, No. 1

Figures 2-14. Shells of Osteopelta and Bruceiella spj) 2-6. Osliopilla pnieccps ii- sp, 2, .1. Holot\pe, length 5,50 nnn. 4.

Immature specimen, Challenger Plateau, MNZ M,9245;5, length 2.20 mm. 5, 6. Protoconch of liolot) pe. 7-9. BruccicUa laevigata

n, sp., holot\pe, height 1.70 mm. 9. Protoconch, with protoconch /teleoconch boundary arrowed. 10-14. Bruceiella prninosa n.

sp., holotype, height 1.43 mm. 10. Detail of inner part of base. 13. Protoconch. with protoconch teleoconch bonndar\ arrowed.

14. Detail of protoconch sculpture. Scale bar 14 = 10 ^m, others = 100 fim.

B. A. Marshall, 1994

Page 5

Figure§ 15-17. Shell of holntvpt- (it XijIotliMula u^teopliila u sp., width l,(iO mm 17. Protdcunch, note inroiled tip. Scale bar

100 Mm.

conch boundary is a sharpK defined line at a distance

behind the vari.x equi\ aleiit to about one vari.x thickness

(figures 9, 13). I also disagree with Waren and Bouchet

(1993) that the varix in their genus VeiUsia is part of the

protoconch and instead consider that it is situated at the

end of the first quarter teleoconch whorl. Species of the

related genus Xyloskenea Marshall, 1988 exhibit a change

in teleoconch sculpture following a sharply defined growth

scar at an equivalent position (Marshall, 1988, figs: 4EJ,

5E; Waren & Bouchet, 1993: fig. 23A). The zone between

the varix and the protoconch/ teleoconch boundary is

analogous to protoconch II of higher gastropods, though

not homologous because protoconch enlargement through

marginal incrementation is unknown in archaeogastro-

pods other than Neritomorpha. These growth distur-

bances probably represent a period of crisis, perhaps a

change in feeding mode during the earliest stages of

postsettlement development

Bruceiella laevigata n. sp.

^figures 7-9, 20-22)

Description: Shell up to 1.70 mm higli, slightK higher

than broad, thin, translucent, gloss\ , with narrow um-

bilical chink, periostracum smooth.

Protoconch 300 fxm w ide, delineated b\ fine groove,

apical fold tip broadly roundetl, sculptured w ith minute

anastomosing dendritic threatis arranged in fine spiral

lines.

Teleoconch of up to 2.20 strongh and rather evenly

con\ ex whorls. First sixteenth w horl minuteU granulate;

next sixteenth whorl occupied b\ strong, rounded \'arix,

immediateK follow ed b\ fine groo\ e; elsew here smooth.

.\perture subcircular, lips thin, parietal contact area nar-

row.

.Animal. Snout subquadrate; cephalic tentacles dorso-

ventralK flattened, tapered, similar, edges ciliated; foot

large, anteriorly indented. Small right and large left sub-

optic tentacle, 3 small right epipodial tentacles beside

operculum, 1 large left epipodial tentacle, 2 small left

epipodial tentacles beside operculum. No eyes.

Radula (figures 20-22) with the formula co -^ .5 -t- 1

-I- 5 -I- oo, teeth longer than broad. Central tooth stout;

cutting area narrow ly angulate, w ithout secondary cusps,

proininently hooded; shaft face subtriangular, suddenly

narrowed at base. Lateral teeth stout, enlarging out-

wards, cutting areas large, hooded, roundly angulate,

coarsely serrate, terminal cusp largest; shafts outwardly

bowed, strongly flanged and con\'oluted to interlock with

adjacent teeth. Marginal teeth slender, outwardly nar-

rowing and with smaller cutting areas and finer cusps,

outermost few pairs w ith spathulate tips and fused shafts.

Type data: Holotype (height 1.70 mm, diameter 1.50

mm, 2.20 teleoconch whorls) MNZ M. 116969; paratypes

(5 MNZ, 1 AMS, 1 MNHN): 43°00.17'S, 174°05.5'W, NE

of Chatham Islands, New Zealand, 1,242 m, 23 August

1989, f.v. Otago Buccaneer tow 31; alive amongst crowd-

ed mytilids (Idaa sp. ) on a whale skull.

Distribution (figure 1): Northeast of Chatham Islands,

New Zealand, on whale bone, 1,242 m.

Remarks: Compared w ith the ty pe species. Bruceiella

laevigata is more tightly coiled with a higher spire, but

otherw ise the two species are similar. .Among the host of

superficially similar skeneimorph gastropods known from

the New Zealand region (MNZ — many undescribed), B.

laevigata is characterised by the combination of distinc-

tise protoconch sculpture, the strong postlarval varix, and

tlie radular morphology.

Etymology: Latin laecigalus (smooth), alluding to the

lack of shell sculpture after the postlar\al \arix.

Bruceiella pruinosa n. sp

(figures 10-14, 23)

Description: Shell (holotype) 1.43 mm high, slightly

higher than broad, thin, translucent, glossy , with narrow

umbilical chink, periostracum smooth.

Protoconch 280 nm w ide, delineated by fine groove,

apical fold tip broadly rounded, sculptured with minute

anastomosing dendritic threads arranged in fine-spiral

lines.

Teleoconch of 2.10 strongly and rather e\enly convex

Page 6

THE NAUTILUS, Vol. 108, No. 1

B. A. Marshall, 1994

Page 7

whorls. First sixteenth whorl niinutel\ granulate; next

sixteenth whorl occupied by strong, roundeil varix, ini-

mediateK lollowed b\ fine groove. Adapical quarter of

spire whorls, and inner half of base w ith minute granules

and very fine axial w rinkles, stronger on base. Aperture

subcircular, lips thin, parietal contact area narrow.

.Animal. Similar to that of B. laevigata but w ith 1 large

right and 2 large left epipodial tentacles, instead of 4

right and 3 left epipodial tentacles,

Radula (figure 23) as in B laevigata.

Type data: Holot>pe M\Z M.11696S (heigiit 1.43 mm,

diameter 1.33 mm, 2.10 teleoconch whorls): 39°53.2'S,

168''01.2'E. Challenger Plateau, New Zealand, 908-912

m, 3 July 1989, f.v. Amaltal Explorer stn 348/129; alive

amongst crowded mytilids {Idas sp.) on a piece of whale

bone.

Distribution (figure 1): Challenger Plateau, New Zea-

land, on w hale bone, 908-912 m.

Remarks: Bruceiella pniinosa differs from both B. glob-

ulus and B. laevigata in ha\ing a teleoconch sculpture

of minute granules and fine axial w rinkles.

Etymology: Latin pruinosus (frosts), alluding to the

fineK granulate surface.

Subclass HETEROBRANCHIA Grav, 1840

Order HETEROSTROPHA Fischer,' 1885

Family XYLODISCULIDAE Waren, 1992

Genus Xylodiscula Marshall, 1988

Xylodiscula Marshall, 1988:988. T\pe species (by original des-

ignation): Xylodiscula vitrea Marshall. 1988; Recent, New

South Wales.

Remarks: Xylodiscula was originalK referred to Or-

bitestellidae for want of a more appropriate position

(Marshall, 1988). Subsequent reevaluation of Orbitestel-

lidae by Ponder (1990), however, suggested that this

placement is untenable, and VVaren (1992) has segre-

gated Xylodiscula in a family of its ov\n.

The new species described below is the first record of

a xylodisculid from whale bone. Other Xylodiscula spe-

cies live at 90-1,100 m in association w itli sunken wood

(Marshall, 1988; Waren, 1992) and old sea grass (Posi-

donia) fibres (Waren. 1992), Most recentK a Xylodiscula

species has been recorded from a h\drothermal vent at

2,000 m depth in the North Fiji Basin (Waren & Bouchet,

1993),

Xylodiscula osteophila n sp

(figures 15-17, 24, 25)

Description: Shell (holotspe) 1,60 mm wide, markedly

w ider than high, spire weakly elevated, thin, translucent,

colorless, umbilicate, periostracum smooth.

Protoconch 270 /um wide, 1.75 whorls; tip infolded,

ver\ small, finely granulate; last w horl smooth

Teleoconch of 2.25 convex whorls, suture shallowly

channelled. First 1.25 whorls evenly convex; last whorl

w ith weakly convex side, and strongK rounded periph-

ery. Base convex, evenly rounded into umbilicus, obscure

spiral lines throughout. Collabral grow th lines geiUK pro-

socline on spire, weakly sigmoidal on base. Umbilical

diameter 17''t of shell diameter. Aperture subcircular.

Outer lip thin; inner lip thickened and angled against

umbilical rim, thin adapically. Parietal area broad, in-

ductura extremely thin,

■Animal unknown (dried). Operculum thin, translu-

cent, chitinous multispiral,

Radula (figures 24, 25) with the formula 2 -I- 1 4- 0

+ 1-1-2, extremeK small. Lateral teeth small, thin,

subquadrate, cutting area almost straight and finely ser-

rate. Marginal teeth similar, large, stout, slender, curved,

\ er\ long fineK' serrate cutting area on both edges.

Type data: Holot\ pe (height 1,07 mm, width 1,60 mm,

2,25 teleoconch whorls) MNZ M, 116971 off Mernoo Bank,

Chatham Rise, New Zealand, ca. 900 m, October 1988,

f,v, Amaltal Explorer; alive on large w hale skull amongst

crowded m>tilids (Idas sp.).

Distribution (figure 1): Off Mernoo Bank, Chatham Rise,

New Zealand, on v\hale bone, ca. 900 m.

Remarks: Xylodiscula osteophila differs from other

named Xylodiscula species in the greater shell height

relative to width, the thickened and angled inner lip,

and the considerabK narrower umbilicus.

Etymology: From the Greek osteon (bone) and philios

(loving).

ACKNOWLEDGEMENTS

I am grateful to the numerous fisheries scientists and

observers (MAF Fisheries, Wellington) who obtained the

samples of whale bone, to .A. Grebneff (I'niversity of

Otago) who drew my attention to the fossil osteopeltid,

and to J. H. McLean (Los Angeles Count) Museum of

Natural History) for constructive comments on the

manuscript. Thanks also to W. St George (Institute of

Geological and Nuclear Sciences, Low er Hutt) for access

to the scanning electron microscope, D. Wilkinson for

w ord processing, and to T, Meek for photographic print-

ing'

Figures 18-2,5. Radulae ex holotypes. 18. 19. Ostcopclta praeceps n, sp,, full v\idth (18) and detail of central and lateral teeth

(19) 20-22. Brticciclla laevigata n, sp,, full width (20), central and lateral teeth (21) and marginal teeth (22), 23. Bruceiella

pruirjo.'ia n sp , central (left), lateral and inner marginal teeth. 24, 25. Xylodiscula osteophila n, sp,, marginal teeth and the small

central tooth (left center in 24), Scale bars = 10 ^m.

Pages

THE NAUTILUS, Vol. 108, No. 1

LITERATURE CITED

Dell. R, K. 1956. The Arcluhenthal Mollusca of New Zealand.

Ddiiiinion Museum Bulletin l.S:l-234.

Dell, R. K. 1987. Mollusca of tlie famih Mytilidae (Bivalvia)

associated with organic remains from deep w ater off New

Zealand, with revisions of the genera Adipicola Dautzen-

berg, 1927 and Idasola Iredale, 1915. National Museum

of New Zealand Records 3:17-36.

Fleming, C. A. 1948. New species and genera of marine

Mollusca from the Southland fiords. Transactions of the

Royal Society of New Zealand 77:72-92.

Haszprunar, G. 1987. .\natomy and affinities of cocculinid

limpets (Mollusca, Archaeogastropoda). Zoologica Scripta

16:305-324.

Haszprunar, G. 1988. .\natom\ and relationships ot the bone-

feeding limpets CoccuhncUa minutissima (Smith) and Os-

tcopclta mirabilis Marshall (.\rchaeogastropoda). Journal

of Molluscan Studies 54: 1-20.

McLean, J, H 1992. Cocculiniform limpets (Cxicculinidae

and P\ ropeltidae) living on whale bone in the deep sea

off California. Journal of Molluscan Studies 58:401-414.

McLean, J. H. and G. Haszprunar. 1987 P\ ropeltidae, a new

familv of cocculiniform limpets from h\drothermal \ents.

The \'eliger 30:196-205.

Marshall. B. .\. 1986. Recent and Tertiary Cocculinidae and

Pseudococculinidae (Mollusca: Gastropoda) from New

Zealand and New South Wales. New Zealand Journal of

Zoology 12 505-546.

Marshall. B .\ 1987 Osteopeltidae (Mollusca: Gastropoda):

a new family of limpets associated with whale bone in the

deep-sea. Journal of Molluscan Studies 53121-127.

Marshall. B. A. 1988, Skeneidae. Vitriueliidae and Orbites-

tellidae (Mollusca: Gastropoda) associated with biogenic

substrata from bathyal depths off New Zealand and New

South Wales, Journal of Natural History 22:949-1004.

Ma,\well, P, \. 1992. Eocene Mollusca from the vicinity of

McCulloch's Bridge, Waihao River, South Canterbury, New-

Zealand: paleontology and systematics. New Zealand Geo-

logical Survey Paleontological Bulletin 65:1-280.

Ponder, W. F, 1990, The anatomy and relationships of the

Orbitestellidae (Gastropoda: Heteroliranchia). Journal of

Molluscan Studies 56:515-532,

Powell, A. W, B. 1979. New Zealand Mollusca: marine, land

and freshwater shells, Collins, .Auckland. 500 p.

Smith, C. R. 1992. Whale falls. Chemoss nthesis on the deep

sea floor. Oceanus 35:74-78.

Smith, C. R.. H. Kukert. R. A. V\ heatcroft. P .\ Jumars, and

J, W. Deming. 1989. N'ent fainia on u hale remains. Na-

ture 341:27-28.

Waren, A, 1989, New and little know n Mollusca from Iceland.

Sarsia 74:1-28.

Waren, .•\, 1992, New and little known "skeneimorph" gas-

tropods from the Mediterranean Sea and the adjacent At-

lantic Ocean, Bollettino Malacologico 27:149-247,

Waren, A. and P, Bouchet, 1993, New records, species, gen-

era, and a new family of gastropods from hydrothermal

vents and hydrocarbon seeps, Zoologica Scripta 22:1-90.

THE NAUTILUS 108(1):9-14, 1994

Page 9

On the Living Habits of Acesta biiUisi (Yokes) in Chemosynthetic

Bottom Communities, Gulf of Mexico

Barry kohl

Harold E. Yokes

tieulijg) Department

Tulane University

New Orleans, LA 70118 USA

ABSTRACT

The supposedly pathologic paratype of Acesta huUisi

(Vokes,1963) has been re-evaluated based on additional infor-

mation from the type locality The specimen is not aberrant,

as originally thought, but represents a normal mode of growth

In uliich the shell is attached over the anterior end of che-

inos\ nthetic tube worms of ihe g,enus Luim'llihrachia sp., which

occur at Indrocarbon seeps in the Gulf of Mexico, It is shown

thai the paratype is a final stage in the ontogen\ of A. bullisi.

which lives attached to the tube worm Lamcllihrachia.

Key W'urds: Acesta, Chemos> nthetic, Lanicllilirucliia. CUill

of Mexico, tube worms.

INTf^ODUCTION

Hydrothermal vents and their oases of previously un-

described species were discovered in 1977 on dives of

the submersible Alvin along the Galapagos Rift in the

Pacific Ocean. Giant tube worms, large clams, and mus-

sels form an assemblage of invertebrates dependent on

hydrogen sulfide-rich waters. These animals contain bac-

terial symbionts that o.xidize the sulfide.

An assemblage of tube worms, clams, and mussels was

reported in the Gulf of Mexico near a saline seep at the

base of the Florida Escarpment (depth 3,266 meters) by

Paull et al., (1984). This was the first record of vent

organisms occurring on a passive margin. The following

year, Kennicutt et al. (1985) described a fauna associated

with oil and gas seeps from trawls along the Louisiana

continental slope. One trawl contained specimens of

Acesta bullisi and an entanglement of vestimentiferan

tube worms of the genus Lamellibrachia , taken near

Green Canvon Block 234, (27°45'N, 9ri4'W) (Figure

1).

Since the discoveries by Paull ct al and Kennicutt et

al.. there have been many projects designed to document,

by manned-submersibles, the occurrences of chemosyn-

thetic communities in the Gulf of Mexico associated with

oil and gas seeps. Shore-based research is investigating

how these organisins derive their energy from oil, meth-

ane, and sulfides, which are toxic to most invertebrates.

In addition to unicjue chemosynthetic species, there

are opportunistic non-chemosynthetic organisms that use

either the carbonate substrate surrounding the vents or

the chemosv nthetic forms for attachment. Acesta bullisi

is one of these opportunistic organisms found attached

to the tube worm Lamellibrachia sp. at many sites in the

Gulf of Mexico (Carnev, 1992; MacDonald & Ambler

1992).

The supposedly "pathologic " hypotype [paratype] of

the species described as Lima (Acesta) bullisi by Yokes

(1963, pi. 2, figs. 1-4), was given to H. E. Yokes in 1962

by H. R Bullis, Jr. It was alive when retrieved in a trawl

from the R/Y Oregon Station 3741, at a depth of 548

meters in the Gulf of Mexico (29°10'N, 88°01.5'W).

This location can be equated to a sampling station in

Viosca Knoll Block 826 (YK 826, Figure 1), 29°11.0'N,

88°00'W, at a depth of 545 meters (L' S. Dept of Interior,

1992, vol. 1, table 1) used by the Geochemical and En-

vironmental Research Group (GERG) of Texas A & M

University. According to MacDonald and Ambler (1992),

this location is the most easterly site within the Gulf of

Mexico of chemosynthetic communities associated with

hydrocarbon seep activity found to date. The chemo-

synthetic fauna at the YK 826 site is represented by living

tube worms {Lamellibrachia sp. and Escarpia sp) and

]X)ssibl\- lucinid clams (MacDonald & Ambler, 1992).

The purpose of this paper is to document that the

paratype of Acesta bullisi described by Yokes in 1963

was a form attached to tube worms from a chemosyn-

thetic commimity. We hope that this paper will also

stimulate an investigation of other "deformed mollusk

species in collections and thereb\ identify locations of

other chemosynthetic communities.

MATERIALS AND METHODS

Specimens of Acesta bullisi were collected by personnel

from Tulane University and Louisiana State University

(LSL') on cruises in the Gulf of Mexico during September

1992 and July 1993. The Johnson Sea-Link (JSL) I &

II, manned-submersibles, were used for bottom sampling

on a total of 24 dives. Photographs and video were taken

Page 10

THE NAUTILUS, Vol. 108, No. 1

Figure 1. Location map showing Garden Banks Block 427,

Green C'anvon Blocks 18.5, 234 and Viosca Knoll Block 826.

of clumps of Lamellibrachia and the attached individuals

of Acesia. Specimens of Acesta bullisi used for the pres-

ent study were collected for research at LSI' and were

photographed on shipboard by B. Kohl to document the

living position of Acesta in relationship to the tube worm

Lamellibrachia. A collection of invertebrates retrieved

on the dives is maintained at LSU.

The sites used for this paper are Green Canyon Block

185, 27°46.93'N, 91°.30.47'W, dive 3301 (1992), depth

541 meters, and Garden Banks Block 427, 27°33.61'N,

92°25.09'\V, dives 3305 (1992) and 3568 (1993), depth

611 meters, (Bgure 1). Ten specimens of A. bullisi at-

tached to Lamellibrachia were retrieved by the JSL's

mechanical arm by selectively collecting individual tube

worms.

RESULTS

Biology of Tube Worms

Lamellilnachia spp., classified as vestimentiferans, lack

a mouth, anus and digestive system. They live in a tough,

thick-walled, chitinous tube about one centimeter in di-

ameter and up to two meters in length. The posterior

end may either be buried in soft sediments or attached

to a rocky substratum. A small red-orange plume, the

obturaculum, (approx. one cm in diameter) is extended

from the anterior opening and serves as a gas exchange

organ. The plume can be withdrawn into the tube at will

(Fisher, 1992; MacDonaid et a/., 1990). Most of the body

volume consists of a trophosome, the organ containing

the bacterial symbionts. The animal absorbs HjS, CO2

and O2 through its obturaculum (plume) and, through a

complex chemical process, the symbiotic bacteria and

enz\ nies convert CO2 to organic carbon. See Fisher (1990)

for a detailed review of the chemistry involved.

New research has suggested that the O2 is absorbed

through the plume and H2S is absorbed through the

posterior part of the tube, which is located below the

sediment,' water interface. It has been observed that the

living animal extends the entire length of the tube. Ab-

sorption of CO2 possibly takes place along the length of

the tube above the sediment/water interface through the

porous chitinous wall (R. S. Carney, personal commu-

nication).

Relationship of A. bullisi to L.\mellibrachl\ sp.

Specimens of A. bullisi attach to tall individuals of La-

mellibrachia approximately one-half meter above the sea

bottom, near the anterior end of the tube. Specimens

collected on the 1992 and 1993 dives show that A. biillisi

frequently occurs attached to the "bushlike ' congrega-

tions of the tube worm Lamellibrachia sp This was also

reported and illustrated by MacDonaid £>/ al. (1989) from

the Green Canyon Block 184-185 area.

Two attachment positions of A. bullisi were observed

on our dives. The first is an attachment by a b\ssus to

the side of the tube worm near the upper portion of the

tube (figure 3). These individuals do not have the notch

or opening and are about half the size of the adult shown

in figures 2 and 4.

The second attachment position can be seen in the

specimens in figure 2, which completely envelop the

anterior region of Lamellibrachia. A detail of one of the

specimens from figure 2 is shown in figure 4. These

specimens are attached by a byssus to the exterior of the

tube worm very near the anterior end.

The shape of the growth lines in the valves of the adult

A. bullisi (figure 9) implies that the opening in the shell

developed late in ontogeny.

DISCUSSION

The specimen described by H. E. Yokes was thought to

have been damaged at a time when the animal was half

grown (Yokes, 1963; 78). Observations of the living an-

Figure§ 2— t. Living specimens of Acesta Imllisi 2. Live specimens of A. bullisi attached to the anterior ends of two Lamcllilyrachia

sp . (Jarden Banks Block 427, dive .'5.305, depth (ill nicters. 3. A live specimen of .A, Imllisi from Green C:anyon Block 185, dive

3.301, deplli 541 meters This specimen is attached by a byssus 32 mm i>el()w (lie anterior enil of the tube worm and lacks the notch

in the anierodorsal margin as seen in ligiires 4 and 9. ll is about 50 mm in height which is half tlie size of the specimen on left in

figure 2. 4. Detail of the live specimen on the right in ligure 2 The specimen is approximatcK , 1 10 mm in height and surrounds

the anterior portion of l.unwUihrarbia sp Note the position of the tuiie worm at center of photograph with the plume of the tube

worm protruding Irom the anterior end (arrow) and its relationship to the notch When the plume is extended, during life position,

it is totally contained within the mantle cavity of A. bullisi The i)yssus holds the A. Imllisi in a position to envelope the lube worm

Specimen from Garden Banks Block 427, dive 3305, depth (il 1 meters. Scale in upper Icll Is in centimeters

B. Kohl and H. E. Yokes, 1994

Page 11

5 cm

*4

%.

N

.vt.

Page 12

THE NAUTILUS, Vol. 108, No. 1

Figures 5-9. Specimen of /\. Imllisi eolleclecl alive Iroiii llie anterior of a lube worm. Green Canyon Block 185, dive 3301, 541

meters. Magnifieation (X 1). This specimen is comparable in size to the one figured 1)\ N'okes (1963, pi 2, figs, 1-4). Note the

B. Kohl and H. E. Yokes, 1994

Page 13

imals retrieved and photographed (figures 2-4) (hiring

recent dives of the JSL (Sept 1992 and JuK 1993) have

shown that the specimen figured h\ \ dkes was actualK

an ecophenotypic variant that grows attached to the tube

worm LamcUihrachia sp^ The inlienchng of the antero-

dorsal margin of A. btillisi and the de\elopment of an

open gape is the result of its attachment to the anterior

end of LamcUihrachia sp. Tliis has been reported from

specimens collected at Viosca Knoll Block 826 (Mac-

Donald & Ambler, 1992: 46)

Yokes (1963: 78) suggested that "the specimen, uhich

was alive when taken, had suftered serious damage to

the shell and mantle in the region of the lunule at a time

when the animal was approximately half grown." With-

out the know leilge that the specimen was originally at-

tached to a tube worm, this was a logical interpretation.

We now know that inbending of the anterodorsal margin

and open gape is a normal development for specimens

growing on the anterior end of LamcUihrachia (Mac-

donaldc? (;/,,1989). Carney (1992) reports that specimens

of A. hiilh.si with thin shells are either attached to car-

bonate rocks or swimming freely near the vent com-

munities v\hereas the forms attached to LamcUihrachia

have a distinctk thicker shell.

We suggest that smaller specimens of A. buUisi ma\

move from an attacfied position on the side of the tube

worm after achieving apprcximately one-half adult size,

and then to an anterior location during the later stages

of life where the gape is developed as it closes over the

tube worm. Members of the genus Acesta can resorb

their byssus filaments. Once unattached they can swim

to another position, near the anterior end of the tube

worm, and re-attach. Since LamcUihrachia has neither

mouth, gut nor anus and is completely dependent on

sulfur-reducing bacteria for its food, it is difficult to un-

derstand the ecological relationship between A. huUisi

and the tube worm. Could there be a symbiotic or com-

mensal association between the two organisms?

Acesta huUisi does not have symbiotic bacteria in its

mantle and appears to be a normal filter feeder. Addi-

tional research on the biology of A. hitUisi. being con-

ducted by C. R Fisher at Pennsylvania State Univ. (R.

S. Carney, personal communication), may answer some

of these cjuestions.

The description of the "pathologic" specimen of A.

huUisi by Yokes (1963), collected at Yiosca Knoll Block

826, was the first record of a mollusk from an oil seep-

chemosynthetic community in the Gulf of Mexico. Care-

ful study of the faunal collections dredged at YK 826 in

1962 by the R/Y Oregon would probably reveal the

existence of tube worms. Subseciuent dives by GERG in

1991 have documented the occurrence of LamcUihrachia

at YK 826 (MacDonald & Ambler, 1992). Based on our

present knowledge of the occurrence of A. hiiUisi in

chemosynthetic communities, there is no doubt that the

paratype of A. huUisi described l)y Yokes from YK 826

was a form attached to the anterior end of a specimen

of LamcUihrachia.

ACKNOWLEDGMENTS

The authors thank Harry H. Roberts, director of the

Coastal Studies Institute at Louisiana State University

and the chief scientist on the cruises of the R/Y Edwin

Link for Septemlter 1992 and the H/ Y Seward Johnson

for July 1993 dives. Also, thanks go to Robert Avent of

Minerals Management Service, Department of Interior

for providing background material on the chemosyn-

thetic faunas and to James M Rrf)oks and Ian R Mac-

Donald of GERG for providing information on Acesta

huUisi occurrences in the Gulf of Mexico.

We also thank Robert S. Carney for the loan of a

specimen of Acesta huUisi from the LSU collection for

photograph} . Emily H. Yokes, of Tulane LIniversity, pre-

pared the plates and reviewed and edited versions of the

manuscript. Marco Taviani, Institute di Geologia Marina,

Bologna, ItaK, who was on the 1992 cruise, assisted with

the shipboard photograph) The authors thank two anon-

\ mous reviewers for their suggestions to improve the

paper.

The dives were supported by the Louisiana Sea Grant

Program, New Orleans Office of Minerals Management

Service, National Oceanic and Atmospheric AgencN and

the National L^ndersea Research Center at the University

of North Carolina at Wilmington.

LITERATURE CITED

Carney, R .S 1992 Heterotrophic Fauna Associated with

CheiiiDsyiitiietic Communities. In: MacDonald, I. (ed.).

Technical Report of Northern Gulf of Mexico Chemosyn-

thetic Ecosystems Study: Literature Review and Data Syn-

thesis, vol. 2, V. S Department of Interior, Minerals Man-

agement Service, Gulf of Mexico OC'S Region, New Oriean.s,

LA, Chapter 9, p. 1-30

Fisher, C. R. 1990 Chemoautotrophic and Methanotropliic

Symbioses in Marine Invertebrates- Reviews in .Aijuatic

Sciences. 2 (3-4): 399-436.

l-'isher, C:, R. 1992. Physiology. Biocl)emistr>, and Ecology

of Vent and Seep Invertebrates with Chemoautotrophic

or Methanotrophic Symbionts. In: MacDonald, I. (ed).

Technical Report of Northern Gulf of Mexico Chemosyn-

thetic Ecosystems Study Literature Review and Data Syn-

thesis, vol. 2, U S Deparlmenl of Interior, Minerals Man-

agement Service, Gulf of Mexico OC^S Region, New Orleans,

LA, Chapter 7, p. 1-17.

Kennicutt. M C. II, J. M. Brooks, R R Bidigare, R. R. Fay,

T L. Wade and T. J. McDonald. 1985. Vent-type taxa

in a hydrocarbon seep region on the Louisiana slope. Na-

ture (London) 317(603.5): 351-353.

position ol die notch, (F'igs. 5,6,9) vviiitli allows A. inillisi to close over the tube worm. This is also evident in Yokes (1963, pi. 2,

figs. 2-3). Specimen on loan troni LSLI.

Page 14

THE NAUTILUS, Vol. 108, No. 1

MacDonald, I. R and J .Ambler. 1992. Regional distribution

of C^heniosynthetic Fauna and Community Ecology hi:

MacDonald, I. (ed.). Technical Report of Northern Gulf

of Mexico Chemosynthetic Ecosystems Study: Literature

Review and Data Synthesis, vol.2, U S Department of

Interior, Minerals Management Service, Gulf of Me.xico

(X;S Region, New Orleans, LA, Chapter 8, p. 1-53

MacDonald, L R., G. S. Borland, J. S. Baker, J. M. Brooks, M.

C. Kennicutt and R. R. Bidigare. 1989 Gulf of Mexico

chemosv nthetic communities II; Spatial distribution of seep

organisms and hydrocarbons at Rush Hill Marine Biology

KM: 235-247,

MacDonald, I R , N L. Guinasso, Jr , J. F. Reilly, J M. Brooks,

W R Callender, and S. G. Gabrielle. 1990. Gulf of

Mexico Hydrocarbon Seep Communities: VI. Patterns in

Community Structure and Habitat, Geo-Marine Letters,

10: 244-252

Paull, C K , B Hecker, R. Commeau, R.P Freeman-Lynda,

C Neumann, W.P. Corso, S. Golubic. J. E. Hook, E. Sikes,

and J Curray 1984. Biological communities at the Flor-

ida Escarpment resemble hydrothernial vent taxa. Science

226: 965-967.

U. S Department of Interior, Minerals Management Service.

1992. Northern Gulf of Mexico Chemosynthetic Ecosys-

tems Study: Literature Reyiew and Data Synthesis. Vol.

1, Executive Summary, 32 pp.; Vol. 2, Technical Report,

238 pp.; Vol. 3, Appendix, 243 pp.; Edited by Ian Mac-

Donald, Geochemical and Environmental Research Group,

Texas A &M University L' S Dept. of Interior, Minerals

Management Service, Gulf ot Mexico OCS Regional Of-

fice, New Orleans, LA,

Vokes, H, E 1963 Studies on Tertiary and Recent Giant

Limidae, Tulane Studies in Geologyl(2): 75-92, 2 pis.

THE NAUTILUS 108(l):15-22, 1994

Page 15

Shell Alignment for the Morphometric Analysis of High-spired

Gastropods.

Paul J. Morris'

Smithsonian Tropical Research

Institute

Apartatdo 2072

Balboa, Republica de Panama

Warren D. Allmon

Palenntological Research Institution

1259 Trumansburg Rd.

Ithaca, NY 14S50 USA

ABSTRACT

.\ significant problem often impedes the morphometric anaKsis

of high-spired gastropods. It is usualK' not possible to obtain

large samples of complete specimens. Without the ontogenetic

landmarks pro\ ided b\ earl\ u horls it is difficult to align in-

complete specuiiens to compare homologous w horls. We discuss

several methods that allow alignment of morphometric data

taken from incomplete specimens of high-spired gastropods.

We evaluated these methods b\ treating a set of complete

specimens as incomplete. Two methods produce reliable align-

ments. One of these uses an ontogenetic function of the whorl

expansion rate to estimate missing whorls, the other aligns whorls

of similar heights. Both produce reasonably precise and accu-

rate alignments, but under different assumptions. Once a set

of specimens has been aligned, it can be subjected to conven-

tional statistical anaKsis using accepted coinentions to deal

w ith missing data

Key words: Morphometries. Gastropoda. Missing Data. Tur-

ritellidae.

INTRODUCTION

Morphometric anal\ ses of very high-spired gastropods

(such as members of the families Cerithiidae, Lo.xone-

matidae, Murchinsoniidae, Nerineidae, Turritellidae, and

Terebridae) are difficult because complete specimens are

usualK unavailable or rare. This is especiaJK true for

fossil taxa. When the juvenile whorls ha\ e been broken

off of most specimens it is difficult or impossible to specif \

homologous points (i.e.. similar whorl number) among

specimens. Therefore, construction of data matrices for

morphometric or statistical anaKses of a large number

of specimens of such incomplete gastropods is usualK

problematic. This difficultv may account for the paucitv

of morphometric studies of high spired gastropods in the

literature (e.g.. .^llmon, 1994).

' Current Address: Paleontological Research Institution.

Trumansburg Rd . Ithaca. \y"i4S50 USA

259

A variety of methods can be used to align the whorls

of incomplete specimens w ith each other \\'e have im-

plemented three algorithms for use on sets of traditional

point-to-point measurements. We assessed the accuracy

of these algorithms by their ability to predict true whorl

number in a data set of complete specimens that we

treated as incomplete. In this data set, we truncated

complete specimens at known whorl numbers. Two al-

gorithms produced good alignments of shells of turritel-

lids in this test data set. Both of these align most speci-

mens to within one whorl of their true position. An

algorithm that uses iterative comparisons of whorl heights

produces slightly more precise alignments. .\n algorithm

that uses an ontogenetic function of the whorl expansion

rate to infer numbers of missing whorls is slightly more

accurate. There are several tradeoffs between these two

algorithms.

SHELL ALIGNMENT

In most gastropods it is possible to identify a homologous

point for aligning whorls. Either the end of the proto-

conch or a varix produced in the adult shell can be used

as landmarks from which whorls can be counted. It is

therefore easy to align whorls that are of the same dis-

tance from this landmark in different specimens. Such

aligned specimens are easiK compared using traditional

point-to-point measures, Raupian parameters, or moving

frame parameters. In most high-spired shells, however,

whorls differ only in size and shape, and lack discrete

landmarks. When specimens have been broken and are

incomplete, there are no clear means for inferring how-

many whorls are missing. Incomplete specimens, which

are particularly common as fossils, are therefore difficult

to align, homologize, and anaK ze morphometricalK . This

problem is depicted in Figure 1 .

There are three basic ways of aligning incomplete high

spired shells. These are: alignment by homologv of or-

nament, inference of missing whorls from whorl expan-

sion rate (W of Raup, 1966), or alignment by compar-

isons of whorl heights among specimens. (One can also

Page 16

THE NAUTILUS, Vol, 108, No. 1

l-i-J..V_ ' \1

Figure 1. The problem of v\horl alignnieiU in two incomplete

specimens of high spired gastropods. In tlie alisence of the

protoconch, juvenile w horls, or other homologous points, it is

necessar) to find some means of aligning w horls in order to

conduct mnrphometric anaKses The two principal \va>s of

making alignments are by matching whorls of similar size, or

by inferring the number of whorls missing from the small end

of each specimen.

use other measures that increase through ontogeny, such

as \\ horl width, but for the remainder of this paper we

refer exclusiveK to whorl height). We ha\e considered

five variations of these methods, alignment by: 1) com-

parison of homologous features of ornament, 2) inter-

polation of missing whorls using the wliorl expansion rate

from earliest preserved whorls of each specimen, 3) in-

terpolation of missing whorls using some function of W

calculated from a composite average specimen, 4) com-

parison of whorl heights between each pair of specimens,

and 5) iterative correction and comparison of whorl

heights with a composite a\erage specimen .\ method

that uses iterati\'e comparisons of whorl heights appears

to be the most generalK useful. None of these, however,

are entireK perfect \\a\s to infer how man> whorls are

missing in an incomplete specimen. There is no ideal

way to align indi\ idual specimens that lack earh whorls

and homologous end points. Each of these approaches

has limitations and makes compromises.

.■\ first approach is alignment using homologous fea-

tures of spiral ornamentation, such as the points of bi-

furcation or origination of keels. This approach (because

it involves homologous points) w ould seem most reliable.

It is limited, howe\er, by an inability to extend homol-

ogies of ornamentation be\ond small groups of related

species, and b\ the need for man\ homologous points

spread throughout ontogeny. As a result, it is useful only

in a few specific cases. It does not lend itself to imple-

mentation as a general algorithm that could appK , for

example, to both turritellids and loxonematids. It also

must avoid the risk of making arbitrary assignment of

homologous points in contiiuiousK growing ornament.

.\ second approach is alignment using w horl expansion

rates to inter the ninnber ol missing w horls between the

protoconch and the first [^reserved whorl in each speci-

men. This could be done 1)> first calculating the whorl

expansion rate from the lirsl preserved whf)rls iil each

specimen. This parameter can then be used to calculate

how main w horls are missing between the first preser\ed

w horl and some smaller first whorl (e.g.. the protoconch,

or the smallest preser\ed whorl in the data set). That is,

iterativeK reducing the height of the smallest preserved

whorl of a specimen b\ its whorl expansion rate. The

number of steps required to reduce this height value to

below the protoconch size provides an estimate for the

number of missing whorls. The advantage of this method

is that the estimated number of missing whorls in each

specimen is independent of the other specimens in the

sample. However, both measurement errors in the small-

est whorls and ontogenetic changes in the w horl expan-

sion rate affect these alignments.

These errors can be reduced by using some ontogenetic

function of the whorl expansion rate rather than an un-

changing W \ alue. This must perforce be calculated from

other specimens in the sample. It is not possible to infer

from a single specimen how W might ha\e \ aried in its

missing whorls. Ideally, this ontogenetic huiction oi W

w ould be continuousK \ ariable, reflecting the real nature

of ontogenetic changes in w horl expansion rate. In man\'

of our samples, however, few specimens preserve the

smallest whorls in the data set. It is therefore necessary

to increase the sample size, and decrease the resolution

of ontogenetic changes in W. We feel that three whorl

expansion rates, produced by a di\ision of the data set

(the composite specimen) into three parts (smallest 'i,

second 'i, largest 'A), pro\ide the best compromise be-

tween resolution of W and sample size. Thus, a whorl

expansion rate is calculated for each pair of whorls in

each specimen, and an average W is calculated from the

whorls in each part of the specimen. These three W

\alues are then used to infer how many w horls are miss-

ing from each specimen.

For example, consider a specimen in w hich the small-

est preserved whorl falls within in the second U of the

range of whorl sizes in a sample. The number of missing

w horls in this specimen are calculated b\ iterati\el\' re-

ducing the height of its first preserved w horl. While this

height value remains within the second 'j, the average

W V alue from the second ' j is used to calculate the height

of the next smaller w horl. When this value falls into the

range of heights of the smallest '4, the average W value

from the smallest U is likewise used to fill in smaller

wliorls. In each specimen, the number of steps required

to reduce the height of the smallest preserved whorl to

less than the height of the smallest w horl in the data set

(or a protoconch height) are an estimate ot the nutnber

of missing w horls.

.\ third possible approach is the aligmnent of w horls

of similar height. This is the most intuitive approach,

and the one we find ourselves naturalK taking when

pliv sicallv lining up specimens next to each other. There

are two obv ions vv av s to carry out these aligmnents; each

starts bv sorting the specimens in order of increasing size

of first whorl. Specimens mav' then be aligned bv pairwise

comparisons or bv constructing a composite average

specimen and itcratlvcly locating and moving poorly

aligned specimens, .\liginnent by height, however, offers

p. J. Morris and W. D. Allmon, 1994

Page 17

several computational problems, Pairwise c<imparisoiis

fail unless each pair of specimens overlaps by several

w liorls. It is therefore necessar\ to compare all the spec-

imens in the sample to some standard specimen that

covers the full range of whorls. If a single specimen is

more complete than an\ other in the sample (ha\ ing the

smallest small w horl and the largest large whorl) all other

specimens can be aligned v\ith it. Such specimens are

rare. A proxy for such a specimen is easily constructed

b\ a\'eragiiig across all the specimens in the data set to

generate a composite a\ erage specimen. This can be used

as the reference to which other specimens are aligned.

It is a variation of this approach that we have tmnul to

be the most generalK reliable.

The specific method that we found to be the most

precise is the alignment of specimens in the data set

through: 1) sorting the specimens in order of increasing

size of first preser\ ed w horl. 2) stepping through the data

set making pairwise comparisons of first w horl size and

moving each larger specimen until its first whorl was

aligned with some whorl on each smaller specimen

(alignment by smallest whcui size), 3) calculating the

average size of each whorl in the data set and the de-

viation of each specimen from the a\erage composite

specimen, 4) moving poorly aligned specimens, and 5)

iterating steps 3 and 4 to minimize the total error (total

deviation of all specimens from the composite average

specimen). The accuracy of these alignments can be im-

proved by removing small w horls represented by only a

single specimen.

We have assessed the accuracy and precision of align-

ment in three of the five methods discussed abov e. These

are: inference of missing w horls using W from the earliest

preser\ed whorls (Ju\enile W), inference of missing

whorls using a crude function of ontogenetic changes in

the w liorl expansion rate (Three W N'alues). and iterative

alignment of whorl heights (Height).

SOURCES OF ERROR

There are five principal sources of errors in alignment.

These are: 1) ontogenetic changes in coiling parameters

or shape, 2) intra-population variabilitv, 3) measurement

error, 4) rotation error, and 5) tlie relationships among

size, age, and homologx . The eftects of some of these

sources of error can be avoided or reduced, whereas

others are less tractable, .\lignment errors introduced bx'

ontogenetic changes in the w horl expansion rate, another

coiling parameter, or whorl shape can be reduced b\

considering some ontogenetic function of W, or b\ mak-

ing alignments to a composite average specimen. Effects

of measurement error can likew ise be reduced 1)\ making

alignments using information from multiple w horls and

specimens. W'ithin-population \ariabilit> can affect

alignments in several ways. Such \ arialiilit\ can produce

a broad spread of alignments, or it can produce non-

intuitive systematic missalignments. It is essential that

4 11 4 11 4

True Whorl humber

Figures 2 — I. .^ccuracN and precision of tiiree methods ot \\ horl

alignment discussed in this paper. Whorl heights uere mea-

sured in ten specimens of Tiirritella gonostoma and used to

construct a data set of 50 truncated specimens containing true

whorl numbers and whorl height. Open circles are aligned

whorl number, filled circles are plus and minus one standard

deviation. Lines represent correct whorl alignment and error

of plus and minus one whorl. 2. Alignment from Ju\enile VV

3. .Alignment using three W \alues, 4. .Ahgnment b\ Height.

.Alignment from Juvenile \V is neither accurate nor precise.

The other twci metlicKls are suhstantialK laetter at correctK

aligning specimens in this test.

the user examine aligned samples to understand how this

variabilitx may have affected the alignments. Rotation

error is the result of taking a set of measurements from

one plane in a continuousK growing spiral shell. In our

test data set it is the principle source of reduced precision

for the tw o best alignment methods. Effects of this source

of error can be reduced In either using three dimensional

coordinates and mo\ ing frame parameters y\ckerly, 1989;

1990; Johnston et ai. 1991), or b\ aligning specimens

before measurement (rotating each specimen until a par-

ticular whorl width (or height) lies in the plane of mea-

surement).

The relationships among size, age, and homologx' pro-

vide a subtle set of philosophical problems. Age, growth

rate, size, and shape are complexly interrelated in gas-

tropods. Environmental factors, such as the availability

of food, readily influence the growth rates of gastropods.

In time averaged samples it is entireK possible that w horls

of the same size (and the same true whorl number) in

different individuals represent different points in their

ontogenies. In most paleontological studies, how ever, the

primary goal is simply to assess how shape and size vary

within and between samples Patterns in the data can

then constrain various taxonomic, evolutionary or pale-

oecologic hypotheses. The two most common questions

for w hich detailed morphometric studies are undertaken

are species discrimination, and trends in variability

tlirough time [i.e.. tempo and mode of evolution). Lines

of ev idence beyond morphologv', such as taphonomv', are

needed to assess the role ol ecophenotypic factors in

increasing variance or creating clusters in the data.

Page 18

THE NAUTILUS, Vol. 108, No. 1

Table 1. Comparison of advantages and pitfalls of three methods of whorl alignment. These methods are; Alignment b> Height

using iterative comparison with a composite average specimen, Alignment using three W values computed from composite average

specimen, and .Alignment using VV calculated from smallest three whorls of each specimen. Alignment by Height is the most suitable

except for cases where 1) many specimens are being aligned and computing time is a concern, or 2) many specimens consist of

just two or three whorls. If the data are three dimensional coordinates suitable for moving frame analysis we expect that a modification

of the Height algorithm will probabK produce the best alignments .'Alignments of a test data set (complete specimens treated as

incomplete) suggest .\lignment by Height as the best overall technique.

.Alignment b\

Height

HISPIRE .3

Three W values

HISPIRE 2

Juvenile W

HISPIRE 1

Specimens aligned using:

.Advantages;

Speed

Alignment

Disadvantages:

Speed

.Alignment

Systematic Missalligment;

Dependence on sample composition;

Recommended for;

Most applications

Many specimens

Few whorls

Principle sources of alignment error;

Measurement error

Ontogenetic del W

Population del W

Few whorls

+ +

+ + +

Self

Fast

Inaccurate

Much

No

+ + +

Test data set, Tiirritella gonostoma

Average of SDs;

Relative precision of alignment

Small whorls

Middle whorls

Large whorls

0.93

high

moderate

1.12

high

moderate

1.73

low

ASSUMPTIONS

A fundamental assumption follows from the previous

paragraph: The questions under consideration can be

tested with information about size and shape without

reference to age (see discussion in Janie.s, 1988), Beyond

this, each alignment method makes its own set of as-

sumptions about the specimens that are to be aligned.

Both the accuracy of alignment and the assumptions

behind the methods affect their suitability for particular

analytic problems. There are three principal assumptions

made by the method of alignment from juvenile W val-

ues. It assumes that there is little measurement error in

the early whorls. In addition, it assumes that the w horl

expansion rate is constant throughout ontogeny. Whorl

number is expected to correlate with the height of the

first preserved whorl. The method of alignment using

three W values allows for ontogenetic variation in the

whorl expansion rate. However, it is still sensitive to

errors in measurements of the earliest preserved whorl.

In addition, this method takes on the assumption that

there is little population variability in the w horl expan-

sion rate. .Alignment by height assumes that whorls of

similar heights are homologous. It does not directly as-

sume that there is little population variabilit> in the

whorl expansion rate, but if this exists, it may produce

non-intuitive alignments.

TESTS OF ALIGNMENTS

We ha\e tested the accuracy of these three methods of

w horl alignment b> evaluating their ability to correctly

align whorls in a sample of complete specimens that we

treated as incomplete. We used ten complete specimens

of Tiirritella gonostoma Valenciennes, 1832, a Recent

turritellid that undergoes relatively large ontogenetic

changes in w horl expansion rate. We photographed these

specimens, enlarged the images, and digitized points at

the suture of each whorl using a Numonics digitizing

tablet. The heights of successive v\ horls were calculated

from these digitized points. We then truncated the data

for each specimen at five different whorls spanning the

region of greatest change in whorl profile. This procedure

provided us w ith a data set of .50 incomplete specimens

p. J. Morris and W. D. Allmon, 1994

Page 19

s

E

.•QO

o

True Whorl Number Alligned Whorl No.

Figures 5, 6. Assessment of accuracy ul w liw I uligimient b> iterative comparisons of whorl height with composite average specimen.

5. Comparison of true and ahgned whorl number. Open circles are aligned whorl number, filled circles are plus and minus one

standard deviation. Lines represent error of plus and minus one whorl from correct whorl alignment, 6. Whorl heights and aligned

whorl number Lines connect the whorls of individual specimens Note that the accuracy of this method is strongly affected by the

alignment of the two smallest specimens in the data set. Elimination of the two smallest aligned whorls will result in substantial

uTipro\ement in accurac> .

of various numbers of whorls, for wliicli whorl height

and true whorl number w ere known in each whorl. This

data set was entered into three conversion programs

(HISPIRE, HISPIRE2, HISPIRE3, see Appendix 2) that

implement the three whorl alignment algorithms.

Figures 2-4 illustrate the different alignments of this

data set b\ the three programs. These figures are graphs

of true whorl number against aligned whorl number.

The mean alignments for these 50 specimens (plus and

minus one standard de\iation) are compared with a cor-

rect alignment (plus and minus one whorl). Clearly align-

ment by ju\enile VV performs least well in this test. On

careful examination of the results, both the ontogenetic

changes in whorl expansion rate and measurement error

contribute to its failure.

The other two alignment methods produce substan-

tially better results. Here, alignment using three W values

produces more accurate results than alignment by height,

.alignment using three W values places the mean align-

ment for each whorl slightly closer to the true whorl

value than alignment by height. However, when the

standard deviations in alignment in each v\ horl (exclud-

ing the first and last whorls in each case) are averaged,

alignment b\ height is more precise (Table 1). It appears

that alignment by height is slightK' more precise (tighter

clustering) and that alignment using three W values is

slightK- more accurate (closer to true).

In order to understand the quality of alignments, it is

important that we consider the reasons for these differ-

ences. The sources of these errors are revealed in an

examination of plots of true against aligned whorl num-

ber, whorl height against true whorl number (e.g.. Fig-

ures 4,5), and the original data. Alignment using three

W values is subject to misalignments produced by mea-

surement errors in the early whorls. Such measurement

errors, along with slight rotation errors and variation in

^^', tend to produce misalignments in steps of one whorl.

L'sing this method, some specimens are well aligned,

others suffer quantum misalignment. In contrast, align-

ment by height produces misalignments that appear to

be the result of rotation error. All specimens are tightly

aligned with each other (Figure 6) and their spread is

some combination of true variation and alignment error.

Careful examination of the graph reveals that the lower

accuracy of this method is also the result of rotation error.

The two specimens that have the smallest whorls in the

data set happen to be aligned to the left relative to the

others. The alignment of these two specimens is the cause

of the reduced accuracy of alignment of the rest of the

specimens. A remecK suggests itself at once. Removal of

early whorls represented by only a few specimens should

make alignment by height the most accurate and precise

of the three methods.

To test this contention further, we qualitatively eval-

uated these programs. We examined the alignments they

produce in fifteen samples of four species of Paleocene

and Eocene turritellids from the U.S. Gulf coastal plain

(Turritella mortoni Conrad, 1830, T. carinata I.Lea,

1S33, T. perdita Conrad, 1S65, and T. praecincta Con-

rad, 1864). .almost all the specimens in these samples are

incomplete. It is therefore not possible to compare the

alignments to true values. Here we assessed the precision

of alignments b\' graphing aligned w horl number against

w horl height. In all the samples we examined, alignment

b\ height consistentK' produced tight clusters. The results

shown in Figures 7,8 are typical. In contrast, alignment

using three W values often placed one or more specimens

outside of a cluster of aligned specimens. This again

reveals a weakness in this method of alignment. Small

errors (in the measurement of the first preserved whorl,

or in fit of the specimen to the three W values) can easily

be magnified to misalignments of one or more whorls.

Page 20

THE NAUTILUS, Vol. 108, No. 1

Alhgned WhorJ No.

o

All igned Whorl No.

Figures 7. 8. Whorl alignments in 7. eight specimens of Tiir-

rilella rnortoni from the Paleocene Aquia Fm. at Piscatavva>

C:reek Prince Georges Co., MD (Appendix 1) and 8. twelve

specimens of Tiirritella carinata from the Eocene Gosport Sand

at Clayfjorne Bluff, Monroe Clo.. Alabama, .\ligned u liorl num-

ber is plotted against whorl height. Lines connect the whorls

of individual specimens. These alignments are t> pical for the

samples of fossil turritellids that we examined.

CHECKING ALIGNMENTS

Given the assumptions of these algorithms and the va-

garies of real data, the alignment programs cannot be

treated as black boxes that produce infallible results. It

is essential that the user e\aluate the accurac)' of the

aligruTients. The programs themseUes pro\ide some

warnings, but graphs of aligned \\li<irl number against

\\ horl fieight (as in Figures 7, 8) should be examined for

all samples. Tfie programs that implement alignments

using whorl expansion rates provide the user with sum-

mary statistics. These include the whorl ex|)ansion rate

at the small end (average of W in lirst and second pre-

served wliorls of each specimen) and large end (\V in

largest preserved whorl of each s[)ecimen). The mean

whorl expansion rate, standard deviation of W, and the

coefficient of variation of W [CV=(SD/Mean)X lOO'V ],

each calculated from the whole data set, are provided

for both tile large end and the small riul Tin' [)rograins

caution the user it the coefficient of \ariation of either

of these values exceeds 5'"i, and provide a warning if

these C\'s exceed 10^7 . Such large C\'s suggest violation

of the assumptions of the method. In addition, the pro-

gram implementing aligimient using three W \ alues as-

sesses the variabilitv' in whorl expansion rate in each of

the three parts of the specimen. Coefficients of \ ariation

greater than IC^'c again suggest \iolation of the assump-

tion of low between specimen differences in whorl ex-

pansion rates. The program that implements alignment

b\ height checks how much variation is present in the

heights of each whorl in the composite average specimen.

It the average CV for all whorls except the first and the

last is less than 7% then alignments are probably reason-

able. If this average C\' exceeds 8*7. then there are prob-

abl\' serious errors in alignment, or considerable variation

in the sample. If this average C\' exceeds ISl, it is es-

sential that the user evaluate the results by plotting whorl

lunnber against whorl height (as in Figure 6).

CHOICE OF METHOD

The choice of alignment method in a particular case

involves four factors. These are accuracy, precision, as-

sumptions, and the goal of the anaKsis. In general, we

feel that the goals of a morphometric anaKsis will best

be met b\ the methods of either three W \alues or it-

erati\e comparison of heights (Table 1). Other methods

are preferred in onl\ a few other cases. If many ho-

mologous points are available in the ornament of all taxa

imder consideration then these may prove the best means

ot alignment. We recommend avoiding use of the whorl

expansion rate in the first preserved whorls of each spec-

imen to align w horls unless it can be clearK demonstrated

thai there is no substantial ontogenetic change in the

w horl expansion rate, and that measurement error is not

large in small w horls. This method of alignment should

not be used if the coefficient of variation of w horl height

from a small w horl of a single specimen measured re-

peatedK ten times is greater than about three percent.

Our examination of the test data set and of alignments

in samples of fossil species suggests that the methods of

aliginnent by three W values and alignment b\ whorl

heights both align specimens to within about one whorl

of their correct position. .Alignment b\ height is more

precise (has less scatter), while aliginnent b\ three W

\ alues is more accurate (is closer to true whorl number).

If comparisons are made between homologous whorls

then aliginnent bv height will introduce less error. On

the other haml, in comparisons of groups of whole spec-

imens, alignment b\ three W values will be superior.

Population \ariabilit\ in whorl expansion rate w ill affect

both of these methods, but in difterent wa\s. If a sample

contains specimens from two populations that differ in

whorl expansion rate then aliginnent In height will tend

tu make good alignments within each cluster. It will.

lidwcNer, systematicalK inissalign one cluster with re-

spect to the other. On the other hand, aliginnent In three

W \ alues will iiicrea.se the spread ol the entire sample.

p. J. Morris and W. D. Allmon, 1994

Page 21

It will reduce the distinctness of the clusters. Alignment

b\ height is better at preserving differences between

indi\ idual specimens. In addition, the accuracy of align-

ment b\ height can be increased b\ removing the small-

est t\\ o \\ horls from the data set. The smaller sample size

of the small (and large) whorls reduces the accuracy' of

alignment for some specimens. Speed max be an addi-

tional concern. ,\t large sample sizes, our algorithm for

alignment b\ height, being iterative, is significantK slower

than our algorithm for alignment bs three W values (17

min. \ s. 1 min. for 50 specimens on an 80386 at 20 niHz).

This brings us to the original purpose of this discussion;

the morphometric anal\ sis of high-spired gastropods that

are missing w horls. The goal of the alignment programs

is to place markers in place of measurements in the

missing whorls. If the data set is subsequent!)' subjected

to multivariate analysis these markers (zeroes in our ex-

ample, .\ppendi.x 1) can be treated as "missing \alues."

Most t\ pes of ordination and cluster anaKsis. howe\er,

deal poorK w ith missing data. It is therefore necessary

either to throw out many whorls and reduce the analysis

to a set of whorls that are complete in all specimens in

the data set, or to appK a technique for estimating miss-

ing values. One efficient wax to do this is to create a

correlation matri.x from the data. Multi\ ariate techniques

can then be applied to this correlation matrix rather than

the incomplete data set (Beale & Little, 1975; Little &

Rubin, 1987). For e.xample; an anaKsis with BMDP (Dix-

on, 1988) could use program SD (Correlation w ith miss-

ing data, Engelman, 1988) to create a correlation matri.x

from the data set. This correlation matrix could then be

used as input for program 4M (Factor anaKsis, which

would normalK exclude all cases w ith any missing data

from the original data set, Frane ct ai. 1988). Program

4M could then be used for \arious sorts of exploratory

variable reduction and factor anaKsis (e.g.. Kleinbaum

et ai, 1988). which could not be performed on the raw

data. Holding in mind the caveats of error introduced

by rotation error and assumed meaningful similarity be-

tween whorls of similar height, the method of alignment

b\ iterative comparisons of whorl heights with a com-

posite average specimen should pro\e suital)le for most

cases in which incomplete high-spired gastroi^otls are to

be aligned for morphometric anaK sis.

ACKNOWLEDGEMENTS

R. E. Chapman provided many insightful commetits on

an earlier \ersion of this paper. He is especially acknowl-

edged for pointing out to us that extant turritellids are

often complete, and that it is possible to assess the ac-

curacy of alignment by treating whole specimens as in-

complete. These tests ha\e show n us that several of our

original assumptions were invalid. S.A. Shellenberg

/3-tested several versions of HISPIRE and raised many

useful questions and suggestions. This manuscript has

also benefited from comments by B.A. Robinson and an

anonvmous reviewer.

LITERATURE CITED

.•\ckerly, S. C. 1990. Using growth functions to identify horn-

ologus landmarks on mollusc shells. In: Rohlf, F.J. and F.

L. Bookstein (eds. ). Proceedings of the Michigan Morpho-

metries Workshop. University of Michigan Museum of

Zoology, .\nn Arbor, MI, pp. 339-344.

.Ackerly. S. C. 1990. The kinematics of accretionarv shell

growth, with examples from molluscs and brachiopods.

Paleobiology 15147-164

.\llmon, W. D. 1994, Systematics and evolution of Cenozoic

American Turritellidae. I. Paleocene and Eocene species

related to "Turritella mortoni Conrad" and "Turritella

humerosa Conrad" from the US Gulf and Atlantic coastal

plains. Palaeontographica Americana 58 (in press).

Beale. E. M. L. and R. J. .\. Little. 1975. Missing values in

multivariate analvsis. Journal of the Roval Statistical So-

ciety .378:129-145.

Dixon. \\ J. (ed.) 1988. BMDP statistical software manual,

2 X'ols. University of California Press, Berkelev, 1234 p.

Engelman, L. 1988. 8D correlations with missing data. In:

Dixon, W. J., (ed. ) BMDP Statistical Sof tw are Manual. Vols

2. University of California Press, Berkeley . pp. 679-688.

Frane, J , R. Jennrich, and P. Sampson. 1988. 4M factor

analysis. In: Dixon. W. J . (ed). BMDP Statistical Software

Manual, \'ol 1, Lniversitv of California Press, Berkelev,

pp, 309-335.

Johnston, M. R., R. E. Tabachnick. and F. L. Bookstein, 1991,

Landmark-based morphometries of spiral accretionarv

growth. Paleobiology 17:19-36,

Jones, D. S. 1988. Sclerochronology and the size versus age

problem. In: McKinney, M. L. (ed.). Heterochrony in Evo-

lution. Plenum Press, NY, pp. 93-110.

Kleinbaum, D. C, L. L. Kupper, and K. E. Muller. 1988.

.\pplied regression analy sis and other multivariable meth-

ods. PWS-KENT Publishing Co., Boston. 718p.

Little. R J A. and D. B. Rubin. 1987. Statistical analysis with

missing data. Wiley , New York, NY, 361p,

Raup, D, M, 1966, Geometric analysis of shell coiling: general

problems. Journal of Paleontology 40:1178-1190,

APPENDIX 1

Sample conversion using the method of alignment by

iterative comparisons of height (program HISPIREo, see

■Appendix 2), Four specimens of Turritella mortoni Con-

rad, ISoO from The Paleocene .Acjuia Fm. at Piscataway

Creed, Prince Georges Co. MD. N'ariables (see .Allmon,

1994) are: SP#, specimen number; WH#, whorl num-

ber; Height, whorl height; CHI. carina height; CH2,

carina height abo\ e suture; SW, w idth at adapical suture;

M\\', width at mid-wliorl; C\\ , width at carina; CAN,

carina angle.

SP# WH# Height CHI CH2 SW

1 1 8.19 6.62 156 14 3

1 2 10.2 8.71 1.56 16.5

1 3 143 10.4 390 19 3

1 4 15,8 115 4,28 21,1

2 1 3,12 2 34 0,779 3,75 4,64 5,39 164

2 2 3,51 2,86 0,6.50 4.85 6.12 6.94 174

2 3 4,29 3,51 0 779 6,08 7,37 8,54 177

2 4 5,45 4 29 1,16 7,94 8,96 11,3 162

M\\ C\\ CAN

16 0 18 9 1.56

18.5 22.4 159

22 0 26.1 160

24.5 28.6 162

Page 22

THE NAUTILUS, Vol. 108, No. 1

Sample Output data file after conversion by HISPIRE3

(alignment b\ height):(For claritN, blank lines have here

been added between specimens. Options selected: single

line per specimen, space delimited, zero for missing value

indicator,)

0000 0 0 00000000000000000000000

0000000000000 S. 19 6.62 1.56 14.3 16.0 18.9

156 10.2 8.71 1.56 16.5 18.5 22.4 159 14.3 10.4 3.90 19.3

22.0 26,1 160 15.8 11,5 4,28 21,1 24,5 28,6 162

0 0 0 0 0 0 0 0 0 0 0 0 0 0 3, 12 2,34 0.779 3.75 4.64 5.39

164 3.51 2.86 0.650 4.85 6.12 6.94 174 4.29 3.51 0.779

6,08 7.37 8.54 177 5.45 4.29 1,16 7,94 8,96 1 1.3 162 6.76

5.60 1.16 10.3 12.1 13.6 175 9.23 7.80 1.43 12.2 14.7 17.5

180 10.4 8.19 2.21 15.0 17.8 21.8 160 0 0 0 0 0 0 0

2.34 1.69 0.648 2,44 3,57 3.98 140 2.99 2.21 0.779 3.28

4.42 5.09 168 3.90 2.86 1.03 4.66 6.11 6.52 157 4.04 3.41

0.629 5.84 7.19 8.18 155 6.10 4.81 1,29 7.35 9.20 10.0

172 7.15 5.85 1.29 9.20 12.7 14.1 164 0 0 0 0 0 0 0 0 0

0000000000000000000

APPENDIX 2

Notes on the computer programs; Three programs (HIS-

PIRE, HISPIRE2, HISPIRE3) carry out the alignments

(b\ Juvenile \V, Three Ws, and Height, respectively)

discussed in this paper. These programs are written in

Microsoft Quickbasic 4,0, which is a structured BASIC,

They contain some extensions of the .ANSI BASIC stan-

dard, these include multiple line (block) IF-THEN-ELSE

statements, DO-LOOP statements, the REDIM state-

ment, the LTRIM$ function and the UCASE$ function,

The\ should be compatible with most microcomputer

BASIC compilers (but not interpreters such as IBM Basic

or GWBasic), They can be compiled by Microsoft Quick-

basic compilers from version 2.0 on. The source code for

each program has been thoroughly documented and they

have, we hope, been written so that little work should

be required to convert them to run under a minimal

BASIC standard. The three programs have successively

less rigid requirements for the data format of the files

they convert, and are successively more flexible in their

output formats. ■•Ml expect at least an input data file with:

one whorl per line, a whorl number on each line, con-

secutive whorl numbers starting with one, a specimen

number on each line, and a whorl height on each line.

Output options in HISPIRE3 include comma or space

delimited free format, user specified missing value in-

dicator, truncation of variables to three significant fig-

ures, specification of the order of variables on each line,

and single whorl or single specimen on each line. None

of these programs produce fortran format data files. \\\

three programs are available as the Quickbasic source

code and a compiled e.xecutable version of each program,

together with a set of sample data files and written doc-

umentation, on disk for IBM compatible per.sonal com-

puters from the first author.

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4.81 4.03 0.777

7.41 9.21 10.6 177 6.63 5.07 1.56 9.82 11.3 12.5 179 7.93

6.76 1.16 11.4 13.4 16,6 172 10,2 8.19 2.08 15.0 17.5 20.3

172 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

THE NAUTILUS 108(l):23-24, 1994

Page 23

On Planorbiila nebraskensis Leonard, 1948, and its position in the

evolutionary sequence of North American planorbuhds

Harold G. Pierce

Research Associate

University of Nebraska State Museum

Lincoln. NE 68588-0514, VS\

Leonard (1948:62) described Planorlmla nebraskensis

from what he considered to be Yannouthian sediments

in Knox County, extreme northern Nebraska, and later

reported this taxon from a second locality in Lyons Coun-

ty, Iowa (Leonard, 1950:18). Leonard commented that

"P. nebraskensis is apparently related to P. wheatleyi"

(Lea, 1858) and that the lamellae were "disposed as in

Planorbiila wheatleyi", without further comment on the

orientation of specific lamellae of the outer lip. Exter-

nally, P. nebraskensis is very similar to the modern la-

mellate species (subspecies?) of Planorlmla In fact, with-

out reference to the apertural lamellae, Planorbiila

armigera (Say, 1821) (Figure 1), P. wheatleyi (Figure

2). and P. nebraskensis can not be separated easily. Sub-

sequent work at Leonard s type localit), near Locality

KX-145 of the University of Nebraska State Museum,

has shown the age of the fauna to be late Kansan. The

fauna is recovered from sediments below a bed of the

Pearlette Type O ash, dated at 600 KA, and is associated

with a sparse, cool to boreal, mammalian fauna. Topo-

types of P. nebraskensis were collected for comparison.

Examination of these topotypes disclosed a substantial

difference in the placement of the upper palatal lamella

(Figure 3). With the shell axis vertical and viewed from

the axis outward, the upper palatal lamella is oriented

vertically, as is the upper palatal lamella of P. wheatleyi

(Figure 2), not nearly horizontalK' as witli P. armigera

(Figure 1). However, the lower palatal lamella of P.

nebraskensis does not have the elongated and very dis-

tinctive vertical segment characteristic of P. wheatleyi.

Subsequent work on the Albert Ahrens Locality (NO-

104) in Nuckols County, extreme southern Nebraska,

resulted in collection of another \ery boreal mammalian

fauna of late Kansan (Irvingtonian) age. This locality is

not directK associated with an ash, but nearby ash de-

posits strongK suggest that it is stratigraphicalK just above

the Pearlette T> pe O ash. and is, therefore, only slightly

younger than the KX-145 locality. Mammals were di-

verse, and of a strongly boreal character (Voorhies &

Corner, 1991 ). Analysis of arboreal pollen confirmed that

a boreal, almost taiga-like, environment existed here

(Bolick, 1991), The molluscan fauna from this locality is

quite similar to that of Leonards t\ pe locality. Exami-

nation of the planorbuliils from NO- 104 locality showed

that they had the unique placement of the labial lamella

characteristic of P. nebraskensis (Figure 4). This sug-

gested that the P. nebraskensis v\as not an unique ab-

erration of the KX-145 locality, and nearby Lyons Coun-

ty, Iowa only, but a widespread taxa. This also prompted

reevaluation of several older (late Pliocene, Blancan, ca.

2 MA) faunas from Nebraska that included planorbulids,

especially those reported by Pierce (1990), Careful re-

evaluation, including break-back of specimens, showed

that the planorbids of late Pliocene Localities KX-109,

KX-143 and CD- 1 04, had been incorrectly identified as

P. armigera. These specimens were clearK' P. nebras-

kensis (Figure 5), and demonstrated a significant range,

both spatially and temporally, for this species.

Fossil planorbulids from strata older than Quaternary

are not common, possiblv due to the unfortunate past

habit of lumping man> disparate planispiral taxa under

a form genus "Planorbis" Ta\lor (1960:36) lists P. ar-

migera as a component ot the Sand Draw fauna (Blan-

can), although I have yet to collect a planorbulid from

that locality for comparison. Taylor (1966) refers, with

question, several Idaho and Wyoming planorbulids, also

of Blancan age, to Planorbiila eampestris (Dawson, 1875).

Hannibal (1912:157-158) described P. mojavensis from

the Miocene Barstow Formation of California, The la-

mellae of P. mojavensis are unique, with only three

lamellae on the outer lip (Taylor, 1954:74). Pierce

(1993) described Planorbula powelli from the late Oli-

gocene (Arikareean) Cabbage Patch beds of southwest-

ern Montana. The lamellae of Planorbiila powelli (Fig-

ure 6) are also of the pattern characteristic of modern

P. wheatleyi. i.e., with a vertically oriented upper palatal

lamella When compared in series, an evolutionary se-

quence, beginning with Oligocene Planorbiila powelli.

and extending through P. nelmiskensis to P wheatleyi

is strongly suggested. This also suggests that P armigera

may be a late offshoot of the older P. wheatleyi lineage.

The slight inclination of the upper palatal lamella noted

with the otherwise typical P. nebraskensis from the

Nuckols County locality may represent the beginning ot

Page 24

THE NAUTILUS, Vol. 108, No. 1

Figures 1-6. Labial lamellae of planorbulid snails, greatK en-

larged. sp=suprapalatal; up=iipper palatal; lp=lo\ver palatal;

b=basal. 1. Plariorbula armigera (Sa\ 1821). 2. P. wheatleyi

(Lea 1858). 3. P. riebraskensis Leonard 1948, hypotype. 4. P.

nelwaskcnsis. Localit\ NO- 104, 5. P. ni'braskrnsis. Locality

KX-109, Blancan 6. Planorlnila pouclli, Oligoeene (Arika-

reean) Cabbage Patch beds. F"igiires 1 and 2 alter Baker 194.5,

Plate 76.

Figures 7, 8. .-Xpertural lamellae of P ncliKiskcnsis, aiiproxi-

niately 20X, both from lot PPLN 1027, Locality K\-145 (Leon-

ard's type locality). 7. Lamellae « ithin aperture ot small spec-

imen. 8. .Aperture of small specimen broken back to radial of

apertnral lamellae.

a divergence that led to the lamellar pattern of modern

P. armigera. Planorbula vulcanata Leonard 1948, a fossil

of mid-Pleistocene age (ca. (5OOKA) from southwestern

Kansas and adjacent Oklahoma, appears to be the oldest

confirmed planorbulid with typical P. armigera denti-

tion. This suggested lineage results, however, in a dis-

tributional inconsistency. Oligocene Planorlnila powelli

were toinid to have existed in a climate similar to that

preferred by modern P . wheatleyi, which is found only

in the humid mesothermal climatic zone of the south-

eastern United States. Planorbula riebraskensis appar-

ently preferred a cooler, humid, microthermal climate.

Planorbula armigera, on the other hand, is a very tol-

erant species, widely distributed from Louisiana to the

Northwest Territories of Canada, currently occupying

both mesothermal and microthermal climatic zones.

LITERATURE CITED

Baker, F. C. 1945. The molluscan famiK Planorbidae: Col-

lation, revision and additions by H. J. \'an Cleve. L'ni-

versily of Illinois Press, Urbana, 5.30 p.

Bolick, M R, 1991. Pollen and spore anaKsis of sediment

from the .\lbert .^hrens Locality, a medial Pleistocene

vertebrate paleontolog\ site (abst). Proceedings of the Ne-

braska .Academy of Sciences 1991:55.

Dawson, G. M. 1875. Land and fresh-water Mollusca, col-

lected during the summers of 1873-74, in the vicinit\ of

the forty-ninth parallel- Lake of the Woods to the Rock\'

Mountains, p. 347-350. (In) North .American Boundary

Commission, 1872-1876, Report on the geolog\ and re-

sources of the region in the vicinity of the forty-ninth

parallel, from the Lake of the Woods to the Rocky Moun-

tains, with lists of plants and animals collected, and notes

on the fossils. B. Westerman & Co., New York.

Hannibal, H. 1912. A synopsis of the Recent and Tertiary

freshwater Mollusca of the Californian Province. Pro-

ceedings of the Malacological Societs 10:112-211,

Lea, I, 1858, Descriptions of a new Helix and two new plan-

orbes. Proceedings of the .^cademv of Natural Sciences of

Philadelphia 10:41,

Leonard, A, B. 1948. Five new Yarmouthian planorbid snails.

The Nautilus 62:41-47,

Leonard. .\, B, 1950, .\ Yarmouthian molluscan fauna in the

midcontinent region of the I'nited States. Uni\ersit\ of

Kansas Paleontological Contributions, Mollusca. .\rticle 3.

48 p.

Pierce. H. G, 1990, Two unusual gastropods from late Plio-

cene lakes in northeast Nebraska, The Nautilus 104(2):53-

56,

Pierce, H, G. 1993. The nonmarine mollusks of the late Oli-

gocene-early Miocene Cabbage Patch fauna of western

Montana. III. Aquatic mollusks and conclusions. Journal

of Paleontology 67(6):980-993.

Say, T. 1821 Descriptions of univalve shells of the United

States. Journal of the .\cademy of Natural Sciences of

Philadelphia 2(1): 149-179.

Taylor, D. W 1954. Nonmarine mollusks from Barstow For-

mation of southern California. U. S. Geological Survey

Professional Paper 254-C:67-80.

Ta\lor, D. W. 1960. Late Cenozoic molluscan faunas from

the High Plains. U. S. Geological Survey Professional Paper

337. 94 p

Ta\lor, 1), W 1966, Summarx of North American Blancan

nonmarine mollusks, Malacologia 4(1): 1-172,

\oorhies. M, R, and R G, Corner 1991. Paleoclimatic im-

plications of a new medial Pleistocene vertebrate site in

south-central Nebraska; Irvingtonian mammals from the

Albert Ahrens Locality (abst). Proceedings of the Nebraska

,\ca(leni\ of Sciences 1991:63.

THE NAUTILUS 108(l):25-26, 1994

Page 25

Effects of Preservatives on Wet-weight Biomass of the Asiatic

Clam, Corbicula fluminea

Michael E. Smith

Gerry A. Lanfair

Department of Biology

Valdosta State University

Valdosta, GA 31698 USA

The Asiatic clam, Corbicula fluminea (Miiller, 1774) was

first collected in North America from Vancouver Island,

Canada in 1924 and in the United States from near the

mouth of the Columbia River during 1938 (Counts, 1986;

McMahon, 1982). Since its introduction, C fluminea has

spread to the east coast of the United States (Counts,

1986) and into South America (Ituarte, 1981) becoming

an important component of freshwater ecosystems. The

Asiatic clam is often found in high densities creating

problems such as biofouling of industrial water intakes

and power plant cooling systems (Isom, 1986). Corbicula

fluminea is also recognized as an important biological

monitor in freshwater ecosystems (Doherty, 1990).

When sampling C. fluminea, individuals are usually

preserved in the field for later laboratory analy sis. Pres-

ervation can cause changes in wet- weight biomass (Don-

ald & Paterson, 1977; Herke, 1973; Howmiller, 1972;

Landahl & Nagell, 1978; Mills et ai. 1982; Stanford,

1973; Wiederholm & Eriksson, 1977; Williams & Robins,

1982) that affect estimates of population parameters such

as standing crop and production. Thus, the purpose of

this study was to determine the effects of two common

preservatives, 70"^' ethanol and 10"^ neutral buffered for-

malin, on wet-weight biomass of the Asiatic clam, Cor-

bicula fluminea.

Specimens of Corbicula fluminea were collected on 6

February 1993 from the confluence of Shellstone Creek

and the Ocmulgee River, Blecklev Countv, Georgia

(32°25'35"N, 83°28'50"W). Induiduals were removed

from their shells and live wet-weights were determined

prior to preservation in 70'^- ethanol (n = 18) or 10*^' neu-

tral buffered formalin (n=18). Formalin and 10'"c ethanol

solutions were prepared b\ diluting 37 ''c formaldehyde

solution and 95% ethanol with distilled water; the 10%

formalin solution included 4 g/l sodium phosphate mono-

basic and 6.5 g/l sodium phosphate dibasic to produce

neutral buffering. All specimens were blotted (Donald

& Paterson, 1977; Wiederholm & Eriksson, 1977),

weighed to the nearest 0.1 mg and placed individually

into 100 ml of preservative at room temperature (25 ±

1°C). Preserved specimens were soaked in distilled water

for 30 minutes (Howmiller, 1972; Mills et ai, 1982),

weighed, and returned to their individual containers.

Comparison of live wet-weights between the two groups

was by the use of a t-test. Means, 95% confidence intervals

and linear regressions were calculated from arcsine trans-

formed data. Homogeneity of regression slopes was test-

ed to determine if there was a difference in weight loss

over time between the formalin and ethanol groups (So-

kal & Rohlf, 1981).

Live wet-weights were not significantly different

(P>0.05) between the formalin and ethanol groups. Mean

weight loss proceeded more rapidly in the ethanol pre-

served specimens and after 163 days, was 41.9%- (40.7-

43.1%; 95% C.I.) and 21.6% (20.3-22.9%; 95% C.I.) for

the ethanol and formalin groups, respectively (Figure 1).

Stabilization of mean weight loss occurred at appro.xi-

mately day 77 for the ethanol group with only a 2.9%

weight loss over the next two samplings (= 86 days). The

formalin group stabilized after day 163; wet- weight bio-

mass was measured at day 385 for the formalin group

only and mean weight loss was 22.1%, a loss of 0.5%' since

da\' 163. Regression equations were y=0.0434-f0.604x,

r-=0.961 for the formalin group and y=0.0265 + 0.0690x,

r-=0.985 for the ethanol group (Figure 1). Equations are

only descriptive for the preservatives and temperature

range used in this study. The slopes of the two lines were

significantly different (P<0.001) indicating that rate of

weight loss was different between the two groups.

Our study demonstrates that the two most commonly

used preservatives for freshwater benthic organisms, 70%

ethanol and 10% neutral buffered formalin, significantly

decrease wet-weight biomass of Corbicula fluminea . Use

of preserved specimen biomass as an accurate determi-

nation of live biomass, therefore is inappropriate. Wet-

weight losses for C. fluminea fall within the range ob-

served in other studies for freshwater invertebrates

(Donald & Paterson, 1977; Howmiller, 1972; Landahl &

Nagell, 1978; Stanford, 1973; Wiederholm & Eriksson,

1977). Howmiller (1972) found, as we did, that formalin

preservation resulted in less wet-weight loss compared

to 70% ethanol.

We recommend that the regression equations be used

to convert preserved wet-weight biomass to live wet-

weight biomass if 10% neutral buffered formalin or 70%

ethanol is used in the temperature range of 25 ± 1°C.

Page 26

THE NAUTILUS, Vol. 108, No. 1

3

O

100 125

TIME (DAYS)

175

30

S2 10

lij

^ 5

3

O

50 75 100 125

TIME (DAYS)

175

0.4 0.6 0.8

1/TIME (DAYS)

Figure 1. Mean cumulative percent weight loss over time for

Corbictda fluminea preserved in 10% ethanol and 10'^^ neutral

buffered formalin Error bars represent 95*^ confidence inter-

vals, n=18. Relationship between the reciprocal of time spent

in preservative and the reciprocal of mean cumulative weight

loss for Corbicula fluminea at 25 ± 1°C.

Also, the regression equations are only valid up to 163

days of preservation. Conversion to live wet-weights

would be especially beneficial for comparison with other

studies where similar equations were generated. If re-

gression equations are not used, we recommend using

10% neutral buffered formalin as the preservative for

Corbicula fluminea in ecological studies based on lower

weight loss.

ACKNOWLEDGMENTS

We thank Dr. W. J. Loughry for preparing the figure.

LITERATURE CITED

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

invasion of the United States by Corbicula fluniinea (Bi-

valvia: Corbiculidae). American Malacological Bulletin

Special Edition No. 2:7-39.

Doherty, F. G. 1990 The .\siatic clam, Corbicula spp., as a

biological monitor in freshwater environments. Environ-

mental Monitoring and .\ssessment 15:143-181.

Donald, G L. and C G. Paterson. 1977 Effect of preservation

on wet weight biomass of chironomid larsae Hydrobiol-

ogia 53:75-80.

Herke, W. H. 1973. Spin-drying of preserved fishes and ma-

croinvertebrates. Transactions of the American Fisheries

Society 102:643-645,

Howmiller, R. P. 1972. Effects of preservatives on weights of

some common macrobenthic invertebrates Transactions

of the American Fisheries Societv 101:743-746.

Isom, B. G. 1986. Historical review of Asiatic clam {Corbi-

cula) invasion and biofouling of waters and industries in

the Americas. American Malacological Bulletin Special

Edition No. 2:1-5.

Ituarte, C. F. 1981. Primera noticia acerca de la introduccion

de pelecipodos Asiaticos en el area Rio Platense. Neotro-

pica 27:79-82.

Landahl, C.-C. and B. Nagell 1978 Influence of the season

and of preservation methods on wet- and dr\ weights of

larvae of Chirononius plumosus L. Internationale Revue

der gesamten Hydrobiologie 63:405-410

McMahon, R. F. 1982. The occurrence and spread of the

introduced Asiatic freshwater clam, Corbicula fluminea

(Miiller) in North America: 1924-1984. The Nautilus 96:

134-141.

Mills, E. L., K. Pittman, and B. Munroe. 1982. Effect of

preservation on the weight of marine benthic inverte-

brates. Canadian Journal of Fisheries and .\quatic Sciences

39:221-224,

Sokal, R. R. and F, J, Rohlf , 1981. Biometrv 2nd edition. \V.

H. Freeman and Co., New York

Stanford, J. \ 1973. A centrifuge method for determining

live weights of aquatic insect larvae, w ith a note on weight

loss in preservative Ecolog\ 54:449-451,

Wiederholm, T. and L. Eriksson. 1977 Effects of alcohol-

preservation on the weight of some benthic in\ ertebrates.

Zoon 5:29-31

Williams, R. and D B. Robins. 1982. Effects of preservation

on wet weight, dry weight, nitrogen and carbon contents

of Calanus helgolandicus (Crustacea: Copepoda). Marine

Biology 71:271-281.

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T H E t7N AUT I L U S

CONTENTS

Volume 108, Number 2

March 7, 1995

ISSN 0028-1344

Bruce A. Marshall

Roland Houarl

A Review of the New Zealand Recent Species of Poirieria

Jousseaume, 1880 (Mollusca: Gastropoda; Muricidae) with

Description of a New Species 27

A New Species of Charitodoron (Gastropoda: Mitridae)

from Mozambique, with Notes on C. veneris (Barnard,

1964) 34

Taxonomic Notes on Kenyonia Brazier and Conopletira

Hinds (Gastropoda: Conoidea) 37

Four New Genera for Northeastern Pacific Prosobranch

Gastropods 39

Population Structure, Growth and Fecundity of Melampus

bidcntatus (Sa\ ) from Two Regions of a Tidal Marsh

Complex in Connecticut 42

48

R. >. Kilburn

Donn L. Tippett

John K. Tucker

James H. McLean

Jessica A. Spelke

Paul E. Fell

Lucille L. Helvenston

News and Notices

THE NAUTILUS 108(2):27-33, 1995

Page 27

A Review of the New Zealand Recent Species of Poirieria

Jousseaume, 1880 (Mollusca: Gastropoda: Muricidae) with

Description of a New Species

Bruce A. Marshall

Museum of New Zealand Te Papa

Tongarewa

P O. Box 467

Wellington, New Zealand

Roland Houart

Research Associate

Institut Royal des Sciences Naturelles

de Belgique

Vautierstraat 29

B-1040 Brussels, Belgium

ABSTRACT

Poirieria syrinx n. sp. is described and compared with P. ze-

landica (Quoy & Gaimard, 1833) and P. hopua Dell, 1956.

Poirieria syrinx is recorded from Early Pleistocene beds at

Palliser Bay, and living at 482-786 m off the east coast of the

North Island. It is locally sympatric with P. zelandica, which

is widely distributed off New Zealand at 0-540 m depth Chal-

lenger Expedition specimens of P. zelandica reputedly from

Tongatabu and the Kermadec Islands are considered to have

been translocated from Queen Charlotte Sound, New Zealand.

Poirieria hopua occurs at 490-1006 m off the east coast of the

South Island. Some groups with similar species are commented

upon, including Pagodula Monterosato, 1884 and Enixotro-

phon Iredale, 1929 The type species of Enixotruphon (Tro-

phon carduelis Watson, 1883) is newly recorded from New

Zealand, and its radula is illustrated for the first time.

Key words: Mollusca, Gastropoda, Muricidae, Poirieria. new

species.

INTRODUCTION

The present contribution was initiated when we recog-

nized an undescribed species of Poirieria from the con-

tinental slope of the North Island of New Zealand. The

new species differs markedly from the well known spe-

cies P. zelandica (Quoy & Gaimard, 1833) and the rare

.P. kopua Dell, 1956 in having tubular shell spines.

ACRONYMS

AMS The Australian Museum, Sydney

BMNH The Natural History Museum, London

MNHN Museum National d'Histoire Naturelle, Paris

MNZ Museum of New Zealand, Wellington

NZGS Institute of Geological and Nuclear Sciences,

Lower Hutt

NZOI National Institute of Water and Atmospheric

Research, Wellington

SYSTEMATICS

Order Neogastropoda Thiele, 1929

Superfamily Muricoidea Rafinesque, 1815

Family Muricidae Rafinesque, 1815

Subfamily Muricinae Rafinesque, 1815

Genus Poirieria Jousseaume, 1880

Poirieria Jousseaume, 1880335. Type species (by original des-

ignation): Murex zelandicus Quoy & Gaimard, 1833; Plio-

cene-Recent, New Zealand.

Remarks: The radula of Poirieria species is morpholog-

ically very similar to those of the Recent European mur-

icine Trophon echinatus (Kiener, 1840) (Bouchet &

Waren, 1985, fig. 333) and a number of bathyal Indo-

West Pacific species (mostly undescrit)ed — AMS, MNHN,

MNZ), among them Trophon carduelis Watson, 1883

(Watson, 1886, pi. 10, fig. 7), the type species of Enix-

otrophon Iredale, 1929 (type locality New South Wales:

here newly recorded from off the North Island and the

west coast of the South Island, New Zealand, living at

676-1217 m— MNZ, NZOI) (figures 1-4). Some of these

have even longer shoulder spines than most Poirieria

species, though all differ in having the shoulder spines

set closer to the rim of each varix, and radular teeth that

are larger in shells of comparable size. Neither Poirieria

species nor Trophon echinatus or T. carduelis appear to

be closely related to Trophon Montfort, 1810 (type spe-

cies Buccinum geversianum Pallas, 1774; Recent, south-

ern South America) or to Boreotrophon Fischer, 1884

(type species Murex clathratus Linnaeus, 1758; Recent,

northeastern Atlantic), both of which differ in details of

central radular tooth morphology (Harasewych, 1984;

personal observation). Trophon echinatus is closely sim-

ilar to Murex vaginatus De Cristofori and Jan, 1832

Page 28

THE NAUTILUS, Vol. 108, No. 2

Figures 1-4. Radulae of New Zealand Poirieria and Pagodula species. 1. Poirieria zelandica (Quoy & Gaimard, 1833), subadult,

off Matakoa Point, Hicks Bay, 99-102 m, MNZ M. 60498, shell height 30 0 mm 2. Poirieria syrinx new species, holotype (adult).

3. Poirieria kopua Dell, 1956, adult paratype, Chatham Rise, .530 m, MNZ M. 10506. shell height 1-4.7 mm 4. Pagodula carduelis

(Watson, 1883), adult, off Gisborne, 913-750 m, NZOI sta. E719, shell height 50.5 mm. Scale bars 1-3 = 100 Mm, 4 = 1.0 mm.

(Pliocene, Italy), which is generally interpreted as the

type species of Pagodula Monterosato, 1884 (Bouchet &

Waren, 1985:126). It seems plausible, therefore, that Pa-

godula is the most appropriate genus group name for T.

carduelis, and thus by implication that Enixotrophon is

a junior synonym.

The type species of Paziella Jousseaume, 1880 (Murex

pazi Crosse, 1869; Recent, Caribbean), and especially

Actinotrophon Dall, 1902 (Trophon actinophorus Dall,

1889; Recent, Caribbean) (see Bayer, 1971, figs. .30, .35D)

are so similar to New Zealand Recent Poirieria species

in shell and radular morphology that separation on these

characters alone seems untenable. Harasewych (1984)

and Kool (1993a, b), however, have shown that shell

characters (unlike anatomy) in Muricidae are subject to

convergence, while Kool (1987) concluded that radular

characters arc largely unrelated to food-type and are

thus a useful clue to phylogenetic relationships. We fol-

low Yokes (1970, 1992) in treating Paziella and Actin-

otrophon as subgenera of Poirieria.

Incidentally, although Paziella was introduced one line

before Poirieria (Jousseaume, 1880:335), Poirieria has

consistently been treated as the senior synonym (e.g.

Yokes, 1970, 1992) following Cossmann (1903), who was

the first reviser according to ICZN Article 24.

Poirieria zelandica (Quoy & Gaimard, 1833)

(Figures 1, 5, 7, 10-15)

Murex zelandicus Quoy & Gaimard, 18.33: 529, pi. .36, fig.

5-7.

Poirieria zelandica Maxwell, 1971 767 (earlier synonymy);

Powell, 1979: 170, pi. .35, fig 2; Beu & Maxwell. 1990:

358, pi. 48K.

Type material: Holotype MNHN: Cook Strait.

Other material examined: About 100 Late Pliocene and

Pleistocene specimens (MNZ, NZGS), and about 1500

Recent specimens in 200 lots (MNZ).

Distribution: (figure 5) Early Pliocene (Opoitian) to Re-

cent. Three Kings Islands southward to Stewart Island,

Challenger Plateau and Chatham Islands, New Zealand,

living at 0-540 ni.

Remarks: This well known species is common through-

out most of its range at 20-150 m depth. Small (imma-

ture) living specimens occur rarely at considerably great-

er depths, particularly in the Bay of Plenty (deepest

record 490-540 m). One of us (B.A.M.) has taken lightly

abraded short-spined specimens on numerous occasions

alive from sand at low tide at the eastern end of Ohope

B. A. Marshall and R. Houart, 1995

Page 29

Figure 5. Map of New Zealand showing distribution of Poir-

ieria zelandica (Quoy & Gaimard, 1833). 200 and 1000 meter

contours indicated.

Figure 6. Map of New Zealand showing distribution of Poir-

ieria syrinx new species (stars) and Poirieria koptia Dell, 1956

(solid circles). 200 and 1000 meter contours indicated.

Beach, but these had probably been transported inshore

by storms. There is considerable variation in the length

of the spines (figures 10-14), which tend to be longest

in specimens from muddy substrata, and shortest in spec-

imens from coarser substrata. Grading southward through

intermediate forms, specimens from off the southern

South Island and Stewart Island (figure 11) attain about

half the size of the largest northern specimens, and are

more stoutly built, often with a few low rounded den-

ticles within the outer lip. In their stout build the Recent

southern form approaches Late Pliocene (Mangapanian)

and Early Pleistocene (Nukumaruan) forms of P. zelan-

dica (figure 15), though the fossils are consistently thick-

er, have more strongly dentate outer lips, and attain the

size of Recent northern specimens. It should be appre-

ciated that all of the fossils are from the southern half

of the North Island, and since shells of equivalent age

are unknown from further north (due to lack of expo-

sures), it is unknown whether northern Late Pliocene-

Early Pleistocene shells were more lightly built. Maxwell

(1971) concluded that P. zelandica evolved gradually

from P. primigena Finlay, 1930, a species in which the

shoulder is set lower on the whorls and that ranges from

Late Oligocene (Duntroonian to Early Pliocene (Opoi-

tian).

Watson's (1886:157) records of P. zelandica from

Challenger stations 171 (north of Raoul Island, Ker-

madec Islands, 1097 m) and 172 (off Tongatabu, 18 m,)

are highly anomalous since, apart from the Challenger

Plateau record, the species has never been obtained from

beyond the New Zealand continental shelf. These spec-

imens (BMNH 1887.2.9.546-7, 1887.2.9.548) are per-

fectly accordant with the specimens from Challenger

station 167A (Queen Charlotte Sound, New Zealand, 18

n, BMNH 1887.2.9.544-5) and numerous additional

specimens from the vicinity (MNZ). Although it seems

clear that the specimens from stations 171 and 172 were

introduced through entanglement in the nets or ropes,

it is significant that they were not detected at stations

168-170, presumably through oversight or the use of a

different trawl. According to the Narrative of the Cruise

(Tizard et ai, 1885) and the Summary of Results (Mur-

ray, 1895), stations 168-171 were sampled by trawl and

station 172 by dredge. According to Tizard et al. (1885:

1012) station 167A was by dredge alone, but in fact there

were also two trawlings at this station (Murray, 1895:

Page 30

THE NAUTILUS, Vol. 108, No. 2

593), which would account for the presence of contam-

inants in both dredge and trawl stations.

Poirieria syrinx Marshall and Houart, new species

(figures 2, 6, 8, 16, 17)

Poirieria zelandica - Dell, 1962:76 (not Quoy & Gaimard, 1833).

Poirieria kopua - Dell, 1963:212 (not Dell, 1956); Powell, 1979:

170 (text in part).

Description: Shell of medium size for the genus, length

up to 45 mm, broadly fusiform, spire slightly higher than

aperture, suture well impressed, glossy, white or salmon

pink. Protoconch 1.20 mm wide, taller than broad, of

l'/4-iy4 convex whorls, smooth, glossy, terminal varix

prominent. Teleoconch of up to 5Vi convex whorls, suture

well impressed. Varices thin, sharp-edged, prominently

spinose, numbering 5 on 1st whorl, 5 on 2nd, 5 or 6 on

3rd, 5 or 6 on 4th, and 4 or 5 on 5th whorl. Spines slender,

sharp, gently curved, bases set behind varical rim, shoul-

der spine strongly posterior; leading edge open to varical

rim at first, becoming closed after 3rd whorl so that

proximal part of each spine is fully tubular. Shoulder

spine row the most prominent, situated medially on ear-

liest whorls, then supramedially. Secondary spines ab-

apical, 3, relative sizes variable, about equidistant, com-

mencing after 3rd whorl, bases centered well in front of

that of shoulder spine, adapical spine row exposed on

spire, set about midway between shoulder spine row and

suture or abapically, abapical spines of previous whorl

in front of aperture. Aperture roundly ovate. Inner lip

thin, smooth, fully contacting or free over abapical half.

Figures 7-9. Outer lip profiles of New Zealand Poirieria

species. 7. Poirieria zelandica (Quoy & Gaimard, 1833), Or-

chard Bay, Marlborough Sounds, 29 m, MNZ M. 45110, shell

height 28.0 mm. 8. Poirieria syrinx new species, holotype, MNZ

M. 117782, shell height 26.0 mm. 9. Poirieria kopua Dell, 1956,

paratype, Chatham Rise, 530 m, MNZ M. 10506, shell height

14.7 mm. Scale bars = 4 0 mm.

Figures 10-18. Shells of New Zealand Poirieria species. 10-

15. Poirieria zelandica (Quoy & Gaimard, 1833) 10. Holotype,

MNHN, shell height 52.0 mm. 1 1. Port Pegasus, Stewart Island,

40-46 m, MNZ M. 26617, shell height 32 7 mm 12. Off Poor

Knights Islands, MNZ M. 83858, shell height 54.0 mm. 13.

Orchard Bay, Marlborough Sounds. 29 m, MNZ M.451 10, shell

height 28.0 mm. 14. Off Waiheke Island, 40 m, MNZ M 35635,

shell height 57.5 mm. 15. Cliffs west of Whangaimoana, Palliser

Bay, Early Pleistocene (Nukumaruan), MNZ M. 40361, shell

height 32.8 nmi. 16, 17. Poirieria si/rini new species. 16. Cliffs

west of Whangaimoana, Palliser Bay, Early Pleistocene (Nu-

kumaruan), MNZ M. 91795, shell height 25.8 mm (incomplete,).

17. Holotype, MNZ Ml 17782, shell height 26.0 mm 18. Poir-

ieria kopua Dell, 1956, paratype, Chatham Rise, 530 m, MNZ

M. 10506, shell height 14.7 mm.

B. A. Marshall and R. Houart, 1995

Page 31

Page 32

THE NAUTILUS, Vol. 108, No. 2

Outer lip thin at rim, internally slightly thickened, smooth,

simple. Siphonal canal long, semitubular, spineless, gent-

ly curved, 2 or 3 previous canals retained. Outer shell

layer (intriticalx) hard, dense, weakly or locally obscurely

spirally lirate, translucent, glossy, becoming chalky

through erosion.

Animal with small subparallel cephalic tentacles, eyes

on outer edges at about midlength, subcylindrical behind

them, tapered before. Male unknown. Radula (figure 2)

similar to that of Poirieria zelandica.

Type material: Holotype (M. 117782,) and 2 paratypes

MNZ: BS761 {R119), 37°22.0'S, 176°40'E, 37 km E of

Mayor I., New Zealand, alive, 616-666 m, January 24,

1979, R.V. Tangaroa (type locality). Other paratypes

(10): 1364, 34°46'S, 174°05.8'E, NE of Cavalli I. alive,

492 m, November 20, 1977, R.V. Tangaroa (1 NZOI);

24 km NE of Plate I., alive, 622-585 m, October 29,

1962,M.V. Ikatere (1 MNZ); BS741 (R99), 37°20.6'S,

176°28.0'E, 17 km E of Mayor I., alive, 482-550 m,

January 22, 1979, R.V. Tangaroa (3 MNZ); 37°31.64'S,

176°55.35'E, SW of White I., alive, 500-518 m, January

20, 1993, F.V. Kaharoa (1 MNZ); VUZ97, 41°33'S,

174°57'E, off Palliser Bay, alive, 786 m, August 28, 1957;

(3 MNZ, 1 R. Houart collection).

Other material examined: Cliffs west of Whangai-

moana, Palliser Bay, New Zealand, 1971-73, B.A. Mar-

shall, Early Pleistocene (Nukumaruan) (1 specimen).

Distribution: (figure 6): Early Pleistocene (Nukumaruan)

of Palliser Bay, and Recent off northeastern North Island

and Cook Strait, New Zealand, living at 482-786 m on

mud.

Remarks: Compared with Poirieria zelandica, which it

most resembles in shell morphology, P. syrinx n. sp. dif-

fers in having the bases of the spines set further behind

each varical rim, and in that the proximal parts of each

spine become fully tubular after the third teleoconch

whorl. Other shell differences include the smaller size

(maximum length 45 mm vs. 71.5 mm), the larger pro-

toconch (width 1.2 mm vs. of 0.9-1.0 mm), the more

prominent spiral microlirae, and the absence or extreme

weakness of spiral swellings between the bases of the

spines. The animal (preserved material) differs markedly

from that of P. zelandica in having considerably shorter

cephalic tentacles that are almost parallel instead of

strongly divergent, with eyes situated closer to the head.

Unfortunately the male of P. syrinx n. sp. was not avail-

able for comparison of the penial morphology. Their

radulae are similar (figures 1, 2). Poirieria syrinx n. sp.

and P. zelandica have overlapping geographic and

bathymetric ranges, and the two species have been ob-

tained living together (37°20.6'S, 176°28.0'E, off Mayor

Island, 482-550 m, MNZ M.60247, 95064).

Poirieria kopua differs from P. syrinx n. sp. in attain-

ing a smaller size (to 19.0 mm high), in having a larger

protoconch (width 1.5-1.7 mm), in having a fully open

shoulder spine on each varix, and in lacking secondary

spines. Judging from their protoconchs, both species have

non-planktotrophic larval development (probably di-

rect). They appear to be allopatric, though they could

conceivably occur together off Cape Campbell. None of

the New Zealand Tertiary species have tubular spine

bases (Maxwell, 1971). The Early Pleistocene (Nuku-

maruan) specimen (figure 16) is indistinguishable from

Recent shells. Reus (1967) conclusion that the Whan-

gaimoana beds were deposited at 400-600 meters depth

is accordant with the known bathymetric range of this

species.

Etymology: Greek syrinx (pipe).

Poirieria kopua Dell, 1956

(figures 3, 6, 9, 18)

Poirieria kopua Dell, 1956:114, pi. 16, figs. 161, 162, 165; Max-

well, 1971:771, figs. 26, 27; Powell, 1979:170 (in part), fig.

42/1.

NOT Poirieria kopua Dell, 1962:76; Dell, 1963:212 (=P. syrinx

n. sp.).

Type material: Holotype MNZ M.9777 and 2 paratypes

(MNZ and Canterbury Museum, Christchurch): CLE.

sta. 59, 43°38'S, 177°19'E, Chatham Rise, alive, 530 m,

February 11, 1954, M.V. Alert.

Other material examined: (11 specimens) BS559, 43°14'S,

173°39'E, wall of Pegasus Canyon, off Kaikoura, New

Zealand, dead, 1006-512 m, September 27, 1976, R.V.

Acheron (2 MNZ); BS201, 44°45.6'S, 171°05'E, off Taia-

roa Head, dead, c. 549 m, Januarv 23, 1957, M.V. Alert

(3 MNZ); BS582, 45°46'S, 171°03'E, off Taiaroa Head,

dead, 660 m, September 1, 1976, R.V. Munida (5 MNZ);

Mu 70-45, 45°50'S, 171°01'E, off Taiaroa Head, dead,

540-490 m, October 22, 1970, R.V. Munida (1 NZGS).

Distribution: (figure 6) Chatham Rise and off the east

coast of South Island, from Kaikoura to Taiaroa Head,

New Zealand, 490-1006 m, taken alive at 530 m.

Remarks: The shell of P. kopua differs from that of P.

zelandica and P. syrinx n. sp. in attaining smaller shell

size (to 19.0 mm), in having a larger protoconch (width

1.50-1.70 mm), in lacking secondary spines below the

peripheral spines, and in that the spines are not fully

tubular at their bases. The radula is similar in all three

Recent species of Poirieria (figures 1-3).

ACKNOWLEDGMENTS

We thank Philippe Bouchet (Museum National d'His-

toire Naturelle, Paris,) and Kathie M. Way (The Natural

History Museum, London) for the loan of material, and

Alan G. Beu (Institute of Geological and Nuclear Sci-

ences, Lower Hutt), Philip A. Maxwell (Waimate) and

Emily H. Vokes (Tulane L'niversity, New Orleans) for

comments on the manuscript. Thanks also to Michael

Hall and Norman Heke (Museimi of New Zealand, Wel-

lington) respectively for the photographic printing and

photography, and Wendy St (ieorge (Institute of Geo-

B. A. Marshall and R. Houart, 1995

Page 33

logiai and Nuclear Sciences, Lower Hutt) for assistance

with the scanning electron microscopy-

LITERATURE CITED

Bayer, F.

R.V

Beu,

M. 197L New and unusual mollusks collected by

John Elliott Pillsbuby and R.V. Gerda in the

tropical Western Atlantic. In: Bayer, F M and G.L. Voss

(eds. ). Studies in tropical American Mollusks. University

of Miami Press, Coral Gables, Florida! 11-236.

A. G 1967 Deep-water Pliocene Mollusca from Palliser

Bay, New Zealand. Transactions of the Royal Society of

New Zealand, Geology 5(3):S9-122.

A. G. and P. A. Maxwell 1990. Cenozoic Mollusca of

New Zealand. New Zealand Geological Survey Paleonto-

logical Bulletin 58, 518 pp.

Bouchet, P and A. Waren. 1985. Revision of the Northeast

Atlantic bathyal and abyssal Neogastropoda excluding

Turridae (Mollusca, Gastropoda) Bollettino Malacologico,

Supplemento 1:121-296

Cossmann, M. 1903 Essais de Paleoconchologie Comparee,

5. Cossmann, Paris 215 pp.

R K 1956 The archibenthal Mollusca of New Zealand

Dominion Museum Bulletin 18, 235 pp.

R. K. 1962. Additional archibenthal Mollusca from New

Zealand. Records of the Dominion Museum 4(6):67-76.

R K. 1963. Archibenthal Mollusca from northern New

Zealand Transactions of the Royal Society of New Zea-

land. Zoology 3(201 );205-216.

Harasewych, M G 1984 Comparativeanatomy of four prim-

itive muricacean gastropods: Implications for trophonine

phylogeny. American Malacological Bulletin 3: 11 -26

Jousseaume, F 1880. Division methodique de la famille des

purpurides. Le Naturaliste 42:335-336.

Kool, S. P. 1987. Significance of radular characters in recon-

Dell,

struction of thaidid phylogeny (Neogastropoda: Murica-

cea). The Nautilus 101:117-132.

Kool, S. P. 1993a. The systematic position of the genus Nucella

(Prosobranchia: Muricidae: Ocenebrinae). The Nautilus

107:43-57.

Kool, S. P. 1993b. Phylogenetic analysis of the Rapaninae

(Neogastropoda: Muricidae). Malacologia 35:1.55-259.

Maxwell, P. A. 1971. Notes on some Cenozoic Muricidae

(Mollusca: Gastropoda) from New Zealand, with a review

of the genus Poirieria Jousseaume, 1880. New Zealand

Journal of Geology and Geophysics 14:757-781.

Murray, J. 1895 A summary of the scientific results obtained

at the sounding, dredging and trawling stations of H.M.S

Challenger Challenger Report. A Summary of the Sci-

entific Results 1. 796 pp

Powell, A. W B 1979. New Zealand Mollusca. Marine, Land

and Freshwater Shells. Collins, Auckland. 500 pp.

Quoy, J C R and P. Gaimard. 1833. Voyage de decouvertes

de I Astrolabe, execute par ordre du Roi pendant les annees

1826-1827-1828-1829 sous le commandement de M. J.

Dumont d'Urville. Mollusques. Zoologie 2[ii]:321-686

Tizard, T H , H. N Moseley, J. Y Buchanan and J Murray.

1885. Narrative of the cruise of H. MS Challenger, with

a general account of the scientific results of the expedition

Challenger Report. Narrative 1(2), Appendix 11:1007-1015.

Vokes, E. H. 1970. Cenozoic Muricidae of the Western At-

lantic region. Part 5 - Pterynotus and Poirieria. Tulane

Studies in Geology and Paleontology 8: 1-50

Vokes, E. H. 1992. Cenozoic Muricidae of the Western At-

lantic region. Part 9 - Pterynotus, Poirieria, Aspella, Der-

momurex, Calotrophon, Acantholabia, and Attiliosa, ad-

ditions and corrections Tulane Studies in Geology and

Paleontology 25:1-108.

Watson, R B. 1886. Report on the Scaphopoda and Gaster-

opoda collected by H.M.S. Challenger during the years

1873-76. Challenger Report, Zoology 15, 756 pp.

THE NAUTILUS 108(2):34-36, 1995

Page 34

A New Species of Charitodoron (Gastropoda: Mitridae) from

Mozambique, with Notes on C. veneris (Barnard, 1964)

R. N. Kilburn

Natal Museum

P/Bag 9070

Pietermaritzburg, 3200, South Africa

ABSTRACT

Charitodoron rosadoi n. sp., from off Bazaruto Island, Mozam-

bique, in 200-350 m, is described. This is the first Recent record

of the genus Charitodoron Tomlin, 1932, from tropical waters.

Mitromorpha veneris, Barnard, 1964, is referable to Charito-

doron, and adult examples are described and figured for the

first time.

Key words: Mitridae, southern African, continental shelf,

Charitodoron.

INTRODUCTION

The genus Charitodoron Tomlin, 1932, is distinguished

from other genera of the Mitridae by the total absence

of columellar pleats. Cernohorsky (1976), who revised

the described taxa, recognized three species, all of which

are endemic to the Agulhas Bank, the temperate-water

continental shelf off South Africa. Subsequently, Kilburn

(1986:635, fig. 21) pointed out that the holotype of Mi-

tromorpha veneris Barnard, 1964, from off Zululand, is

actually a worn, juvenile example of a species of Char-

itodoron, but did not discuss its identity further. Most

recently Lozouet (1991) recorded the genus for the first

time from the Upper Oligocene of France, and illustrated

several South African species.

During the Natal Museum Dredging Programme, a

numf)er of samples of a distinctive species of Charito-

doron were collected off Natal and Zululand, confirming

that the range of the genus extends into subtropical wa-

ters. The early whorls in tiiis material agree with the

holotype of the supposed turrid Mitromorpha veneris

(Barnard, 1964). Subsequently, Mr. Jose Rosado of Ma-

puto brought to my attention some specimens taken from

crayfish traps off Mozambicjue, which prove to be an

undescribed species of Charitodoron, and provide tlie

first record of the genus from tropical East Africa.

ABBREVIATIONS

a/h = ratio of aperture length (measured along main

axis) to total shell length.

b/h = ratio of maximum protoconch breadth to its

height.

1/h = ratio of shell breadth to total length.

MHNM = Museu de Historia Natural, Maputo, Mozam-

bique.

NMSA = Natal Museum, Pietermaritzburg.

NMDP = Natal Museum Dredging Programme.

SAMC = South African Museum, Cape Town.

TAXONOMY

Family MITRIDAE

Charitodoron Tomlin. 1932

Charitodoron rosadoi n. sp.

Figures 1, 2

Diagnosis: Whorls almost flat, forming a very narrow

shoulder immediately below suture, left side of base of

body whorl shallowly concave; aperture with greatest

width posterior to middle, columella medially convex,

siphonal notch shallow; base of body whorl with 10-12

irregular spiral lirae, sculpture elsewhere weak and in-

conspicuous, initially of extremely fine axial riblets, be-

coming somewhat stronger on later whorls (20-30 on

penultimate one), but remaining weak and irregular,

forming a row of feeble nodules below suture; cream to

brownish-white, with axial flames of brownish-orange,

interrupted by a pale band at mid-bod\ whorl; base of

body whorl white. Protoconch breadth 0.98-1.00 mm.

Maximum length 16.9 mm.

Description: Shell fusiform (1/h = 0.29-0.35), aperture

narrow, a/h = 0.39-0.42; whorls almost fiat, forming a

very narrow shoulder immediateK below suture, left side

of base of bod\' whorl shallowK concave; base slightly

oblique, outer lip slightly foreshortened relative to base

of columella. Teleoconch whorls approximately 5.4 in

number. Aperture w ith greatest width posterior to mid-

dle, gradualK tapering anteriorly; parietal region shal-

lowly concave, columella medialK con\ex, without trace

of pleats; inner lip w ith callus deposit only in columellar

region, bordered by a longitudinal depression (scarcely

a false umbilicus); outer lip thin [edge damaged in all

examples examined]; sipiional notch shallowly and

obliquely U-shaped.

R. N. Kilburn, 1995

Page 35

3 > ^^

Figures 1-4. Charitodoron rosadoi n. sp. and C. veneris, (Barnard, 1964), 1-2. Charitodoron rosadoi, holotype NMSA L1078,

16.9 X 5.5 mm 3-4. C. veneris, NMSA D4223, off Durban, 130 m, 24.0 x 7 2 mm

Surface superficially smooth, other than spiral ridging

on base of body whorl; under magnification whorls are

seen to be axially ribbed, ribs initially extremely fine,

indeed little stronger than growth lines, on later whorls

strengthening, but remaining weak and irregular; ribs

slightly arcuate, moderately prosocline, spacing irregu-

lar, forming a row of feeble nodules where they termi-

nate on the subsutural berm; 20-30 ribs on penultimate

whorl, becoming obsolete on later part of body whorl.

Extremely fine and faint spiral grooves visible on 2nd

and 3rd whorls; base of body whorl with 10-12 irregular

spiral lirae.

Protoconch somewhat papilliform, limit ill-defined,

evidently about 1.7 whorls; smooth, but fine axial riblets

developing towards termination, and continuing onto 1st

teleoconch whorl; dimensions: breadth 0.98-1.00 mm,

height 0.80-0.88 mm (b/h = 1.14-1.23).

Ground color cream to brownish-white, patterned with

axial flames of brownish-orange, interrupted by a pale

band at mid-body whorl; base of body whorl white

Dimensions: 16.9 x 5.5 mm (holotype).

Range: Known only from the type locality.

Type material: Holotype NMSA L1078/T1 168, off Ba-

zaruto Island (21°40'S; 35°25'E), Mozambique, in c. 200-

350 m, in a crayfish trap. Paratypes 1 and 2, same data.

in MHNM and private collection J. Rosado respectively.

All type specimens appear to have been inhabited by

pagurids.

Remarks: Of described species, Charitodoron rosadoi

shows closest resemblance to C. agulhasensis , (Thiele,

1925) of the Agulhas Bank; in particular, the spire whorls

of the latter species have a similar profile (although slight-

ly more convex, with a weaker subsutural shoulder) and

the color pattern is similar although much more muted;

however, in C. agulhasensis the early whorls are char-

acterized by much stronger spiral sculpture and the body

whorl is much more convex. The ranges of C. rosadoi

and C. agulhasensis are widely separated; the northern-

most locality at which the latter species was found during

the NMDP was off Mendu Point, Transkei (32°22.6'S;

29°00.4'E, in 250-260 m).

Etymology: Named in honor of its discoverer, Mr. Jose

Rosado.

Charitodoron veneris (Barnard,

Figures 3-4

1964)

Mitromorpha veneris Barnard, 1964; 16. Type locality: Off Cape

Vidai, Zululand, 80-100 fathoms [=' 145-180 m].

Charitodoron veneris: Kilburn, 1986:635, fig. 21.

Page 36

THE NAUTILUS, Vol. 108, No, 2

Diagnosis: Whorls convex, suture deep, left side of base

of body whorl markedly concave; aperture with greatest

width anterior to middle, columella flattened, siphonai

notch deep; sculpture of strong, flat-topped spiral lirae,

their interstices with coarse collabral threads, feebly in-

dicated on tops of lirae, 8-1 1 lirae on penultimate whorl,

plus a thin thread below suture; white with axial blotches

of brownish-orange, markings occasionally very pale.

Protoconch breadth 1.08-1.15 mm. Maximum length 24

mm.

Description: Shell fusiform (b/h = 0.29-0.30), with nar-

row aperture (a/h = 0.42-0.49); whorls convex, with

greatest width at basal 0.3 of each whorl, suture deep;

left side of base of body whorl markedly concave; base

slightly oblique, outer lip slightly foreshortened relative

to base of columella. Teleoconch whorls up to 6. Aperture

with greatest width anterior to middle; parietal region

shallowK' concave, columella straight, callus deposit thin;

siphonai notch deep and asymmetrically U-shaped; outer

lip thin.

Sculptured by flat-topped spiral lirae, their interstices

with coarse collabral threads, feebly indicated on tops of

lirae. First whorl with 6-9, close-set lirae, becoming more

widely-spaced on later whorls (intervals often as wide as

lirae); 8-11 lirae on penultimate whorl, plus a thin lira

below suture, base of body whorl with 8-11 narrow lirae

plus 13-18 somewhat angular and irregular lirae on ros-

trum.

Protoconch papilliform, 1.7 whorls, smooth with fine

axial plicae near termination, termination sharply indi-

cated; breadth 1.08-1.15 mm, height 1.00-1.25 mm (b/

h = 0.92-1.05). White with axial blotches of brownish-

orange, markings occasionally very pale.

Dimensions: 24.0 x 7.2 mm (largest individual exam-

ined).

Type material: Holotype SAMC A8750 (Kilburn, 1986:

fig. 21).

Distribution: Continental shelf of Zululand and Natal

(Cape Vidal to Amanzimtoti), 98-320 m (fresh shells).

Material examined: (all NM: NMDP unless otherwise

stated): ZULULAND: S. E. of Neill Peak (Cunge), 320-

340 m, sandy mud (£4023); S. E. of Port Durnford, 310-

320 m, glutinous sandy mud (E3190); same locality, 153

m, mud, stones (E4585); off Matiguhi River mouth, 300

m, soft mud (E8903); same locality, 145 m, mud, shell

rubble (E8791); same locality, 200-220 m, mud, coarse

sand (P:9024). NATAL: off Tongaat Bluff, 100 m, coarse

sand mud (E9817); same locality, 120 m, sandv mud

(E9968); off Sheffield Ik-ach, 150 m, muddy sand (E9237);

same locality, 1 10 m, muddy sand (E9303); S. E. of

Sheffield Beach, 100-105 m, glutinous gray mud (E5038);

N. E. of Umhianga liocks, 98 m. fine sand (S256); same

locality, 107 m (B()297: A. Council); oH^ Durban, 130 m,

sandstone gravel, some rocks (D4223); same locality, 95

m, fine, slightly muddy sand (D4059); same locality, 1 10-

120 m, coarse muddy sand (D3816); same locality, 100

m, very fine muddy sand (B5875); same locality, 104 m

(B6287: A. Council); same locality, 104-110 m, muddy

sand, broken shells (D4018); same locality, 150 m, sand-

stone gravel and some sponge, living (D4172); same lo-

cality, 100 m, slightly muddy sand (D3898); off Umlaas

Canal, 150 m, coarse sand, numerous spatangoids, peb-

bles (D789); same locality, 150 m, coarse sand (D865);

off Amanzimtoti, 260-270 m, medium sand (Dl 188);

same locality, 115-125 m, medium sand (Dl 285).

Remarks: Juveniles of the present taxon were compared

(Kilburn, 1986) with the worn and broken holotype of

Mitromorpha veneris, and found to agree.

Charitodoron veneris is characterized by stronger spi-

ral sculpture than any other Recent member of the genus.

There is a distinct but superficial resemblance in shape,

sculpture and color pattern to a number of members of

the conid genus Daphnclla Hinds, 1844 (s.l), but char-

acters of protoconch and outer lip distinguish them at a

glance.

Empty shells of C. veneris are not rare, although in

adults the outer lip is almost always damaged. The fore-

gut anatom\ of the only example to be taken alive is

presently being examined by Y. Kantor.

ACKNOWLEDGMENTS

This study was supported by a grant from the Foundation

for Research Development (FRD). 1 thank Mr. Jose Ro-

sado of Maputo for his hospitality and for making ma-

terial available. For the loan of type material, 1 am in-

debted to Ms. Michelle Van der Merwe (SAMC). Mrs.

Linda Davis helped in the preparation of plates. Dr. D.

G. Herbert kindly read the manuscript.

LITERATURE CITED

Barnard, K. H. 1964. The work of the s.s. Pieter Faure in

Natal waters, with special reference to the Crustacea and

Mollusca; with descriptions of new species of Mollusca

from Natal Annals of the Natal Museum 16:9-29.

Cernohorsky, W 0. 1976. The Mitridae of the World. Part

1. The subfamily Mitrinae. Indo-Pacific Mollusca 3(17);

273-.528.

Kilburn, R. N. 1986. Turridae of southern Africa and Mo-

zambique. Part 3. Subfamily Borsoniinae. .\nnals of the

Natal Museum 27(2):633-720.

Lozouet, P. 1991. Mollusca Gastropoda: Eumitra recents de

la region neo-caledonieiine et Charitodoron fossiles de

roligocene superieur d Aquitaine (Mitridae). In: Crosnier,

A and P Bouchet (eds.) Resultats des Campagnes MU-

SORSTOM 7 Memoires du Museum national d Histoire

nalurelle [A] 150:205-222.

THE NAUTILUS 108(2):37-38, 1995

Page 3

Taxonomic Notes on Kenyonia Brazier and Conopleura Hinds

(Gastropoda: Conoidea)

Donn L. Tippetl

10281 Gainsborough Road

Potomac. Maryland 20854, USA

John K. Tucker

Illinois Natural History Survey

Long Term Resource Monitoring

Program-Pool 26

1005 Edwardsville Road

Wood River, Illinois 62095, USA

INTRODUCTION

Kenyonia Brazier, 1896 is a monotypic genus that has

been assigned to the families Conidae or Turridae by

various authors. Brazier's description of the type species,

Kenyonia pulcherrima, based on a single, unfigured.

specimen, suggested a morphology so unique that he only

tentatively placed the genus as a relative of Conus. He

stated "This is connected with Conus and Pleurotoma

and may be placed under the former genus for the pres-

ent untill the animal is known." Thought for some time

to be lost, the holotype and only known specimen of K

Figures 1-3. Kenyonia pulcherrima Brazier, 1896 I. Holotype (SAM D6181), New Hebrides, shell length = 27 8 mm. 2.

Radiograph of holotype showing resorption of internal shell structure. 3. Apical view of holotype showing pitting of shoulder sulcus.

Figures 4-7. Conopleura striata Hinds, 1844. 4. USNM 869535, Panglao, Bohol Islands, Phhilippines, shell length = 14 7 mm. 5.

Radiograph showing absence of internal shell resorption. 6. Apical view showing honeycomb pitting on shoulder sulcus. 7. Operculum,

length = 3 mm

Page 38

THE NAUTILUS, Vol. 108, No. 2

pulcherrima was located and figured by Zeidler (1985:

71, fig. 2b), who considered its family allocation uncer-

tain. Powell (1966, with query) included Kenyonia in

the synonymy of Conopleura, also a monotypic genus,

t)ecause he presumed the pits on the shoulder slope re-

ported in the description of Kenyonia to be similar to

those of Conopleura (see figures 3,6.)

Regarded as a turrid unequivocally, the relationships

of Conopleura within the family have been uncertain.

This genus had been grouped with the drilliine genera

by older authors such as Fischer (1883) and Tryon (1884).

However, Thiele (1929) removed it to Cytharinae ( =

Mangeliinae), commenting that this assignment was un-

certain. Powell (1966) referred Conopleura to the sub-

family Mangeliinae. A specimen of C. striata Hinds,

1844, originally in the junior author's collection (USNM

869535), was discovered to contain an operculum. This

finding together with a reevaluation of the shell mor-

phology of this species suggests a different subfamilial

affinity Radiographic examination of the holotype of

Kenyonia indicates a reclassification of that genus is re-

quired. The purpose of this paper is to report the results

of these findings.

RESULTS

An X-ray of the holotype of Kenyonia pulcherrima (fig-

ure 2) clearly demonstrates that it has undergone exten-

sive interior shell remodeling (sensu Kohn et ai, 1979).

Its columella is exceedingly thin. In contrast, a similar

X-ray reveals the columella of Conopleura striata to be

thick and stout (figure 5). The internal portions of body

wall in K. pulcherrima are also resorbed and thin, while

those of C. striata are thick. The operculum of Cono-

pleura striata (figure 7) is leaf-shaped, with a terminal

nucleus.

DISCUSSION

Among the toxoglossate mollusks, extensive interior re-

modeling of the shell is a derived character state (Kohn,

1990) found in the Conidae (sensu Cossmann, 1896 =

Conorbinae/Coninae of Taylor et a/., 1993). Therefore,

K. pulcherrima and the genus, Kenyonia, belong in the

family Conidae since they share this state. The presence

of an operculum in Conopleura striata supports removal

of the genus from Mangeliinae since members of that

subfamily lack opercula. The characteristics of the oper-

culum of C. striata are similar to opercula in members

of the subfamily Drilliinae. Furthermore, the general

shape of the shell and the presence of a subtubular sinus

with parietal tubercle are typical of that subfamily. Based

on these characters, we transfer the genus Conopleura

to the subfamily Drilliinae.

ACKNOWLEDGEMENTS

We thank Karen Gowlett-Holmes of the South Australian

Museum for loan of the holotype of Kenyonia pulcher-

rima. Marc Moore of the Armed Forces Radiobiology

Research Institutute, provided the radiographs. Photo-

graphic prints were made by Richard Harris.

LITERATURE CITED

Brazier, J. 1896. A new genus and three new species of Mol-

lusca from New South Wales, New Hebrides, and Western

Australia. Proceedings of the Linnean Society of New South

Wales 21:345-347.

Cossmann, M 1896 Essais de Paleoconchologie comparee.

Cossmann, Paris. 21-179, figs 1-48, pis. 1-8.

Fischer, P. 1880-87. Manuel de conchyliologie et de paleon-

tologie conchyliologique. F. Savy, Paris, 1369 pp., 23 pis.

Hinds, R. B. 1844-45. The zoology of the voyage of the

H.M.S. Sulphur Volume II. Mollusca. Smith, Elder & Co.,

London, v + 72 pp., 21 pis.

Kohn, A. J. 1990. Tempo and mode of evolution in Conidae.

Malacologia 32(l):55-67, figs. 1-4.

Kohn, A. J, E. R. Myers, and V. R. Meenakshi. 1979 Interior

remodeling of the shell by a gastropod mollusc. Proceed-

ings of the National Academy of Sciences (USA)76(7):

3406-3410, figs. 1-4.

Powell, A. W. B. 1966. The molluscan families Speightiidae

and Turridae. Bulletin of the Auckland Institute and Mu-

seum 5:1-184, pis. 1-23, figs. 1-179.

Taylor, J. D., Y. I. Kantor, and A. V. Sysoev. 1993. Foregut

anatomy, feeding mechanisms,relationships and classifi-

cation of the Conoidea (=Toxoglossa) (Gastropoda). Bul-

letin of the Natural History Museum (Zoology) 59(2): 125-

170, figs. 1-27.

Thiele, J 1929. Handbuch der systematischen weichter-

kunde. Gustav Fisher, Jena. I(l):l-376.

Tryon, G. W. 1884. Family Pleurotomidae. Manual of Con-

chology 6(23-24):151-413, pis. 1-.34

Zeidler, W 1985 Mollusc type-specimens in the South Aus-

tralian Museum 2 GastropodaConidae. Records of the

South Australian Museum 19(5):69-75, figs. 1-3.

THE NAUTILUS 108(2);39-41, 1995

Page 39

Four New Genera for Northeastern Pacific Prosobranch

Gastropods

James H. McLean

Los Angeles County Museum of

Natural History

900 Exposition Blvd.

Los Angeles, CA 90007 USA

ABSTRACT

In order to make the names available for use in a faunistic

revision of the northeastern Pacific gastropod fauna, four new

genera are proposed Naticidae: new genus Benthobulbus. type

species Choristes carpenteri Dall, 1896 Eulimidae; new genus

Pseudosabinella. type species Sabinella bakeri Bartsch, 1917.

Muricidae, Trophoninae: new genus Ocenotrophon, type spe-

cies Murex (Ocinebra?) painei Dall, 1903. Buccinidae: new

genus Retimohnia, type species Mohnia frielei Dall, 1891.

Key words: Northeastern Pacific Gastropoda; systematics; new-

genera.

INTRODUCTION

In the course of preparing a checklist of the northeastern

Pacific gastropod fauna, in which all generic assignments

are being re-evaluated, I have been unable to reconcile

the allocation of a number of species to the genera in

which they have previously been placed. The following

new genera have been recognized and are here intro-

duced in advance of that work.

Family NATICIDAE Forbes, 1838

Benthobulbus McLean, new genus

Type species: Choristes carpenteri Dall, 1896.

Included species: Type species and Choristes coani Mar-

incovich, 1975. Marincovich (1977) treated both species

in detail and illustrated their radulae; both were further

discussed by McLean (1992).

Diagnosis (modified from that of Marincovich, 1977:338,

for "Choristes" Carpenter): Shell small to medium in

size, globose, thin; whorl moderately inflated, spiral

sculpture of weak lirae, suture narrowly channeled. Um-

bilicus narrow to broad, simple. Inner lip slightly thick-

ened, simple, lacking umbilical callus. Parietal callus thin.

Operculum chitinous, entirely filling aperture. Radula

with monocuspate rachidian, one monocuspate lateral

tooth and two monocuspate marginal teeth per half row.

Remarks: McLean (1992:291) assigned the type species

of Choristes Carpenter in Dawson, 1872, to the synon-

ymy of Atnauropsis islandica (Gmelin, 1791), leaving

two living eastern Pacific species treated by Marincovich

without a genus. Marincovich placed Choristes next to

Bulbus Brown in J. Smith, 1839 (type species Bulbus

smithii Brown in J. Smith, 1839; = Natica fragilis Leach,

1819), noting that only Bulbus and the two deep-water

eastern Pacific "Choristes" species had the monocuspate

rachidian tooth. Further remarks on the naticid radula

are given by Bouchet and Waren (1993:753), who noted

that the first marginal of Bulbus has three terminal den-

ticles. The unsatisfactory alternatives to placement of

these two species in a new genus would be to place them

in Bulbus. in which the umbilicus is narrow and filled

with callus, or in Euspira Agassiz, 1838, in which the

rachidian is tricuspate and shells may be larger and thick-

Family EULIMIDAE Troschel, 1853

Pseudosabinella McLean, new genus

Type and only known species: Sabinella bakeri Bartsch,

1917 (synonyms: Alaba catalinensis Bartsch, 1920; Alaba

serrana Smith & Gordon, 1948). See Abbott (1974:126,

fig. 1382).

Diagnosis: Shell thin, whorls rounded, with faintly an-

gulate shoulder; suture deep; final lip inflated and pro-

duced anteriorly; umbilical chink present; lip scars slight-

ly raised, variable in position; early whorls dark brown,

surface of teleoconch tan.

Remarks: Regarding "Sabinella" bakeri, Waren (1992:

189) commented: "This species probably is an eulimid

despite having a rather fragile and irregular shell. I have

examined a specimen with dried soft parts and it has a

ptenoglossate radula, similar to species of Eulirna. It can

provisionally be placed in Eulitna." Pseudosabinella

bakeri differs from typical Eulima in having rounded

whorls, a projecting and flared final whorl and a brown

cast to the shell. In contrast, typical species of Eulima,

as defined by Waren (1984:43; 1992:179), have straight-

sided shells, often with color banding. Waren elected not

Page 40

THE NAUTILUS. Vol. 108, No. 2

to provide a genus for this species until the echinoderm

host becomes known, but now endorses the need for a

genus (pers. comm.). The species is a fairly common and

characteristic member of the Californian faunal prov-

ince, for which the provision of a genus may perhaps

provide impetus toward the eventual discovery of its host.

Familv MURICIDAE Rafinesque, 1815

Subfamily TROPHONINAE Cossmann, 1903

Ocenotrophon McLean, new genus

Type and only known species: Miirex {Ocinebra?) pai-

nei Dall, 1903. See Radwin & D'Attilio (1976:123, pi.

20, pi. 20, fig. 8).

Diagnosis: Shell small (length about 15 mm), sculpture

of numerous lamellar cords of unequal prominence,

crossed by sharply raised ribs, forming nodes at inter-

sections; canal short, closed.

Remarks: The generic allocation of the type species has

been uncertain, starting with Dall's original assertion that

'"it resembles one of the austral trophons in miniature."

Dall (1921:159, pi. 6, fig. 1) retained it in Tritonalia

(ICZN rejected name for what is now Ocenebra Gray,

1847), evidently because of the sealed canal, which is

characteristic of Ocenebra but not of trophonine genera.

Radwin & D'Attilio (1976:123) followed Dall and re-

tained it in Ocenebra but noted that: "The lamellose

shell sculpture and short, stout form suggest affinity with

the southern trophons (e.g. T. geversianus Pallas, 1774);

the fused canal implies an ocenebrine relationship; and

the radula favors neither of these possibilities." D'Attilio

(1980:6) again treated the species and illustrated the rad-

ula, concluding that the radula is in fact close to that of

Trophon geversianus, and reallocated the species in Tro-

phon. That allocation is less than satisfactory because T.

geversianus , type species of Trophon, is large, has a dark

colored interior, has an open canal and occurs in the

intertidal zone of Argentina in the southern hemisphere.

Kool (1993) treated the radula and anatomy of Trophon

geversiansus. Ocenotrophon differs from species of Bor-

eotrophon Fischer, 1884, (in which spiral sculpture is

lacking or does not override the axial lamellae) and from

species of Trophonopsis Bucquoy & Dautzenberg, 1882

(in which spiral sculpture overrides the axial lamellae)

in having a thick final lip and in having a shorter, sealed

canal rather than a relatively long open canal. Species

of Boreotrophon and Trophonopsis tend to merge at the

subgeneric level, whereas Ocenotrophon painei stands

well apart from all other boreal trophonine species.

Family BUCCINIDAE Rafinesque, 1815

Retimohnia McLean, new genus

Type species: Mohnia frielei Dall, 1891. The holotype

has been well figured by Kosuge (1972, pi. 13, fig. 3),

showing also the characteristic pattern of the operculum.

Included species: Urosalpinx carolinensis Verrill, 1884;

Neptunea caelata Verrill & Smith, 1880; Sipho glyptus

Verrill, 1882; Mohnia clarki Dall, 1907; Mohnia micra

Dall, 1907; Mohnia hondoensis Dall, 1913; Mohnia ja-

ponica Dall, 1913; Mohnia robusta Dall. 1913; Mohnia

vernalis Dall, 1913; Cohis (Latisipho) clementinus Dall,

1919.

Diagnosis: Shell small, fusiform; whorls rounded, canal

short, twisted to left; periostracum thick, adherent; dom-

inant sculpture of strong to faint axial ribs on upper half

of whorl, becoming obsolete in some species at later

growth stages; spiral sculpture of fine to moderate spiral

incisions. Initial whorl of protoconch small, not project-

ing; operculum subspiral, nucleus away from anterior

edge.

Remarks: Species grouped in Retimohnia have previ-

ouslv been assigned to Mohnia Friele in Kobelt, 1878,

by Dall (1913), Radwin (1972), and Tiba & Kosuge (1992).

Bouchet & Waren (1985:205) only tentatively included

in Mohnia three northeastern Atlantic species "for which

we could not find a better genus, although we are not

satisfied with their position here " The latter three species

(Urosalpinx carolinensis Verrill, 1884, Neptunea caelata

Verrill & Smith, 1880, Sipho glyptus N'errill, 1882) were

treated in detail by Bouchet and Waren and are here

allocated Retimohnia.

The type species of Mohnia. Fusus mohni Friele, 1877,

has been illustrated by Bouchet & Waren (1985:205, figs.

435, 482, 530-531) and by Tiba & Kosuge (1992:1). In

species of Mohnia (and the subgenus Tacita Lus, 1971,

as treated by Bouchet & Waren, 1985:210), the whorls

are rounded and the suture deep; spiral sculpture pre-

dominates, whereas axial sculpture is limited to fine axial

riblets that produce a fine clathrate effect in early stages.

The characteristic low protoconch was illustrated by

Bouchet & Waren (1985).

Axial sculpture of projecting ribs is the principal sculp-

ture in Retimohnia, although in fi. clementinus it may

be weak. The protoconch of Pacific species of Reti-

mohnia is eroded in specimens I have examined, but is

known from the three Atlantic species illustrated by

Bouchet & Waren. The operculum is similar in both

genera and the genera are evidently related. As is true

of the Atlantic species treated by Bouchet & Waren, the

Pacific species of both genera occur in deep water, at

abyssal depths or the lower continental slope.

Tiba and Kosuge (1992) placed three species in Mohn-

ia that are assignable neither to Mohnia or Retimohnia:

"Chrysodomus" brunneus Dall, 1877, "C." virens Dall,

1877, and "Mohnia" siphonoides Dall, 1913. I retain the

first two in the genus Retifusus Dall. 1916 (type species

Tritonium jessoense Schrenck, 1867, Japan), in which

they were retained by Dall (as a subgenus of Plicifusus

Dall, 1902). These two species are not restricted to deep

water and have stronger spiral sculpture than species of

Retimohnia. The operculum of Retifusus brunneus, as

figured by Tiba & Kosuge (1992, fig. 10) does not have

the nucleus sufficiently distant from the margin to be

considered comparable to that of Retimohnia. Dall (1913:

502) remarked that the species he then described as

J. H. McLean, 1995

Page 41

Mohnia siphonoides from 987 fathoms off the Pribilof

Islands "would certainly be referred to Tritonofusus"

[now Coins Roding, 1798] if it were not for the oper-

culum. I assign the latter species to Coins, and assume

that Dall mixed the operculum with that of a different

species. With the e.xceptions above and the type species

of Mohnia. all other species illustrated b> Tiba & Kosuge

(1992) as Mohnia, are here considered t\pical species of

Retimohnia. Bouchet & Waren (1985) described and

illustrated several typical species of Mohnia. New species

of Mohnia that I have recognized from the northeastern

Pacific will be described separately.

LITERATURE CITED

Abbott, R. T. 1974. American Seashells, second edition. New

York: Van Nostrand Reinhold, 663 pp.

Bouchet, P. & .\. Waren 1985. Revision of the northeast

Atlantic bathyal and abyssal Neogastropoda excluding

Turridae (Mollusca, Gastropoda) Societa Italiana di

Malacologia, Bollettino Malacologica, Suppiemento 1, 12.3-

269.

Bouchet, P. & A. Waren. 1993 Re\ision of the northeast At-

lantic bathyal and abyssal Mesogastropoda. Societa Ualiana

di Malacologia, Bollettino Malacologico, Suppiemento 3,

579-840.

Dall, W. H. 1903. Diagnoses of new shells from the Santa

Barbara Channel, California. Proceedings of the Biological

Society of Washington 16:171-176.

Dall, W. H. 1913. New species of the genus Mohnia from

the North Pacific. Proceedings of the Academy of Natural

Sciences, Philadelphia, 65(2):501-504.

Dall, W. H. 1921. Summary of the marine shell-bearing mol-

lusks of the northwest coast of America, from San Diego,

California, to the polar sea, mostly contained in the col-

lection of the United States National Museum, with illus-

trations of hitherto unfigured species. United States Na-

tional Museum, Bulletin 112, 217 pp , 22 pis.

D'Attilio, A. 1980 Trophon painei (Dall, 1903): An anom-

alous murex. The Festivus 12(l):6-9.

Kool,S. P. 1993. Phylogenetic analysis of the Rapaninae (Neo-

gastropoda: Muricidae). Malacologia 35(2): 155-260.

Kosuge, S. 1972. Illustrations of type specimens of molluscs

described by William Healey Dall (North- Western Pacific

Gastropoda). National Science Museum, Tokyo, 29 pis. and

captions.

Marincovich, L. N. 1977. Cenozoic Naticidae (Mollusca: Gas-

tropoda) of the northeastern Pacific. Bulletins of American

Paleontology 70(294): 165-494, pi. 17-42.

McLean, J. H. 1992. Systematic review of the family Chor-

istellidae (Archaeogastropoda: Lepetellacea) with descrip-

tions of new species. The Veliger 35(4):273-294.

Radwin, G. E. 1972. The systematic position of Urosalpinx

carolinensis Verrill, 1884, with comments on the genus

Mohnia Friele, 1878. Transactions of the San Diego So-

ciety of Natural History 16(16):339-342.

Radwin, G. E. and A D'Attilio. 1976. Murex shells of the

world. Stanford University Press, 284 pp

Tiba, R. and S. Kosuge. 1992. Genus Mohnia Friele, 1877.

North Pacific Shells, no. 18, Occasional Publication of the

Institute of Malacology, Tokyo, 26 pp.

Waren, A. 1984. A generic revision of the family Eulimidae

(Gastropoda, Prosobranchia). The Journal of Molluscan

Studies, Supplement 13, 96 pp.

Waren, A. 1992. Comments on and descriptions of eulimid

gastropods from Tropical West America. The Veliger 35(3):

177-194.

THE NAUTILUS 108(2):42-47, 1995

Page 42

Population Structure, Growth and Fecundity of

Melampus bidentatus (Say) from Two Regions of a

Tidal Marsh Complex in Connecticut

Jessica A. Spelke

Paul E. Fell'

Lucille L. Helvenston

Department of Zoology

Connecticut College

New London, CT 06320, USA

ABSTRACT

We examined papulations of the tidal marsh snail, Melampus

bidentatus Say, within two regions of the Barn Island Wildlife

Management Area in Stonington, Connecticut: a Spartina pat-

ens-dominaied bayfront marsh and a restored impounded val-

ley marsh covered by Spartina alterniflora and forbs. On the

bayfront marsh, 87% of the snails collected from late May

through early November were less than 8 mm in shell length,

whereas 99% of the snails on the restored impounded marsh

exceeded 8 mm in shell length. Snails from both marsh regions

reared in the laboratory on Spartina alterniflora /(orh turf from

the restored impounded marsh grew more rapidly than snails

reared on Spartina patens turf from the bayfront marsh. In

laboratory culture, Melampus exhibited a regular lunar (4-

week) egg-laying periodicity with cycles of egg-laying occur-

ring in late May, late June and late July. Not only did large

snails (9.1-10.0 mm) from the restored impounded marsh de-

posit nearly twice as many egg masses as small snails (6.1-7.0

mm) from the bayfront marsh, but the egg masses produced

by the large snails were significantly larger than those laid by

the small snails.

Key words: Melampus bidentatus, tidal marsh snail, popu-

lation structure, growth, fecundity.

INTRODUCTION

Melampus bidentatus Say, 1822, is a pulmonale snail

that inhabits the higher elevations of tidal salt marshes

along the Atlantic and Gulf of Mexico coasts of North

America (Hausman, 1932; Holle & Dineen, 1957; Russell-

Hunter et al., 1972). This snail often occurs in large

numbers in regions of high marsh covered by Spartina

patens (Ait) Muhi, 1817 (saltmeadow hay), Distichlis

spicata (L.) Greene, 1887 (spikegrass) and Juncus spp..

' To whom correspondence should be addressed.

as well as in well-drained regions dominated by stunted

Spartina alterniflora Loisel., 1807 (saltwater cordgrass).

It is frequently a dominant member of the high marsh

community (Russell-Hunter et al., 1972; Price, 1980;

Hilbish, 1981; Fell et al, 1982; Joyce & Weisberg, 1986).

It has been shown that on a restored impounded tidal

marsh dominated by stunted Spartina alterniflora , the

size of Melampus bidentatus is substantially larger than

on bayfront, Spartina pa^fns-dominated marshes in the

same system. On the restored impounded marsh, 83% of

the snails exceeded 8 mm in shell length and the modal

size was 9.1-10.0 mm. In contrast, on the undisturbed

Spartina patens -dominated marshes, 95% of the snails

were 8 mm in length with a modal size of 6.1-7.0 mm

(FeW etal., 1991). Considering specific areas within these

marshes, the same difference in the sizes of snails was

observed. During the summer of 1990 and the late spring/

early summer of 1991, the most abundant size class of

Melampus within the Spartina patens meadows of an

undisturbed bayfront marsh was 6.1-7.0 mm, whereas

the most abundant size class of this snail in well-drained

areas of the restored impounded marsh covered by stunt-

ed Spartina alterniflora and forbs (herbaceous plants

other than grasses, rushes and sedges) was 9.1-10.0 mm.

Because of this size difference, the shell-free biomass of

Melampus in the first area was only about half that in

the second, even though snail density was somewhat

higher (Peck et al., 1993). Such size differences could be

due in large part to quantitative and/or qualitative dif-

ferences in the food resources available in the two marsh

regions (Rietsma et al., 1988; Peck et al., 1993). How-

ever, the size differences could also result to some extent

from greater size specific predation (Vince et al., 1976;

Joyce & Weisberg, 1986) and/or factors that promote

greater longevity of snails (Crowe & Covich, 1990) on

the restored impounded marsh. Melampus feeds exten-

sively on detritus derived from the marsh grasses but

also consumes algae and animal tissues (Rietsma et al.,

1982; Thompson, 1984). One objective of the present

study was to examine growth of Melampus from the

J. A. Speike et al, 1995

Page 43

Spartina patens meadows of an undisturbed bayfront

marsh and from stunted Spartina alterni flora -dominat-

ed regions of tfie restored impounded marsh when sam-

ples from both groups of snails were maintained in the

laboratory on marsh turf from these two areas. Melam-

pus is a simultaneous hermaphrodite with an ovotestis.

It deposits egg masses on the surface of the marsh peat,

as well as on the stems and leaves of grasses and on the

shells of other Melampus. These egg masses are flattened

hemispheres about 1 to 2 mm in diameter and approx-

imately 0.5 mm high. Each egg mass may contain from

about 500 to more than 1200 eggs (Hausman, 1932; Apley,

1970; Russell-Hunter et al., 1972). A preliminary study

(Balboni-Tashiro et al., 1985) indicates that Melampus

exhibits age (size) specific fecundity. Older snails pro-

duce more egg masses containing more eggs. A second

objective of this study was to compare egg mass depo-

sition by large Melampus from stunted Spartina alter-

nt^Jora -dominated regions of the restored impounded

marsh and by smaller snails from the Spartina patens

meadows of an undisturbed bayfront marsh.

MATERIALS AND METHODS

The study areas are part of the Barn Island Wildlife

Management Area in Stonington, Connecticut, which is

located in the southeastern corner of the state (figure 1).

The marsh complex consists of a series of valley marshes

that border Little Narragansett Bay. One study area was

the westernmost valley marsh which was impounded in

1946-1947 in an attempt to create waterfowl habitat.

This marsh (figure 1, IP) subsequently converted to a

Typha angustifolia L., 1753 (narrow-leaved cattail-

dominated brackish marsh. In 1978 a 5 ft. -diameter cul-

vert was placed in the impoundment dike, and in 1982

a 7 ft. -diameter culvert was added. As a result of the re-

established tidal exchange, typical tidal salt marsh an-

giosperms have recolonized much of the area, with stunt-

ed Spartina alterniflora and forbs providing most of the

plant cover (Sinicrope et al., 1990). The other study area

was an undisturbed Spartina patens -dominated bayfront

marsh situated below the westernmost impoundment

(Wequetequock Cove marsh, figure 1, WC).

In order to examine the size structure of Melampus

populations in the Spartina patens meadows of the bay-

front marsh and stunted Spartina alterniflora-dominai-

ed areas of the restored impounded marsh, all of the

snails contained within two randomly placed 0.25m'

quadrats were collected from each marsh area at about

30-day intervals from late May until early November.

Shell lengths were measured to the nearest 0. 1 mm with

vernier calipers.

For experimental studies, small (6.1-7.0 mm) Melam-

pus were collected from Spartina patens -dominated

regions of the bayfront marsh and large (9.1-10.0 mm)

snails were gathered from an area of the restored im-

pounded marsh covered by stunted Spartina alterniflora

and forbs. The snails were maintained in the laboratory

on clipped marsh turf. They were placed in covered 11

IITTIE

NARRAGANSETT BAY

Figure 1. Map of the western portion of the tidal marsh

complex within the Barn Island Wildlife Management Area in

Stonington, Connecticut (short vertical arrow on inset at upper

left) IP marks the collecting site on the restored impounded

marsh and WC indicates the location of the Wequetequock

Cove Marsh.

cm-diameter finger bowls containing a piece of turf ap-

proximately 6 X 4 X 1.5 cm situated on a 9 cm disc of

Whatman No. 1 filter paper moistened with 15 ppt sea

water. In most cases, 8 snails were placed in each bowl.

The cultures were kept at about 21°C and exposed to

natural photoperiod. The filter paper and bowls were

changed every 2 days and the marsh turf was replaced

every 6 days.

For growth experiments, some of the snails from each

marsh region were placed on Spartina patens turf taken

from the bayfront marsh and others were cultured on

Spartina alterniflora /forh turf from the restored im-

pounded marsh. Each treatment was replicated 5 times

(a total of 40 snails per treatment). The snails were mea-

sured to the nearest 0.1 mm with vernier calipers at

monthly intervals. The experiments were begun 8 June

1992 and continued until 1 November 1992.

In order to quantify egg mass deposition, snails from

each of the two marsh regions were cultured separately

on Spartina patens turf from the bayfront marsh (15

snails per bowl in 1991 and 8 snails per bowl in 1992).

Cultures were initiated 23 May 1991 and 20 May, 10

June and 25 June 1992. Each set of cultures was repli-

Page 44

THE NAUTILUS, Vol. 108, No. 2

o

SHELL LENGTH (mm)

Q

7 8 9 10 11 12 13

SHELL LENGTH (mm)

Figure 2-3. Size-frequency distribution of Melampus biden-

tatus 2. In Spartina patens meadows of the bayfront marsfi

during the reproductive period (20 May to 3 August 1992). N=

1024. 3. In regions of the restored impounded marsh covered

by stunted Spartina alterniflora and forbs during the repro-

ductive period (20 May to .3 August 1992). N = 702.

cated 3 times. Newly deposited egg masses were removed

and counted each day through 31 July.

RESULTS

On the bayfront marsh, 87% of the snails collected

throughout the study were less than 8 mm in shell length,

and the most abundant size class was 6.6-7.5 mm. In

contrast, 99% of the snails on the restored impounded

marsh exceeded 8 mm in shell length, and 10.6-1L5 mm

was the most abundant size class. During the reproduc-

tive period, the minimum length of Melampus on the

bayfront marsh was 4.1-4.5 mm and the most abundant

size class was 6.6-7.5 m (figure 2). Minimum length of

snails from the restored impounded marsh during the

reproductive period was 6.6-7.0 nmi, and less than 1%

of the snails were of this size. The majority of snails (61%)

were 10.1-11.5 mm in shell length (figure 3). Young-of-

the-year snails were not observed on either marsh region

during the study.

Snails from both marsh regions reared in the laboratory

on Spartina alterniflora /f orb turf from the restored im-

pounded marsh grew more rapidly than snails reared on

Spartina patens turf from the bayfront marsh (Table 1).

Mann Whitney tests with values of tied rank (Zar, 1974)

indicated that the difference in growth of Melampus

from the bayfront marsh on turf from the two marsh

regions was statistically significant (U = 0, df = 2, p =

0.01) as was the difference in growth of snails from the

restored impounded marsh on these substrata (U = 23.5,

df = 2, 0.05 > p > 0.02).

In laboratory culture, Melampus exhibited a regular

lunar egg-laying periodicity with cycles of egg-laying

occurring in late May, late June and late July (figure 4).

Egg-laying was usually initiated at the time of the full

moon and continued for about 9 to 14 days. Deposition

of egg masses ceased shortly before the new moon, and

little or no egg-laying was observed between the new

moon and subsequent full moon. Aggregation of snails

was noted 2-3 days before egg-laying began. Egg masses

were deposited most commonly on the surface of the

peat, but they were also attached to the stems of Spartina

patens and occasionally to the shells of other Melampus.

Egg-laying by Melampus collected from the bayfront

marsh on 20 May 1992 was not in synchrony with that

of other snails gathered from the two marsh regions. For

this group, egg-laying began 9 June, six days before the

full moon, and continued at a low rate until 25 June, ten

days past the full moon. A second cycle of egg-laying

was initiated 6 July, eight days before the full moon, and

lasted until 15 July.

The snails from the restored impounded marsh de-

posited nearly twice as many egg masses as those from

the bayfront marsh (Table 2). During the summer of

1992, large snails (9.1-10.0 mm) from the restored im-

pounded marsh maintained under laboratory conditions

produced an average of 128 egg masses per animal dur-

ing the reproductive season, whereas small snails (6.1-

7.0 mm) from the bayfront marsh deposited an average

of 63 egg masses per animal over the same span of time.

Analysis of the data from the third spawning period, for

which there were three different sets of snails from each

marsh region, indicated that the difference in the number

of egg masses deposited by the larger snails of the restored

impounded marsh and the smaller snails of the bayfront

marsh is statistically significant (two-sample t-test: t =

2.9, df = 15, p = 0.012).

The mean size of egg masses produced by snails from

the restored impounded marsh was 3.3 mm ± 0.42 mm

SD (N = 31) and that of snails from the bayfront marsh

was 2.5 mm ± 0.39 mm SD (N = 28). Using a two-

.sample t-test, the difference was found to be highly sig-

nificant (t = 7.58, df = 56, p < 0.0001).

During 1991, large snails from the restored impounded

marsh, maintained at a density of 15 snails per bowl.

J. A. Spelke et al., 1995

Page 45

Table 1. Growth (increase in shell length) of Melampus hidentatus (Say) from a restored impounded marsh (IP) and an undisturbed

bayfront marsh (Wequetequock Cove Marsh, WC) on turf from these two marshes in the laboratory. The experiment extended

from 8 June to 1 November 1992.

deposited an average of 96 egg masses per animal during

the first cycle of egg-laying and the study was terminated

before the cycle was completed. In contrast, during 1992,

large snails cultured 8 per bowl produced an average of

no more than 71 egg masses per animal for any cycle.

The relationship between snail density and egg-laying

should be systematically investigated.

DISCUSSION

On the Barn Island marshes in southeastern Connecticut,

Melampus exhibits a clear lunar periodicity in egg-lay-

ing. Egg mass deposition begins at about the time of the

full moon and continues until shortly before the time of

the following new moon. This pattern of egg-laying dif-

fers from that described for snails at the Little Sippew-

isset Marsh in Falmouth, Massachusetts (Cape Cod). Sev-

eral studies conducted over a 10-yr. period at that location

have demonstrated a semilunar pattern of egg-laying by

Melampus (Apley, 1970; Russell-Hunter et al., 1972;

Table 2. Production of egg masses by Melampus hidentatus

(Say) in the laboratory Small snails were collected from an

undisturbed bayfront marsh (Wequetequock Cove Marsh, WC)

and large snails were gathered from a restored impounded

marsh (IP).

Price, 1979). The semilunar periodicity is well defined

with egg-laying normally taking place for about 4 days

beginning on about the second day following a new or

full moon (figure 5). Spring tides that occur at these times

often immerse the newly deposited egg masses and cover

them with a layer of fine detritus that keeps the eggs/

embryos moist during the period of neap tides (Russell-

Hunter et al, 1972).

The pattern of egg-laying described here does not

appear to be an artifact resulting from laboratory con-

LUNAR PHASES

HEIGHT OF HIGH TIDES

EGG-LAYING PERIODS

n

FULL

o

A LITTLE SIPPEWISSET MARSH

B BARN ISLAND MARSH

Figures 4-5. Egg-laying by Melampus hidentatus 4. Snails

from the restored impounded marsh in relation to the occur-

rence of full moons. Each point represents the average number

of egg masses produced by three bowls of snails. Cultures were

initiated 23 May 1991 and 20 May, 10 June and 25 June 1992.

5. Snails on the Little Sippewisset Marsh in Falmouth, Mas-

sachusetts (after Russell-Hunter et al., 1972) and on the Barn

Island marshes in Stonington, Connecticut in relation to phases

of the moon and the relative height of high tides (marsh sub)-

mergence).

Page 46

THE NAUTILUS, Vol. 108, No. 2

ditions. First, egg-laying by Melampus maintained in

the laboratory follows the same rhythm as that of field

populations of this snail (Russell-Hunter et ai, 1972 and

our unpublished observations). Second, cultures of Me-

lampus were initiated a few days before both new and

full moon phases, and the pattern of egg-laying was the

same in both cases: egg-laying began at the time of the

full moon but not at the time of the new moon. It would

be interesting to simultaneously compare egg-laying by

Melampus from Barn Island and Cape Cod under iden-

tical laboratory conditions.

How the timing of reproduction by these animals is

set is unknown. When Melampus spawns with a lunar

periodicity, each spawning cycle is about 2-3 times lon-

ger than when it spawns with a semilunar rhythm and

the period between spawning cycles is nearly doubled.

Evidently more eggs are produced per spawning cycle

and more time may be required to recover before com-

mencing another cycle of egg-laying.

Melampus is a largely terrestrial snail; its marsh hab-

itat may be unsubmerged for more than 95% of the time.

However, this snail possesses an aquatic veliger larva

(Russell-Hunter et al., 1972). Although egg-laying is usu-

ally initiated during spring tides both on the Little Sip-

pewisset and Barn Island marshes, flooding of the marsh

surface is not required for egg-laying to occur. On the

other hand, the hatching of veligers and their subsequent

settlement back on to the marsh depend upon inundation

of the marsh during spring tides (Russell-Hunter et al.,

1972). The observed differences in the patterns of egg-

laying by the two populations of Melampus do not ap-

pear to necessarily affect later stages in the life cycle of

the snail. For example, larval hatching may occur from

about 10 days to 24 or more days after egg mass depo-

sition, depending upon tidal submergence (Russell-Hun-

ter, 1972).

The minimal reproductive size of Melampus is from

about 5 mm shell length (Apley, 1970) to 5.8 mm shell

length (Balboni-Tashiro et al., 1985). However, not all

snails of this minimal size reproduce (Balboni-Tashiro et

al., 1985). Most of the Melampus collected from both

marsh regions at Barn Island were greater than 6 mm

in shell length.

In the present study, Melampus maintained in labo-

ratory culture produced an average of from 63 to 128

egg masses per animal during the reproductive season,

depending upon snail size. On the other hand, Russell-

Hunter e< a/., (1972) reported that under their laboratory

conditions, snails taken from the Little Sippewisset Marsh

produced on average only 39 egg masses during the

reproductive period. This is consistent with the obser-

vation that egg-laying by the Cape Cod population of

Melampus is typically restricted to fewer days compared

to the populations of this snail at Barn Island. On Cape

Cod, there are about 12 to 16 days of egg-laying by

Melampus each year (Ru.ssell-Hunter et al., 1972),

whereas at Barn Island snails deposit egg masses during

about 20 to 30 days within the reproductive period. How-

ever, the snails normally aggregate prior to mating and

egg-laying (Apley, 1970; Russell-Hunter et al., 1972);

and the number of egg masses produced may also be

influenced by snail density in laboratory cultures and

under field conditions.

Not only did the large snails from the restored im-

pounded marsh at Barn Island deposit more egg masses

than the smaller snails from the nearby bayfront marsh,

but the egg masses were significantly larger. In gastro-

pods for which size specific fecundity has been well doc-

umented, the larger egg capsules/ masses produced by

large individuals contain more eggs than the smaller egg

capsules/masses produced by small individuals (Hendler

& Franz, 1971; Spight & Emlen, 1976). In addition, a

preliminary study by Balboni-Tashiro et al. (1985) has

shown that older (larger) Melampus deposit more eggs

per mass than younger (smaller) snails. Therefore even

though egg numbers were not determined in the present

study, it seems highly probable that large Melampus

from the restored impounded marsh produced more eggs

than smaller ones from the bayfront marsh, indicating

that fecundity is size specific for this species. However,

interpretation of the data is complicated by the fact that

the different sized snails were taken from different

marshes. Although snails from the two marshes were

maintained on the same substratum (food resources) in

the laboratory, an influence of prior nutrition on repro-

duction beyond a larger size, resulting from a more rapid

growth rate, cannot be ruled out (Spight & Emlen, 1976).

Differences in growth rates of snails in the laboratory

on turf from the two marsh regions were significant.

Melampus from both marsh regions grew more rapidly

on Spartina alterniflora/iorh turf from the restored im-

pounded marsh than on Spartina patens turf from the

bayfront marsh. These results are consistent with the

hypothesis that the snails on the restored impounded

marsh grow more rapidly than the snails on the undis-

turbed bayfront marsh because of a larger/better food

resource. It would be interesting to determine which

potential foods are being used by Melampus on the two

marshes by examination of gut and fecal pellet contents

(Thompson, 1984). It is well documented that Melampus

prefers old detritus with a low phenolic content and

grows more rapidly on this type of diet (Rietsma et al.,

1988). However, other factors, such as the structural

characteristics (toughness and texture) of the food and/

or its moisture content, may also importantly affect pal-

atability. Finally, large microbial populations are often

found on detritus particles and they constitute a rich

energy source for organisms capable of removing them

(Odum & De La Cruz, 1967).

In conclusion, snails inhabiting a restored impounded

tidal marsh in southeastern Connecticut are larger, grow

more rapidly and produce more eggs than snails occu-

pying an undisturbed bayfront marsh within the same

marsh complex. The rapid growth and high fecundity

of Melampus on the restored impounded marsh indicate

that the marsh has become a highly favorable habitat

for this snail and provide additional evidence that res-

toration efforts have been successful.

J, A. Spelke et al., 1995

Page 47

ACKNOWLEDGMENTS

The authors are indebted to Dr. Robert Askins for help

with the statistical analysis and to Myron Peck for pro-

ducing figures 4 and 5. This study was supported by a

grant from the Andrew Mellon Foundation.

LITERATURE CITED

Apley, M. L. 1970. Field studies on life history, gonadal cycle

and reproductive periodicity in Melampus bidentatus

(Puimonata: Ellobiidae) Malacologia 10(2):381-397.

Baiboni-TashircJ.. P. WalbornandB. Crise. 1985. Diapause,

degrowth, and age of first reproduction in Melampus bi-

dentatus Biological Bulletin 169533

Crowe, T. A. and A. P. Covich. 1990. Predator-induced life-

history shifts in a freshwater snail. Science 247949-951.

Fell, P. E., N C Olmstead, E. Carlson, W Jacob, D Hitchcock

and G. Silber. 1982 Distribution and abundance of mar-

coinvertebrates on certain Connecticut tidal marshes, with

emphasis on dominant molluscs. Estuaries 5:234-239.

Fell, P. E , K A Murphy, M. A. Peck, and M. L. Recchia.

1991. Re-establishment of Melampus bidentatus (Say)

and other macroinvertebrates on a restored impounded

tidal marsh: comparison of populations above and below

the impoundment dike. Journal of Experimental Marine

Biology and Ecology 15233-48

Hausman, S. A 1932. A contribution to the ecology of the

salt marsh snail, Melampus bidentatus Say. The American

Naturalist 66:541-545.

Hendler, G. and D R. Franz. 1971 Population dynamics and

life history of Crepidula convexa Say (Gastropoda: Pros-

obranchia) in Delaware Bay. Biological Bulletin 141:514-

526.

Hilbish, T. J 1981. Latitudinal variation in freezing tolerance

of Melampus bidentatus (Say) (Gastropoda: Puimonata).

Journal of E.xperimental Marine Biology and Ecology 52:

283-297.

Holle, P. A. and C. F. Dineen. 1957. Life history of the salt-

marsh snail, Melampus bidentatus Say. The Nautilus 70:

90-95.

Joyce, A. A. and S. B. Weisberg. 1986. The effects of predation

by the mummichog, Fundulus heteroclitus (L), on the

abundance and distribution of the salt marsh snail, Me-

lampus bidentatus (Say). Journal of Experimental Marine

Biology and Ecology 100:295-306.

Odum, E P and A. A. De La Cruz 1967. Particulate organic

detritus in a Georgia salt marsh ecosystem. Estuaries 83:

383-388.

Peck, M. A., P. E. Fell, E. A. Allen, J. A. Gieg, C. R. Guthke,

and M. D. Newkirk 1994. Evaluation of tidal marsh

restoration: comparison of selected macroinvertebrate

populations on a restored impounded valley marsh and an

unimpounded valley marsh with the same salt marsh sys-

tem in Connecticut, USA, Environmental Management 18:

283-293.

Price, C. H. 1979 Physical factors and neurosecretion in the

control of reproduction in Melampus (Mollusca: Puimon-

ata). Journal of Experimental Zoology 207:269-282.

Price, C. H. 1980. Water relations and physiological ecology

of the salt marsh snail Melampus bidentatus . Journal of

Experimental Marine Biology and Ecology 45:51-67.

Rietsma, C S., I. Valiela, and A. Sylvester-Serianni 1982.

Food preferences of dominant salt marsh herbivores and

detritivores. Marine Ecology 3(2): 179-189.

Rietsma, C. S., I. Valiela, and R. Buchsbaum. 1988. Detrital

chemistry, growth, and food choice in the salt marsh snail

(Melampus bidentatus). Ecology 69:261-266.

Russell-Hunter, W D., M. L. Apley, and R. D. Hunter 1972.

Early life history of Melampus and the significance of

semilunar synchrony. Biological Bulletin 143:623-656.

Sinicrope, T L., P G Hine, R S. Warren, and W. A. Niering.

1990 Restoration of an impounded salt marsh in New

England. Estuaries 13(l):25-30.

Spight, T. M. and J. Emien. 1976. Clutch sizes of two marine

snails with a changing food supply. Ecology 57: 1 162-1 178.

Thompson, L. S. 1984. Comparison of the diets of the tidal

marsh snail, Melampus bidentatus and the amphipod,

Orchestia grillus. The Nautilus 98(l):44-53

Vince,S., I. Valiela, N. Backus and J. M. Teal. 1976. Predation

by the salt marsh killifish Fundulus heteroclitus (L.) in

relation to prey size and habitat structure: consequences

for prey distribution and abundance. Journal of Experi-

mental Marine Biology and Ecology 23:255-266.

Zar, J. H. 1974. Biostatistical analysis. Prentice-Hall, Engle-

wood Cliffs, New Jersey, 620 pp.

NOTICE THIS MATERIAL MAY BE

PROTECTED BY LAW

(TITLE 17 U.S. CODE)

THE NAUTILUS 108(2):48, 1995

Page 48

News and Notices

SMITHSONIAN FUNDS FOR MALACOLOGY

STUDENTS

The Division of Mollusks, Department of Invertebrate

Zoology, National Museum of Natural History, Smith-

sonian Institution announces the availability of the Houb-

rick-Rosewater Fellowship (up to $750) to be awarded

to graduate students of systematic malacology This award

provides support for students conducting systematic stud-

ies of Mollusca (leading to publication) who require ac-

cess to collections and libraries of the Division of Mol-

lusks, National Museum of Natural History. Funds can

be used for travel, subsistence, and research costs. Inter-

ested students should submit a succinct proposal (1-2

pages), including budget, with indication of any match-

ing funds, and a support letter from faculty advisor(s).

The application deadline is March 15, 1995. Award(s)

will be announced on April 15, 1995. Applications should

be sent to:

Dr. Clyde F. E. Roper

Division of Mollusks/NHB stop 118

National Museum of Natural History

Smithsonian Institution

Washington, DC 20560 USA

THE AMERICAN MALACOLOGICAL UNION

ANNUAL MEETING

The 1995 meeting of the American Malacological Union

will be held at the University of Hawaii at Hilo, in Hilo,

Hawaii, June 8-12, 1995 (arrive Hilo June 7 — leave June

13). The meeting is a celebration of islands, beginning

with a keynote address by Hawaii's well known naturalist

and photographer Bill Mull, who will describe with tales

and slides some of the wonders of Hawaiian animals and

plants, and especially land shells. Three symposia are

scheduled: biogeography convened by Gustav Paulay,

cephalopods convened by Richard Young, and conser-

vation convened by Elaine Hoagland. Plan to contribute

a paper and/or poster for other sessions. The annual

auction of books and shells will be exciting. Social events

include the President's Welcome, a sunset reception at

Lyman House Museum, a barbecue, and final banquet.

Field trips on June 1 1 include tidepools and snorkeling,

lava tubes, rainforest, kipukas (oases with snails on Mauna

Kea), and the volcano.

Accommodation will be in University of Hawaii at Hilo

dormitories on campus with a meal plan. Accommoda-

tion will run $25 per night per bed in a four room suite,

and the meal plan, covering most meals and coffee breaks

from arrival June 7 to leaving June 13 will be about $150.

If you prefer a hotel, you may make reservations from

a list with rates to be included with registration forms.

Van service to and from the airport will be provided.

Special day trips for accompanying guests and family

are scheduled.

Please indicate on a postcard, by FAX or e-mail if you

wish to receive a registration form and plan to attend:

E. Allison Kay

President, AMU

Department of Zoology

University of Hawaii

2538 The Mall

Honolulu, Hawaii 96822

Phone (808) 956-8620

FAX (808) 956-8612

E-Mail: [email protected]

SYMPOSIUM ON CONSERVATION AND

MANAGEMENT OF FRESHWATER MUSSELS

October 16-18, 1995. Symposium on The Conservation

and Management of Freshwater Mussels IL Initiatives

for the Future. Embassy Suites Hotel, St. Louis, Missouri.

Sessions devoted to endangered and at-risk species, re-

location and refugia, reproduction and propagation, sam-

pling methods, the mussel industry, and developing part-

nerships in preservation. For information, contact: Kurt

Welke, Wisconsin Department of Natural Resources, 111

West Dunn St., Prairie du Chien, WI 53821, USA. Phone

(608) 326-0233.

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THE NAUTILUS

Volume 108, Number 3

April 19, 1995

ISSN 0028-1344

A quarterly devoted

to malacology.

,7/

c Iruiiiution

Hoio, IbM 0<U>43

EDITOR-IN-CHIEF

Dr. M. G. Harasev\Tch

Division of Molliisks

National Museum of

Natural Histor\'

Smithsonian Institution

Washington, DC 20560

ASSOCIATE EDITOR

Dr. R. Tucker Abbott

.■\merican Malacologists, Inc.

P.O. Box 2255

Melbourne, FL 32902

CONSULTING EDITORS

Dr. Riidiger Bieler

Department of Invertebrates

Field Museum of

Natural History

Chicago, IL 60605

Dr. Robert T. Dillon, Jr.

Department of Biology

College of Charleston

Charleston, SC 29424

Dr William K. Emerson

Department of Living Invertebrates

The American Museum of Natural

History

NewYork, NY 10024

Dr Robert Hershler

Division of Mollusks

National Museum of

Natural History

Smithsonian Institution

Washington, DC 20560

Mr. Richard I. Johnson

Department of Mollusks

Museum of Comparative Zoology

Harvard University

Cambridge, MA 02138

Dr. Aurele I^a Rocque

Department of Geology

The Ohio State University

Columbus, OH 43210

Dr. James H. McLean

Department of Malacology

Los Angeles ("ounty Museum of

Natural History

900 Exposition Boulevard

Los Angeles, C A 90007

Dr. Arthur S. Merrill

% Department of Mollusks

Museum of Comparative Zoology

Harvard University

Cambridge, MA 02138

Dr. Paula M. Mikkelsen

Department of Malacology

Delaware Museum of Natural Histor\'

P.O. Bo.x 3937

Wilmington, DE 19807

Dr. Donald R. Moore

Division ol Marine Geology

and Geophysics

Rosenstiel School of Marine and

Atmospheric Science

University of Miami

4600 Rickenbacker Causeway

Miami, FL 33149

Dr. Ciustav Paulay

Marine Laboratory

University of Guam

Mangilao, Guam 96923

Mr. Richard E. Petit

RO. Box 30

North Myrtle Beach, SC 29582

Dr. Edward J. P(>tuch

Department of Geology

Florida Atlantic University

Boca Raton, FL 33431

Dr. David H. Stansbery

Museum of Zoology

The Ohio State University

Columbus, OH 43210

Dr. Ruth D. Tinner

Department of Mollusks

Museum of C^omparative Zoology

Harvard University

Cambridge. MA 02138

Dr. Geerat J. Vermeij

Department of Geology

University of C'alifornia at Davis

Davis, CA 95616

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TH E€:7NAUTI LUS

CONTENTS

Volume 108, Number 3

April 19, 1995

ISSN 0028-1344

Nathan S. Dayan

Robert T. Dillon. Jr.

Florida as a Biogeographic Boundary: Evidence from the

Population Genetics of Littorina irrorata

49

Mark E. Gordon

Venustaconcha sima (Lea), an Overlooked Freshwater

Mussel (Bivalvia: Unionoidea) from the Cumberland River

Basin of Central Tennessee

55

M. G. Harasewych

Alan R. kabat

Richard S. Houbrick (1937-1993): Biographical Sketch and

Malacologicai (contributions

61

Yu. A. Filippova

D. N. Khromov

K. N. Nesis

I. V. Nikitina

New Data on the Distribution and Morphology of some

Western Indian Ocean Sepiid Cuttlefishes (Cephalopoda:

Sepiida)

67

James H. McLean

Three Additional New Genera and two Replacement

Names for Northeastern Pacific Prosobranch Gastropods

80

News and Notices

WESTERN SOCIETY OF MALACOLOGISTS

ANNUAL MEETING

The twenty-eighth Annual Meeting of the Western So-

ciety of Malacologists will be held at the Resort at Chena

Hot Springs, near Fairbanks, Alaska, from June 2 to June

6, 1995. The agenda will include contributed papers on

all areas of molluscan studies, freshwater, marine, ter-

restrial, living and fossil. Symposia on ecology and pa-

leoecology are being organized \s ith the help of Howard

Fader and David Hopkins. Also in the planning stages

are an auction, reprint sale, and banquet. The University

of Alaska Museum Aquatic Collection will be available

for visitors before and after the meeting. For more in-

formation contact WSM President, Nora R. Foster, Uni-

versity of Alaska Museum, 907 Yukon Drive, Fairbanks,

Alaska 99775 USA. Phone: (907) 474-9557. E-Mail: FYA-

QUA(a) aurora.alaska.edu. Or contact Conference and

Special Events, 117 Eielson Building, University of Alas-

ka Fairbanks, Fairbanks, Alaska 99775.

INDO-PACIFIC MALACOLOGICAL MEETING

PRELIMINARY NOTICE

The Council of the Malacological Society of Australasia

has agreed to hold an international meeting on mollusks

of the Indo-Pacific region in Perth, Western Australia,

in January or February 1997. This notice is intended to

alert malacologists worldwide of the upcoming meeting

and to solicit suggestions for possible symposia; all sug-

gestions of possible topics are welcome. The meeting will

include all aspects of malacology, including mollusks of

marine, freshwater and terrestrial habitats. If sufficient

interest develops a post-conference field trip to the south-

west of Western Australia can be arranged. As plans

develop the conference will be advertised in malacolog-

ical journals. To be placed on the list for direct receipt

of future notices please write: Dr. F. E. Wells, Western

Australian Museum, Perth 6000. Western Australia (Fax:

61-9-328-8686).

NEW MOLLLISK JOURNAL LAUNCHED

The Malacological Society of Australia has a history of

over forty years of service to the study of mollusks in

Australia, and has published the Journal of the Mala-

cological Society of Australia since 1957. Like most sci-

entific societies, including AMSA, the Malacological So-

ciety has been continuously evolving. In recent years

there has been increased emphasis and membership of

the Society in New Zealand and countries to the north

of Australia. In recognition of its changing role the So-

ciety changed its name to the Malacological Society of

Australasia at the annual general meeting held in Sydney

in November 1993.

With the change in name and emphasis of the Society a

re-examination of the role oi the Journal of the Mala-

cological Societtj of Australia was appropriate. After

extensive di.scussion within the MSA Council it was de-

cided to broaden the content of the Journal and change

its name to Molluscan Research to reflect the modifi-

cations. The cover and pages have been redesigned. To

ensure continuity, the volume numbering remains the

same.

Theoretical papers considering any molluscan topic are

welcome for consideration for publication in Molluscan

Research. Papers considering specific geographical areas

or new taxa should be restricted to the Indo-West Pacific

region, which includes all of Australia. The Journal has

previously been an annual publication, but if sufficient

quality papers are available, Molluscan Research will

appear twice a year.

The first issue of Molluscan Research has been published

in early November, 1994.

For further information about Molluscan Research please

contact the editor. Dr. F. E. Wells, Western Australian

Museum, Perth, WA 6000, Australia. For information

afjout the Malacological Society of Australasia contact

the Department of Malacology, Australian Museum, P.O.

Box A285, Svdnev south, NSW 2000, Australia.

THE NAUTILUS 108(3):49-54, 1995

Page 49

Florida as a Biogeographic Boundary: Evidence from the

Population Genetics of Littorina irrorata^

[Nathan S. Dayan

Graduate Program in Marine Biology

Grice Marine Biological Laboratory

205 Ft. Johnson. Charleston, SC 29412

USA

Robert T. Dillon. Jr.-

Department of Biology

C;ollege of Charleston

C;harleston. SC 29424 USA

ABSTRACT

The marsh periwinkle, Littorina irrorata (Say, 1822), ranges

from New York to Texas with an apparently large disjunction

around southern Florida. We examined gene frequencies at

eight polymorphic enzyme loci in populations from Virginia,

South Carolina, Atlantic Florida, Gulf Florida, and Louisiana.

The within-population deviation from Hardv -Weinberg ex-

pectation was small at all loci examined (F|,,=0 049) Small but

significant gene frequency differences among populations were

detected at four loci, indicating some isolation by distance

Hierarchical gene diversity analysis suggested, however, that

very little of the population divergence present (Fy, =0.033) is

attributable to a division between coasts (Ft ;h=0.004). Nei ge-

netic distances calculated between pairs of sites sharing the

same coast were comparable to such distances between Atlantic

and Gulf pairs Thus we find no evidence that the F"lorida

peninsula constitutes a significant barrier to L irrorata, in spite

of the apparently extensive gap in its range. We offer three

hypotheses, not mutually exclusive: that the barrier is not real,

that it was broached in the not-too-distant past, or that bal-

ancing selection may be ongoing to hold polymorphisms con-

stant at multiple enzyme-encoding loci simultaneously.

Keij words: Gastropoda, electrophoresis, allozymes, range dis-

junction, balancing selection

INTRODUCTION

Shallow, protected regions along the southern Atlantic

coast of the United States and the northern coast of the

Gull of Mexico are characterized by temperate climate,

depositional environment, and the dominant salt marsh

cordgrass, Spartina. The molluscan faunas of these two

regions are described as "Carolinian (Rehder, 1954;

Coomans, 1962), sharing over 60% of their shallow water

gastropod species (query to database of Rosenberg, 1993).

However, striking climatic and geological variation along

' Contribution number 120 from the Grice Marine Biological

Laboratory

^ Address for Correspondence

the Florida peninsula seems to impose a biogeographic

boundary between these regions (Briggs, 1974). Due to

tropica! conditions, mangroves replace salt marshes in

protected bays and estuaries around 27°- 29° N latitude

(Kangas & Lugo, 1990). This transition occurs between

Cedar Key and Tampa Bay on the Gulf coast of Florida,

and between St. Augustine and Cape Canaveral on the

Atlantic coast. Coralline sands become the dominant sed-

iment type along south Florida coasts, replacing terrig-

enous silt and mud.

Scheltema (1989) surveyed the ranges of 88 mesogas-

tropod and neogastropod species, dividing the western

Atlantic coast into eight regions from Arctic Canada to

Brazil. He reported that 58 ol 72 species inhabiting his

"region IV ' (Beaufort, North Carolina to Miami, Flor-

ida) also occurred in his "region V" (Gulf of Mexico).

However, 18 of the 58 shared species did not occur in

Scheltema s "region VI", encompassing the southern tip

of Florida and the Greater Antilles. For at least these 18

species (including such common species as Fasciolaria

hunteria. Poliniccs duplicatus, and Littorina irrorata)

Florida would seem to constitute a potential barrier This

general distributional pattern extends beyond the near-

shore molluscan fauna to include many other elements

of the flora and fauna of the southeastern United States.

Recent molecular techniques have commonly detected

substantial genetic differentiation between animal pop-

ulations of the southern Atlantic coast and those of the

northern Gulf of Mexico. Most of the 19 such species for

which mtDNA surveys have been completed, including

horseshoe crabs, toadfish, black sea bass, diamondback

terrapins, and seaside sparrf)ws, show distinct differen-

tiation associated with the Florida peninsula (Avise, 1992).

To this list could be added the coastal North American

tiger beetle, Cicindela dorsalis, where mitochondrial

DNA sequence data were used to assign four subspecies

to either an Atlantic coastline lineage or a Gulf of Mexico

coastline lineage (Vogler & DeSaile, 1993). Significant

divergence at enzyme-encoding loci has been detected

between Atlantic and Gulf populations of such diverse

taxa as the sea anemone, Bunodosoma cavernata

(McCommas, 1982), and the marsh crab Sesarnia reti-

culatum (Felder & Staton, 1994).

Page 50

THE NAUTILUS, Vol. 108, No. 3

Figure 1. The range of Littorina irrorata (shaded), showing

sample sites for the present study.

No Florida disjunction is apparent in the range of the

American oyster, Crassostrea virginica, nor was any di-

vergence in allozyme frequencies detected by Buroker

(1983) in his survey of 19 populations from Massachusetts

to Texas. Thus the report of substantial mtDNA diver-

gence between Atlantic and Gulf oyster populations by

Reeb and Avise (1990) was greeted with unusual interest.

Karl and Avise s (1992) re-e.\amination of the issue using

restriction fragment length polymorphisms in anony-

mous single-copy nuclear genes confirmed the mtDNA

results, suggesting that Florida does, in fact, constitute a

barrier to the disper.sal of oysters. Karl and Avise went

on to propose that the similarity in allozyme frequencies

reported by liuroker might result from balancing selec-

tion at multiple enzyme-encoding loci. The importance

of natural selection to the preservation of enzyme poly-

morphism has been a central question in evolutionary

biology for over 25 years (Lewontin, 1991). Thus if the

findings of Karl and Avise can be generalized beyond

oysters, there will be implications for our understanding

of evolution as a whole.

The purpose of this study is to document divergence

in allozyme frequency between Atlantic and (iulf pop-

ulations of an heretofore genetically unsurveyed inter-

tidal mollusk, the marsh periwinkle Littorina (or Lit-

toraria) irrorata (Say, 1822). The snail is primarily an

inhabitant of salt marshes dominated by the cord grass

Spartina altcrniflora, leaving the marsh surface with the

incoming tide to climb the vegetation. It ranges from

Jamaica Bay, Long Island, New York (Jacobson, 1965)

to Port Isabel, Texas (Bequaert, 1943), with a disjunction

around southern Florida (Figure 1). As such it would

seem an excellent candidate for an attempt to confirm

the Karl and Avise phenomenon.

Littorina has been the object of considerable popu-

lation genetics research worldwide, with much effort

directed toward questions of systematics (Ward & War-

wick, 1980; Maestro et al . 1991; Boulding el ai, 1993;

Zaslavskaya et al., 1992). Other workers have prospected

for environmental dines (Newkirk & Doyle, 1979;Janson

& Ward, 1984; Johannesson et al., 1993), founder effects

(Janson, 1987), or correlates of heterozygosity (Noy et

al., 1987; Foltz et al , 1993). However, the only previous

examination of L. irrorata allozymes, prior to this report,

was that of Berliner (1981) in a Virginia salt marsh. He

found no significant difference in heterozygosity between

young and old size classes, but noted a lower value of

heterozygosity in snails of median age. Berliner's study

did not extend beyond his single population.

MATERIALS AND METHODS

Approximately 50-60 L. irrorata per site were collected

from three sites on the Atlantic coast of the southeast

United States and two from the Gulf of Mexico: the

Virginia Institute of Marine Science Laboratory (VIMS)

at Wachapreague, Virginia, the Grice Marine Biological

Laboratory (GMBL) at Charleston, South Carolina, Cres-

cent Beach, Florida (CBFL), Tampa Ba\ , Florida (TBAY),

and the Louisiana University Marine Consortium Lab-

oratory (LUMC) at Cocodrie, Louisiana (Figure 1) All

sites except TBAY were typical salt marshes dominated

by the salt marsh cord grass, S. alterniflora. The Tampa

Bay site was a small patchy area of S. alterniflora growing

on sand, rather than mud. At all sites, the snails were

found on the stalks of the salt marsh cord grass or on the

substrate at the base of the stalks, generalK in the mid-

marsli to high marsh areas. The snails were transported

to Charleston alive, where tissues were frozen at — 60°C

in 150-300^1 of 0.05 M Tris Tissue buffer pH 7.5 (Dayan,

1994).

Horizontal protein electrophoresis was conducted us-

ing methods and equipment previously described (Dil-

lon, 1985; 1992; Dayan, 1994). The 12% starch gels were

a 1:1 mixture (by volume) of Electrostarch (Otto Hillar,

Madi.son, Wl) and Sigma starch (Sigma C'hemical, St.

Louis, MO). We initially compared the zymograms of

individuals from GMBL and LUMC using 19 enzyme

stains and 9 buffer systems. Results were poor or unin-

terpretable for six enzyme-s and appeared invariable for

seven others. LUtimately we were able to resolve allo-

zymes interpretable as the products of codominant Men-

delian alleles at eight loci (encoding six enzymes) using

N. S. Dayan and R. T. Dillon, Jr., 1995

Page 51

Table 1. Allele frequencies at eight enzyme-encoding loci in

five populations of Liltorina irrorata. (n) = Sample size

Locus

Estl

(n)

Est2

(n)

Pgm

(n)

Mpi

(n)

Isdhl

(n)

Isdh2

(n)

Sdh

(n)

Lap

All-

ele VIMS

Population

GMBL CBFL TBAY LUMC

A

B

C

A

B

A

B

C

D

A

B

A

B

C

0.571

0 429

0.000

56

0,984

0.016

61

0.648

0.270

0.082

0.000

61

0 992

0 008

63

0 934

0.057

0.008

61

A 0.992

B 0.008

60

A

B

C

A

B

C

D

(n)

0814

0.161

0.025

59

0917

0.075

0.008

0 000

60

0.689

0.292

0.019

53

0,989

0.011

44

0 441

0.480

0.078

0.000

51

0.952

0.048

83

0.965

0.035

0.000

57

0.990

0.010

50

0.788

0.205

0.008

66

0.700

0.255

0.036

0.009

55

0.638

0.362

0.000

58

0 992

0,008

62

0 490

0.288

0.202

0.019

52

0.992

0.008

62

0.877

0.098

0.025

61

1 000

0.000

62

0.856

0.136

0.008

59

0 900

0 067

0 025

0.008

60

0 556

0 444

0.000

62

0.968

0.032

62

0.411

0.218

0.371

0.000

62

0.968

0.032

62

0 966

0.017

0017

59

1 000

0.000

62

0.805

0.161

0.034

59

0 860

0 061

0.018

0.507

0,485

0.007

67

1 000

0.000

63

0 490

0.127

0 343

0 039

51

0.971

0.029

70

0.918

0 061

0.020

49

0 990

0010

49

0.698

0.250

0.052

48

0.862

0.078

0.043

0.017

58

three buffer systems. The AP6 buffer (Clayton & Tretiak,

1972) was used to resolve esterases {Est, two loci), and

phosphoglucomutase (Pgrii). The VVWI (Ward & War-

wick, 1980) buffer was also used to resolve Pgm, as well

as isocitrate dehydrogenase {Isdh, two loci), and sorbitol

dehydrogenase (Sdh). The TC6 buffer (Dillon, 1985) was

used to resolve leucine aminopeptidase (Lap) and man-

nose-phosphate isomerase (Mpi). Working with L. sax-

atilis. Ward et al. (1986; 1991) have confirmed Men-

delian inheritance at all these loci except Est and Sdh

Data analysis was by Biosys-1 (Release 1.7, Swofford

& Selander, 1981) unless otherwise specified. We tested

the fits to Hardy- Weinberg expectation for each locus at

each population using goodness of fit x" statistics, com-

bining rare genotypic classes as necessary. We then per-

formed two separate gene diversity analyses using

Wrights (1978) F-statistics. In the more conventional

analysis, the total deviation from Hardy- Weinberg ex-

pectation over all loci (Fi^) was divided into a component

Table 2. Wright's (1978) F-statistics, averaged over eight loci,

measuring gene diversity attributable to individuals (/), pop-

ulations (S ), and coast (C) Values from the hierarchical analysis

of coastal variance are set under Fsj, the variance attributable

to population structure

C^omparison

Coefficient

F,s

FsT

Fsc

FcB

>^sn

Fn

0.049

0.033

0.020

0.004

0.024

0.080

within populations (F,^) and a component between the

five populations (Fsy). We also performed a hierarchical

analysis (Dillon & Manzi, 1992; Dillon & Wethington,

1995), grouping the three Atlantic and two Gulf popu-

lations to determine the proportion of gene diversit\'

attributable to coast. Mean F-statistics calculated in this

way we labeled F^^ (between populations within coasts),

F( B (between coasts) and Fsb Note that Fjb is expected

to be less than F^-,. since that proportion of the variance

between populations between coasts remains unattribut-

ed in the hierarchical analysis.

For each locus at which n alleles were identified, di-

vergence among populations was tested with a 5 x n x"

contingency test. To avoid the necessity of combining or

eliminating rare alleles for this analysis, we estimated

the significance of our values of x" using the Monte Carlo

approach of Roff and Bentzen (1989). Unbiased genetic

identity and distance between all pairs of populations

was calculated using the method of Nei (1978).

RESULTS

Gene frequencies at eight enzyme-encoding loci from

five populations of Littorina irrorata are shown in Table

1. The fits to Hardy- Weinberg expectation within pop-

ulations were very close in most cases. Goodness-of-fit

values of x' nominally significant at the 0.05 level were

obtained only at Pgm in VIMS, and at Isdh in TBAY,

well within expectation for type I error. The mean value

of F|s over all loci, measuring deviation from Hardy-

Weinberg within-sites over the entire studv, was small

(F,s=0.049, Table 2).

The mean value of F^t = 0.033, measuring deviation

from Hardy- Weinberg expectation between sites, was

lower than the deviation within-sites. However, the data

of Table 1 reflect significant divergence among the five

populations at four loci. The value of x' testing homo-

geneity in Pgm allele frequencies was 103.6 (p<0.001),

with notably high frequencies of Pgm-A in VIMS, Pgm-

B in GMBL, and Pgm-C in the three most southern

populations. The GMBL population was distinguished

by a significantly high frequency of Lap-B (overall

X^=46.4, p<0.001), the LUMC population by high Sdh-

B (overall x^=21.7, p=0.034), and the CBFL population

Page 52

THE NAUTILUS, Vol. 108, No. 3

Table 3. Neis (1978) unbiased genetic identity (above diag-

onal) and distance (below diagonal) between all pairs of five

L. irrorata populations.

by high Isdhl-B (overall x-=19.6, p=0.047). Differences

at the remaining four loci were not significant.

The contribution of the coastal level to the hierarchical

gene diversity analysis was negligible. Table 2 shows that

F^;i5, the deviation from Hardy-Weinberg expectation

between coasts, was 0.004. This was lower than the vari-

ance between populations within coasts. The values of

Neis unbiased similarity and distance among all pairs

of sites are shown in Table 3. The four within-coast values

of Neis distance ranged from 0.001 to 0.014, only slightly

less than the range for the six values between coasts (0.004

to 0.026).

DISCUSSION

The data of Table 1 constitute substantial evidence of

low-level genetic divergence among populations of Lit-

torina irrorata separated by distances on the order of

hundreds of kilometers. A general relationship between

interpopulation divergence, mode of embryonic devel-

opment, and dispersal capability has often been noted

in marine mollusks (Burton & Feldman, 1981; Scheltema,

1989; Yamada, 1989). Littorina irrorata spawn at high

tide, females releasing eggs just beneath the water level

(Gallagher & Reid, 1974). Egg capsules are planktonic,

but slightly negatively buoyant in calm seawater (Bing-

ham, 1972). They hatch into swimming veliger larvae

after one to two days of further development, but the

time to settlement is unknown. Settlement occurs on

Spartina shoots (Boothe, 1969), where juveniles remain

hidden in curled blades until they reach about 5 mm

(Stiven & Hunter, 1976; Crist & Banta, 1983). Berger

(1973) compared population divergence in three North

American Littorina species, L. littorea (pelagic larval

development), L. ohtusata (juveniles hatch from gelat-

inous egg masses), and L. saxatilis (ovoviviparous). The

levels of genetic differentiation we report here in L.

irrorata are, as might be expected, comparable to those

of L. littorea and much less than those of L. ohtusata

or L. saxatilis.

Such interpopulation genetic divergence as we have

identified does not, however, seem to reflect a barrier to

dispersal around the Florida peninsula. Table 1 shows

no allele uni(|uc to cither coast; even the live rarest alleles

{Est2-R. !sdh2-B, Lap-C, Lap-D. and Mpi-R) were found

in both Atlantic and Gull populations. Nor did our hi-

erarchical gene diversity analysis or our inspection of

interpopulation genetic distances suggest any evidence

of a barrier to gene flow corresponding to the Florida

peninsula. The divergence between Atlantic and Gulf

populations of L. irrorata is indistinguishable from dif-

ferences among populations sharing the same coast. We

offer three (not mutually-exclusive) hypotheses for this

unexpected result: no-barrier, past-dispersal, and bal-

ancing selection.

It is possible that, in spite of the climate shift in south-

ern Florida and the disappearance of salt marsh habitat,

no barrier currently exists between Atlantic and Gulf

populations of L. irrorata. We are unaware of any col-

lections of adult snails south of Ft. Pierce, Florida. But

it is possible that sparse populations do exist on man-

groves or concrete bulkheads along the remainder of the

southern Florida coast. Long distance dispersal of larvae

or juveniles (perhaps on dead Spartina rack) is also pos-

sible. Only a few days might be required to transport

larvae on the Gulf Stream from the west coast of Florida

into the eastern Atlantic, although onshore currents would

still be necessary to carry larvae back into estuaries. It

is difficult, however, to imagine how such a passive dis-

persal mechanism might be effective for Littorina larvae

and not for toadfish, black sea bass, horseshoe crabs, sea

anemones, or oyster larvae, all of which do exhibit ge-

netic divergence between coasts.

A second possible explanation for the absence of di-

vergence between Atlantic and Gulf L. littorina popu-

lations would invoke higher levels of gene flow in the

not-too-distant past. The Suwannee Straits (or Okeefen-

okee Trough) most recently connected the two coasts of

northern Florida during the Pliocene epoch (Avise, 1992).

Fossil L. irrorata found in North Carolina, South Car-

olina, and Florida have been dated from the upper Mio-

cene and Pliocene, while fossils found in Louisiana and

Texas are Pleistocene in age (Bequaert, 1943). Faunal

exchange between Atlantic and Gulf may also have oc-

curred in the Pleistocene epoch, although it is difficult

to predict under what environmental conditions. L ir-

rorata could certainly have extended its range southward

during glacial periods, but lowering sea levels would have

elongated the Florida peninsula. The distance between

Atlantic and Gulf shortened as interglacial seas rose, but

the snail may have been driven north by the advance of

tropical conditions. It would appear, however, that the

past-dispersal hypothesis is no better than the no-barrier

hypothesis in accounting for the difference betv\een Lit-

torina and those (many) s[)ecies that do show intercoastal

divergence.

The third hypothesis would recall Karl and Avise's

(1992) work with oysters. Although allozyme frequencies

were homogeneous, Karl ami .\\ ise were able to establish

that a barrier has existed between Atlantic and Gulf

oyster populations using nilDNA and anonymous nuclear

DNA markers. They concluded that balancing selection

may be holding Atlantic and Gulf cnster populations

undifferentiated in their enzyme poK niorphism in the

absence of gene flow, and that caution should be used

N. S. Dayan and R T Dillon, Jr . 1995

Page 53

regarding the assumption of neutrality for ailozyme

markers.

The present data are insufficient to distinguish among

these three hypotheses. Further, more detailed surveys

of L. irrorata's range in south Florida would be helpful,

along with an expansion of isozyme studies if necessary.

Some examination of the larval behavior and develop-

ment time of L. irrorata would clarifv the likelihood of

long distance gene Dow. And further surveys of genetic

divergence among Atlantic and Gulf samples of L. ir-

rorata using mtDNA or nuclear DN A markers are strong-

ly indicated.

ACKNOWLEDGMENTS

We thank E. W. McMillan, K. Metzner-Roop, and A. R

Wethington for assistance in the laboratory, and C. Choi-

let and C. Walton for help in the field. Drafts of this

manuscript have been read to good effect by C. K. Biern-

baum, R. C. Brusca, R. A. McCarthy, and J. W. Smiley

Gary Rosenberg provided insights on the distribution of

mollusks, R. W. Chapman wrote the program for the

Roff-Bentzen analysis, and Dr. & Mrs. M. Dayan pro-

vided moral support. This work is based on a thesis sub-

mitted in partial fulfillment of the requirements for an

M.S. in Marine Biology from the University of Charles-

ton, SC.

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Page 55

Ventistaconcha sima (Lea), an Overlooked Freshwater Mussel

(Bivalvia: Unionoidea) from the Cumberland River Basin of

Central Tennessee

Mark E. Gordon

Zoology Section

Campus Box 315

University of Colorado Museum

Boulder, CO 80309 USA

ABSTRACT

Unio simus Lea, 1838, generally has been considered to be a

junior synonym of Villosa nebulosa (Cxinrad, 1834) Exami-

nation of recently collected specimens indicates that it is a valid

species of Venustaconcha restricted to the upper Caney Fork

River system in central Tennessee. Venuslacuncha sima may

represent the Cumberlandian counterpart of V. pleasii (Marsh,

1891) of the southern Ozark Plateaus.

Key words: Bivalvia, Unionoidea, Venustaconcha.

INTRODUCTION

During recent studies of the mussel fauna of the Caney

Fork River basin (e.g., Farzaad, 1991; Layzer et ai,

1993), a major tributary system of the Cumberland River,

a diminutive purplish-nacred mussel was found, which

possessed a posterioventral emargination with an asso-

ciated radial sulcus in female shells and relatively heavy

hinge dentition. Stansbery (personal communication)

considered it to be an undescribed species of Villosa.

however, the above morphological characters match di-

agnostic criteria in Haas (1969) for Venustaconcha. A

review of the multitudinous species descriptions of Isaac

Lea and subsequent examination of type specimens in

the National Museum of Natural History, Smithsonian

Institution, resulted in the identification of this mussel

as Venustaconcha sima (Lea, 1838).

SYSTEMATICS

Tribe Lampsilini

Venustaconcha sima (Lea, 1838)

(Figures 1-7; Table 1)

Margarita (llnio) simus Lea (1836:29) nomen nudum.

Unio simus Lea (1838:26, figure 20 on plate V'llI).

Margaron (Unio) simus Lea (1852:31).

Lampsilis simus Simpson (1900:556).

Lampsilis sima Simpson (1914:123).

Eurynia (Micromya) nebulosa (in part) Ortmann (1918:577).

Micromija nclndosa (in part) Ortmann (1924:102).

Lampsilis (Liguniia) nebulosa (in part) Frierson (1927:78).

Villosa nebidosa (in part) Burch (1975:173).

Description: Shell small, broadly elliptical (males) to

somewhat ovate (females), barely inflated (W/HsO.63;

for interpretation of shell proportions, see Wu, 1978);

solid, thinner posteriorly; anterior rounded; dorsal mar-

gin slightly convex, oblique, consequently shell may ap-

pear humped or somewhat winged, juncture with an-

terior margin sometimes angular; ventral margin slightly

convex to virtually straight; posterior bluntly pointed to

biangulate; posterio-ventral region of female shells may

be swollen with some distention of the extreme posterior

of the ventral margin, an emargination and associated

radial sulcus may develop posteriorly to the distention;

posterior ridge low, vaguely double, somewhat flattened

between ridges but may appear rounded; posterior slope

slightly concave with a shallow radial furrow, furrow

may cause an indentation of the posterio-dorsal margin;

ligament low, short; umbo compressed, low, barely ele-

vated above dorsal margin; umbonal sculpture ". . ir-

regular, somewhat doubly-looped ridges. . ." (Simpson,

1914); periostracum rather smooth and somewhat shiny

in younger specimens, may be dull and rough in older

shells, annual growth lines well-marked, yellowish to dark

brown or black with narrow dark green rays, raying most

prevalent on posterior half of shell and may be wavy

and clustered closely together.

Pseudocardinals thick, heavy, serrated, double in left

valve, single in right valve but often with small anterior

and posterior denticles opposite adjacent sulci; interden-

tum moderately wide, relatively short; lateral teeth short,

straight, lamellar, may be slightly serrate, double in left

valve, single in right valve; anterior adductor and re-

tractor muscle scars confluent, rather small, deeply im-

pressed; protractor muscle scar distinct, straight to cres-

cent-shaped; posterior muscle scars typically confluent,

impressed, adductor large, retractor small and positioned

under distal base of lateral tooth; dorsal muscle scars

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THE NAUTILUS, Vol. 108, No. 3

Figures 1-11. Shells of Venustacoiicha. 1-5. Typf sptvuiR-ii!, ul \ cntislaconcha sinm 1. Hnidtype (male, I'SNM 85342)

Mai.' paratypes (ANSP .56474). 5. Female paratype (ANSP 56474). 6-7. Venustaconcha sima from C:ollins River, Mt. Olive, Grundy

County, Tennessee, collected 15 Julv 1989. 6. Male. 7. Female. 8-9. Venustaconcha ellipsiformis from Osage Fork of Gasconade

Hivcr, Dryknoh, Laclede Countv. Missouri, collected 3 October 1983. 8. Male. 9. Female 10-11. Venustaconcha pleasii from

James River, east of Springfield, Greene County, Missouri, collected 4 October 1983 10. Male 11. Female Scale bars = 1 cm.

M. E. Gordon, 1995

Page 57

Table 1. Shell dimensions of type specimens and representative shells of Venustaconcha sima (L = length, H = height, W =

width, M = male, F = female; specimens other than types were collected from the Collins River, Tennessee, Grundy County, Mt.

Olive, 15 July, 1989).

Specimen

Se\

H

W

H L

\V/H

Holotype

Paratype

Collins River

Paratype

Collins River

deeply impressed on underside of interdentum, occa-

sionally causing a notch in interdentum, extend from just

posterior of the umbo to base of pseudocardinals; pallial

line impressed, lighter posteriorly; beak cavity moder-

ately developed; nacre variable, tends to be purple in

hving individuals but may be lighter, pinkish or whitish,

and blotched with brow n, color fades rapidly in dead

shells.

Type locality: "Cumberland River, Tennessee . . . This

shell was procured by Professor Troost from the Cum-

berland River, but whether near Nashville or not, I am

not informed (Lea, 1838).

Type specimens: Holotype, National Museum of Nat-

ural History, Smithsonian Institution (USNM) 85342

(male). Paratypes, Academy of Natural Sciences of Phil-

adelphia (ANSP) 56474 (three males, one female). All

type specimens have whitish nacre as noted by Lea

(1838)', evidently reflecting the tendency for the purple

coloration to fade in dead shells. Only one additional lot

of this species was located at the USNM (782358) labelled

"Villosa trahalis perpurpurea (Lea, 1861) ', but contain-

' Lea (1862:62) stated that V. sima had a salmon colored nacre.

Since examination of available specimens at LISNM and ANSP

indicated that Lea only had access to specimens in the type

lots (all with white nacre), he may have confused this particular

attribute of the similarly-shaped Unio noiatus Lea, 1838 (in-

certae sedis) with .sima.

ing a mixed collection of Venustaconcha sima and Pleu-

robema gibberum (Lea, 1838) from the Collins River,

Grundy County, Tennessee. Additional lots are located

at the Carnegie Museum, Museum of Comparative Zo-

ology, and Ohio State University Museum. Voucher spec-

imens from the present study have been deposited in the

USNM, University of Michigan Museum of Zoology, Mu-

seum of Comparative Zoology, Carnegie Museum, and

University of Colorado Museum.

Distribution: The distribution of this species is difficult

to delineate due to the lack of historical records; however,

recent collecting indicates that it is restricted to the Ca-

ney Fork River basin above Great Falls. Much of its

presumed former habitat is inundated by Great Falls

Reservoir.

Habitat: Venustaconcha sima appears to be associated

with riffle systems in small to medium-sized streams,

including headwaters. It is most abundant in sandy sub-

strate between cobbles and boulders with slow to mod-

erate current, although it also occurs in courser substrate

and faster currents (see Farzaad, 1991).

Etymology: Venustaconcha- L. venusta [adj.] -I- con-

cha [n., feminine]), pretty shell; sima-L. [adj.], snub-

nosed.

DISCUSSION

Following its original description, Venustaconcha sima

was known only by its inclusion in synoptic lists of fresh-

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THE NAUTILUS, Vol. 108, No. 3

Figure 12. Distribution of Venustacuncha sima within the

Caney Fork River system, Tennessee.

water mussels (e.g., Troschel, 1839; Conrad, 1853; Lea,

1870). Call (1885) alluded to an affinity with -Xhuo iris.

Lea," 1829, which Simpson (1900, 1914) subsequently

followed. Simpson (1914) did note its resemblance to

"Lampsilis" nelmlosa (Conrad, 1834) and "L. ' ellipsi-

formis (Conrad, 1836). Ortmann (1918) relegated sima

as a junior subjective synonym inider "Eunjnia {Micro-

mya)' nebidosa. Veimstaconcha sima is a heavier shell

than Villosa iris or V. nebidosa. It tends to be relatively

shorter and broader than either of the latter and, par-

ticularly in the case of females, more closely resembles

Venustaconrha ellipsiformis by virtue of its posterior-

ventral swelling with associated emargination and radial

sulcus, heavy p.seudocardinal teeth, wide interdentum,

and fine, wavy, closely-spaced rays (Figs. 8-9). These

characteristics were among those employed by Haas

(1969) to distinguish Venttstaamcha Frierson, 1927-, from

Villosa Frierson, 1927.

Within the Cumberland River basin, Venustaconcha

sima is similar in appearance to and occasionally may

be confused with species in the Villosa iris / nebiilosa

comple.x (see above) and V. trabalis (Conrad, 1834) (e.g.,

(USNM 782358). Compared to Venustaconcha sima, the

latter species is a relatively heavier, larger and more

elongate shell. Its periostracum tends to be darker, the

posterior-ventral emargination of the female is not as

acute, pseudocardinal teeth are relatively larger, and the

nacre tends not to be blotched. These two species also

appear to be distributed allopatrically. Interestingly, the

shells of V. trabalis and V. ellipsiformis are more similar

to each other than either is to V. sima, and Frierson

(1927) included trabalis in his original list of species

under Venustaconcha. Although further investigation

into generic relationships of advanced lampsilines is re-

quired (Hoeh and Frazer, personal communications),

classification of trabalis under Venustaconcha may be

more representative of actual relationships that an as-

sociation with Villosa.

The distribution of Venustaconcha sima appears to

be restricted to the portion of the Caney Fork River

basin draining the Highland Rim upstream from Great

Falls, the cataract at the escarpment between the High-

land Rim and Nashville Basin (Fig. 12). Call (1885) listed

its range as the "Cumberland river, Tennessee, and

Swamp Creek, Whitfield County, Georgia." To this,

Simpson (1900, 1914) added '. . .Tennessee river sys-

tem(s); Othcalooga Creek, northwest Georgia." As noted

above, only two lots of V. sima were observed at the

USNM. It is unclear upon which specimens Simpson

based this range. Call (1885) and Simpson (1900, 1914)

possibly may have confused V. sima with species of Vil-

losa (e.g., V. iris complex; V. vaniixemii [Lea, 1838]',

particularly the umbrans Lea, 1857, morph).

Lea (1838) noted the Cumberland River, Tennessee,

as the type locality for U. simus, but commented that

he did not know the exact collection site. Lea's locality

information often was increadibly vague (e.g., Unio

grayanus Lea, 1834: type locality - China) or reflected

the address of the collector (e.g., Lampsilis reeveiana

Lea, 1852: see Gordon & Kraemer, 1984). The latter may

be the case with Venustaconcha sima. Since the type

specimens were sent by Prof. Troost from Nashville, Lea

may have assumed that they had been collected from

the Cumberland River. With respect to its known dis-

tribution (Fig. 12), the type specimens may have origi-

nated from the Collins River or possibly its principle

- On several occasions (e.g., Vokes, 1980; Oesch, 1984; Stan.s-

bery, xeroxed ephemera), authorship of Venustaconrha has

been attributed to Thieie, 1934 or 1935. Venustaconcha was

proposed as a replacement name by E'>ier.son (1927) in the

"errata el corrigenda" for Vcnusta Frierson, 1927 (nan Boett-

ger, 1877, nee Barrande, 1881).

^ Lea (1838) staled that he named this species after Prof

Lardner Vanuxem, yet the original spelling ended with the

suffix for a locality name rather than the masculine genetive.

The spelling of vanuxeniensis was corrected to vanuxemii by

Lea (1858). This change is in accordance with Articles 19a (i),

32c (ii), 32d, ;33b (ii) and .Appendix D of the International Code

of Zoological Nomenclature (thirtl edition) John.soii (1974) pre-

viously noted this correction of the lapstis calami.

M. E. Gordon, 1995

Page 59

tributary. Barren Fork, in the vicinity of McMinnville,

Warren Count>, Tennessee (the largest town in that area

in 1838).

Vcniistaconcha sima does not appear to have a coun-

terpart in tlie Tennessee River or drainages northward

into Kentucky. As previously observed, Simpson (1914)

noted similarities between V. sima and V. ellipsijormis.

However, in both shell morphology and habitat prefer-

ence, V. sima seems to be allied more closely to V. plcasii

(Marsh, 1891), a small mussel endemic to the Ozark

Plateau drainages of the White River system, Arkansas

and Missouri (Figs. 10-11) (personal observations; Gor-

don, 1980). This hypothetical association would be con-

sistent with previously observed molluscan affinities be-

tween the fauna of the Cumberlandian and Interior

Highlands regions (e.g., Ortmann, 1917; van der Schalie

& van der Schalie, 1950).

.\lthough major surveys of the Cumberland River have

been conducted (Wilson & Clark, 1914; Neel & Allen,

1964), its mussel fauna has received relatively little at-

tention in comparison with that of the Tennessee River

system. Starnes and Bogan (1988) listed 85 species from

the CAuriberland River drainage, while Gordon and Lav -

zer (1989) reported 94 species. Previous faunal compi-

lations for the Caney Fork included 27 species (Miller,

1984) and 14 species (Starnes & Bogan , 1988); however,

Layzer et al. (1993) have found that the historical fauna

was considerably more diverse than the former accounts

indicate. It is apparent that the species richness of the

Cumberland River mussel fauna is greater than previ-

ously considered,

Ortmann (1924) concluded that the Cumberland River

system lacked an endemic mussel fauna. Starnes and

Bogan (1988) similarly stated that "all of the mussel

species recorded from the Cumberland River occur in

the Tennessee River system despite the inclusion in their

synoptic table of two mussels (Alasmidonta atropiirpii-

rea [Rafinesque, 1831]^ and Pleurobema gibberum) that

are restricted to portions of the Cumberland River drain-

age. A distinct endemic mussel fauna did evolve in the

upper CAmiberland River system. Despite massive hab-

itat destruction within the basin (e.g., reservoir construc-

tion, acid coal mine run-off), a fragment of this fauna

persists, as represented by A. atropnrpttrea, Quadrula

tiiberosa (Lea, 1840: possilby e.xtinct), P. gibberum. and

Vcniistaconcha sima.

■* Clarke (1981) reported A. atropurpurea from the Collins

River at a site within the Highland Rim province. This speci-

men appears to have been a misidentified shell of A. marginata

Say, 1818 (Anderson, personal communication) Alasmidonta

atropurpurea is restricted to the Cumberland River drainage

on the Cumberland Plateau upstream from the hypothesized

pre-erosional localilv of Cumberland Falls (Gordon & Lavzer,

1993).

ACKNOWLEDGMENTS

Partial funding for this study was provided by the Na-

tional Ecology Research Center, U.S. Fish and Wildlife

Service, Fort Collins, Colorado. I would like to thank

Andrew G. Gerberich, Paul R. Greenhall, and Robert

Hershler for their assistance and for access to the collec-

tions at the National Museum of Natural History. Robert

M. Anderson, Indiana Department of Natural Resources;

Kenneth S. Frazer, Bar Harbor, Maine; Walter R. Hoeh,

Dalhousie l^niversity; and David H. Stansbery , Ohio

State University Museum, provided pertinent informa-

tion.

LITERATURE CITED

Burch, J. B. 1975. Freshwater unionacean clams (Mollusca:

Pelecypoda) of North America. Malacological Publica-

tions, Hamburg, Michigan. 204 pp.

Call, R E. 1885. A geographic catalogue of the Unionidae

of the Mississippi Valley Bulletin of the Des Moines Acad-

emy of Science 1:5-57.

Clarke, A H 1981 The tribe Alasmidontini (Unionidae: An-

odontinae), part 1: Pegias. Alasmidonta, and Arcidens.

Smithsonian Contributions to Zoology 326:1-101.

Conrad, T. A. 1853. A synopsis of the family of naiades of

North America, with notes, and a table of some of the

genera of the family, according to their geographical dis-

tribution, and descriptions of genera and subgenera of the

familv. Proceedings of the Academy of Natural Sciences

of Philadelphia 6:243-269.

Farzaad, M 1991. Microhabitat utilization of Venustaconcha

sima (Bivalvia; Unionidae) and its host, Cottus carolinae,

in the Collins River, Grundy County, Tennessee MS the-

sis, Tennessee Technological LIniversity, Cookeville. 94 pp.

Frierson, L. S. 1927. A classified and annotated check list of

the North American naiades. Baylor University Press, Waco,

Texas 111 pp

Gordon, M E 1980 Freshwater Mollusca of the Elk River,

White River above Beaver Reservoir, and Frog Bayou

drainages of the southwestern Ozarks. M.S. thesis. Uni-

versity of Arkansas, Fayetteville. 366 pp.

Gordon, M. E. and L. R. Kraemer. 1984. Lanipsilis reeveiana

and Lampsilis strecheri (Bivalvia: Unionacea): some clar-

ifications. Malacological Review 17:99-100.

Gordon, M. E. and J. B. Layzer. 1989. Mussels (Bivalvia:

LInionoidea) of the CAimberland River: review of life his-

tories and ecological relationships. U.S. Fish and Wildlife

Service Biological Report 89(15):l-99.

Gordon, ME. and J. B. Layzer 1993 Glochidial host of

Alasmidonta atropurpurea (Bivalvia: Unionoidea: Union-

idae). Transactions of the American Microscopical Society

112:145-1.50.

Haas, F. 1969. Das Tierreich, Lieferung 88: Superfamilia

Unionacea. Walter de Gruyer & Co., Berlin. 663 pp

Johnson, R. I. 1974. Lea's unionid types or recent and fossil

taxa of Unionacea and Mutelacea introduced by Isaac Lea

including the location of all the extant types. Special Oc-

casional Publication of the Department of Mollusks, Mu-

seum of Comparative Zoology 2:1-159

Layzer, J B , M. E. Gordon, and R M Anderson. 1993. Mus-

sels: the forgotten fauna of regulated rivers Regulated

Rivers: Research and Management 8:63-71.

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THE NAUTILUS, Vol. 108, No 3

Lea. I. 1836 A s\ nopsis of ttic family of naiades. Carey, Lea,

and Blaiuhard, Fhiladelpliia 59 pp.

Lea, I. 1838. Descriptions of new fresh- water and land shells

Transactions of the American Philosophical Society 6:1-

154.

Lea, I. 1852. A synopsis of the family of naiades. Blanchard

and Lea, Philadelphia. 88 pp.

Lea, I. 1858. New Unionidae of the LInited States Journal of

the Academy of Natural Sciences of Philadelphia 451-

95.

Lea, I. 1862. New Unionidae of the United States. Journal of

the Academy of Natural Sciences of Philadelphia 5:53-

109.

Lea, 1. 1870. A synopsis of the famiU of naiades Henry C

Lea, Philadelphia. 184 pp.

Miller, A. C. 1984. A survey for mussels on the lower Caney

Fork River, Tennessee, 18-20 October, 1983 U.S. Army

Waterways E.xperiment Station, Vicksburg, Mississippi 18

pp.

Neel, J. K. and W. R. Allen. 1964. The mussel fauna of the

upper Cumberland River before its impoundment

Malacologia 1:427-459.

Oesch, R. D. 1984. Missouri Naiades: a guide to the mussels

of Missouri. Missouri Department of Conservation, Jeffer-

son City. 270 pp.

Ortmann, A. E. 1917. A new type of the nayad-genus Fus-

conaia. Group of F. barncsiana Lea. The Nautilus 31:58-

64.

Ortmann, A. E. 1918. The nayades (freshwater mussels) of

the upper Tennessee drainage with notes on synonymy

and distribution Proceedings ol the American Philosoph-

ical Society 57:521-626.

Ortmann, A. E. 1924. The naiad-fauna of Duck River in

Tennessee. American Midland Naturalist 9:18-62.

Simpson, C. T. 1900. Synopsis of the naiades, or pearly fresh-

water mussels. Proceedings of the U.S. National Museum

22:501 -1044.

Simpson, C. T. 1914. A descriptive catalogue ot the naiades

or pearly fresh-water mussels. Brvant Walker, Detroit,

1540 pp.

Starnes, L. B. and A. E. Bogan. 1988. The mussels (Mollusca:

Bivalvia: Unionidae) of Tennessee. American Malacolog-

ical Bulletin 6:19-37.

Troschel, F H. 1839. Die Familie der Najaden. Archiv fiir

Naturgeschichte 5:235

van der Schalie, H and A van der Schalie 1950 The mussels

of the Mississippi River American Midland Naturalist 44:

448-466.

Yokes, H E 1980. Genera of the Bivalvia: a systematic and

bibliographic catalogue (revised and updated). Paleonto-

logical Research Institute, Ithaca, New York, 307 pp.

Wilson, C. B., and H. W. Clark. 1914. The mussel fauna of

the Cumberland River and its tributaries. Report of the

U.S. Commissioner of Fisheries for 1912, Special Paper:

1-63.

Wu, S.-K. 1978. The Bivalvia of Colorado, part 1. The fingernail

and pill clams (family Sphaeriidae). University of Colorado

Museum Natural History Survey of Colorado 2:1-39.

THE NAUTILUS 108(;3):61-66, 1995

Page 61

Richard S. Houbrick (1937-1993): Biographical Sketch and

Malacological Contributions

M.C. Harasewyeh

Alan R. Kabal

Department of Invertebrate Zoology

National Museum of Natural History

Smithsonian Institution

Wasfiington, DC. 20560 USA

I. BIOGRAPHY

The malacological community lost one of its most emi-

nent and respected members and prolific authors with

the death of Richard S. Houbrick on August 26, 1993 at

the age of 56, after a long and valiant struggle with

leukemia and hepatitis

Richard Steven Houbrick was born on March 16, 1937

in Trenton, New Jersey to Stephen J. and Barbara A.

Houbrick. The family moved to Hollywood, Florida when

Joe was 12. As a young student, Joe was an avid field

biologist, shell collector, and occasional shell show ex-

hibitor. After completing Broward High School, he en-

tered St. John's Seminary in Little Rock, Arkansas, then

transferred to St. Bernard College, Cullman, Alabama,

where he graduated with a Bachelor of Arts degree in

1959.

He then entered St. Leo Theological School, St. Leo,

Florida, and was ordained to the priesthood in March,

1963. As a member of St. Leo Abbey (Benedictine Order)

he took the name Joseph, and has since been known as

"Joe' to his many friends and colleagues. He was an

Instructor in general biology, zoology, invertebrate zo-

ology, embryology and botany at St. Leo College. He

began attending summer sessions of the University of

Miami and the University of Florida, taking courses in

Biology and Chemistry. In 1964, Joe entered the Grad-

uate School of the L'niversity of Miami. He earned a M.S.

degree from the Rosenstiel School of Marine and At-

mospheric Science, University of Miami in 1967, having

completed his thesis on "A survey of the littoral marine

mollusks of eastern Costa Rica.

In the summer of 1968, he attended the Biology of

Mollusks Training Program, in Hawaii, where he studied

moUuscan anatomy with Vera Fretter and others. This

course had a profound influence on his future research,

as shown by his extensive use of anatomy and repro-

ductive biology in his systematic publications.

Joe enrolled in the newly established LIniversity of

South Florida in 1968, and received the first Ph.D. to be

awarded by this University in 1971, for his dissertation

on "Taxonomy, anatomy, and life history studies on the

genus Cerithium (Gastropoda: Prosobranchia) in the

western Atlantic. His thesis advisor was Dr. Joseph L.

Simon, a marine ecologist. Upon completing his doctor-

ate, Joe joined the staff of the Smithsonian Oceanograph-

Richard S Houbrick at the Smithsonian Marine Station at Link

Port, Florida, February, 1983.

ic Sorting Center, and was subsequently appointed Su-

pervisor of the Benthic Section, and later Acting Director.

In 1977, Joe was appointed Associate Curator in the

Division of Mollusks at the National Museum of Natural

History, where he became Curator in 1981, a position in

which he remained active until shortly before his death.

Noted primarily for his expertise in the systematics,

comparative anatomy, fimctional morphology and re-

productive biology of the Cerithioidea, he contributed

significantly to the body of descriptive and synthetic

literature on this group. Since his work emphasized the

Page 62

THE NAUTILUS, Vol. 108, No. 3

study of living animals and llieir life-histories, he traveled

wideK , conducting field work in many parts of the world,

including the tropical western Atlantic, the tropical Pa-

cific and .\ustralia. He was a frequent invited participant

in s\niposia and workshops.

Since joining the Division of Mollusks, Joe spent an

extremely productive portion of each year working with

live animals at the Smithsonian Marine Station at Link

Port, Florida. The many colleagues who had an oppor-

tunity to work with Joe at the Marine Station soon dis-

covered his special interest in and enormous knowledge

of the flora and fauna of Florida that he had accumulated

since childhood. Joe was an active supporter of the Ma-

rine Station, serving on its Task Force and Advisory

Group.

All who knew Joe recognized that he was above all an

educator. He was an Adjunct Professor at George Wash-

ington University, and served on thesis committees of

graduate students at this and other universities. He also

served as Advisor to several Smithsonian Post-doctoral

Fellows, who have since risen to prominent positions in

the field of systematic malacology. Whether student, post-

doc or professional colleague, his own or someone else's,

Joe would always take the time to show an interest in

their work, to learn from them as well as to make helpful

comments and offer his own insights.

Joe was active in numerous professional organizations,

among them the American Malacological Union (of which

he was president in 1980-81), the American Society of

Zoologists, the California Malacozoological Society, the

Malacological Society of London, Unitas Malacologia, the

Biological Society of Washington, and the Smithsonian

Senate of Scientists.

Joe will be remembered as a quiet scholar whose re-

search and publications stand as a lasting monument to

his scientific dedication and insight. As a person, he was

known for his sense of humor, for his enjoyment of the

fine arts along with music, and he was invariably most

congenial to all. He is survived by his parents, and by

his brother James and his family.

ACKNOWLEDGMENTS

We thank James Houbrick, Robert Hershler, and CHyde

Roper for their helpful comments on this manuscript.

II MALACOLOGICAL TAXA

Of R.S. Houbrick s fifty-six publications from 1967 to

1993 (not counting meeting abstracts or notes), twelve

included descriptions of twenty-two new taxa. These

comprise two family-level taxa, three genus-level taxa

(two in the C>erilhiidae and one in the Plana.\idae) and

seventeen species-level taxa (twelve in the C^erithiidae

and the remainder in other prosobranch gastropod fam-

ilies). Two of the species-level taxa are replacement names

for junior homonyms.

Abbreviations

AMS Australian Mu,seuni, Sydney

ANSP Academy of Natural Sciences of Philadelphia

LACM Los Angeles County Museum of Natural His-

tory

MNHNP Museum National d'Histoire Naturelle, Paris

NM Natal Museum, Pietermaritzburg

NMFS National Marine Fisheries Service [USA]

USBF U.S. Bureau of Fisheries

USNM U.S. National Museum [= National Museum

of Natural History, Smithsonian Institution]

WAM Western Australian Museum, Perth

Family-level taxa

CERITHIDEIDAE Houbrick 1988, Malacological Re-

view, Supplement 4:88 jf. (diagnosis on page 118). For

(by implication) Cerithidea Swainson 1840. Subsequent-

ly svnonymized with the Potamididae H. & A. Adams

1854 by Houbrick (1991, Malacologia, 33(l-2):333). Cer-

ithioidea.

PLESIOTROCHIDAE Houbrick 1990, Proceedings of

the Third International Marine Biological Workshop: The

Marine Flora and Fauna of Albany, Western Australia,

pages 237 ff. (diagnosis on page 248). For Plesiotrochus

Fischer 1878. Originally placed in the Cerithioidea. Sub-

sequently transferred to the Campaniloidea by Healy

(1993).

Genus-level ta.xa

Ittibittium Houbrick 1993, Malacologia, 35(2):283. Type

species Bittium parcuni Gould 1861; monotypy. Ceri-

thiidae.

Longicerithium Houbrick 1978, Monographs of Marine

Mollusca, 1:85. Type species Cerithium longicaudata A.

Adams and Reeve 1850; original designation. As a sub-

genus of Rhinoclavis Swainson 1840. Cerithiidae.

Simulathena Houbrick 1992, The Veliger, 35(l):64-65.

Type species Simulathena papuenais Houbrick 1992;

monotypy. Planaxidae.

Species-level taxa

abditum, Cerithium Houbrick 1992, Smithsonian Con-

tributions to Zoology, 510:18-20, figures 6-8. Holotvpe

USNM 286404; five paratypes USNM 860482. USBF'a/-

batross Station 5569, 554 meters, off Simaluc Island,

Tawitawi, Philippines. Cerithiidae.

admirabilis, Clypeontorus Houbrick 1985, Smithsonian

C:ontributions to Zoology, 403:99-104, figures 47-49. Ho-

lotype WAM 3346-83; seven paratypes USNM 784652;

thirty-eight paratypes ANSP 233431; three paratvpes

AMS 139779. False Cape Creek (near mouth), 100 miles

(161 kilometers) S of Broome, Western Australia, .Aus-

tralia. Cerithiidae.

africanum, Cerithium Houbrick 1992, Smithsonian

Contributions to Zoology, 510:20-23, figures 9-11. Ho-

lotype NM HI 861; five paratypes USNM 629034. South-

western Conducia Bay, NW of Choca, Mozambique.

Cerithiidae.

M. G. Harasewych and A. R. Kabat, 1995

Page 63

argentea, Strombina (Cotonopsis) Houbrick 1983, Pro-

ceedings of the Biological Society of Washington, 96(3):

349-352, Bgures 1-2. Holotvpe USNM 810345; nine para-

types USNM 821851. NMFS R/V Oregon. Station 5432,

384-430 meters, 20°48'N, 70°46'W, near Silver Bank, 60

miles NE of Luperon, Dominican Republic. Columbel-

lidae.

diadema, Rhinoclavis (Rhinoclavis) Houbrick 1978,

Monographs of Marine Molkisca, 1:64-66, figures 33-35.

Replacement name for Cerithitim nitidum Honibron

and Jacquinot 1854 non MacAndrew and Forbes 1847

et al.. Lectotype and one paralectotype (of CerUhiiim

nitidum), MNHNP (not numbered), selected by Houb-

rick (1978: 66). Mangareva, Gambier Islands. Cerithi-

idae.

exiinium, Ataxocerithium Houbrick 1987, The Nautilus,

101(4): 157-159. figures 1-16. Holotype AMS C142394;

five paratvpes AMS C153005; two paratypes L'SNM

862328. 421 meters, 34°21-16'S, 151°24-28'E, SE of Bot-

any Bay, New South Wales, Australia. Cerithiopsidae.

garciai. Cerithioclava Houbrick 1986, Proceedings of the

Biological Society of Washington, 99(2):257-260, figure

1. Holotype USNM 849023; two paratypes USNM 849024.

24 meters, off Great Corn Island, Nicaragua [Caribbean

Sea]. Cerithiidae.

gloriostun. Cerithium Houbrick 1992, Smithsonian Con-

tributions to Zoology, 510:94-97, figures 61-63. Holo-

type, MNHNP (not numbered); 200 paratypes MNHNP

(not numbered); seventy-nine paratypes USNM 862327.

Benthedi Station DR-06, 500-600 meters, 11°28'S,

47°12'E, lies des Glorieuses, Indian Ocean. Orithiidae

inflatiis, Acamptochetus Houbrick 1984, Proceedings of

the Biological Society of Washington, 97(2):421-423, fig-

ure 1. Holotype USNM 229183; one paratype USNM

820185. USBF Albatross Station 5419, 320 meters,

9°58'30"N, 123°46'E, off Lubigon, Bohol, Philippines.

Buccinidae.

nympha, Clypeomorus Houbrick 1985, Smithsonian

Contributions to Zoology, 403:113-120, figures 56-59.

Replacement name for Cerithium variegatum Quoy and

Gaimard 1834 non Fischer 1807 et al.. Lectotype and

one paralectotype (of Cerithium variegatum), MNHNP

[not numbered]; selected by Houbrick (1985: 113). Ton-

gatapu [Tonga Islands]. Cerithiidae.

pacificum Cerithium Houbrick 1992, Smithsonian Con-

tributions to Zoology, 510:143-146, figures 103-104. Ho-

lotype USNM 584662; four paratypes USNM 862491.

55-61 meters, Bikini lagoon, Bikini Atoll, Marshall Is-

lands. Cerithiidae.

papuensis, Simulathena Houbrick 1992, The Veliger,

35(l):65-67, figures 1-13. Holotype AMS C166326; two

paratypes USNM 859456. 8°50'S, 146°32'E, Kairuku, Yule

Island, Central District, Papua New Guinea. Planaxidae.

persica, Clypeomorus bifasciata (Sowerby 1855) netv

subspecies Houbrick 1985, Smithsonian Contributions to

Zoology, 403:41-43, figures 17-18. Holotype, ANSP

263194; eight paratypes USNM 838028. Ras fanura. Per-

sian Gulf, Saudi Arabia. Cerithiidae.

purpurastoma, Clypeomorus Houbrick 1985, Smithson-

ian Contributions to Zoology, 403:83-89, figures 38-40.

Holotype L'SNM 774749; sixteen paratypes USNM

784650 Napot Point, Morong, Bataan, Luzon, Philip-

pines. Cerithiidae.

rehderi, Cerithium Houbrick 1992, Smithsonian Con-

tributions to Zoology, 510:156-158, figures 112-113. Ho-

lotype USNM 859954; five paratypes USNM 798574. 56-

72 meters, Haava Straits, between Tahuata and Hiva Oa,

Marquesas. Cerithiidae.

reidi. Cerithidea Houbrick 1986, The Veliger, 28(3):280

ff., figures 1-16. Holotype WAM 3380-84; one paratype

AMS C144144; ten paratypes USNM 828823. Willies

Creek, N of Broome, Western Australia, Australia. Po-

tamididae.

scobiniforme, Cerithium Houbrick 1992, Smithsonian

Contributions to Zoology, 510:177-179, figures 129-130.

Holotype USNM 859930; two paratypes USNM 859931;

six paratypes LACM 76-702. 1.5 miles S of Estango Is-

land, Port Dimalosan, Cagayan Province, Luzon, Phil-

ippines. Cerithiidae.

III. PUBLICATIONS

R.S. Houbrick authored or co-authored a total of fifty-

six research papers and notes, along with eighteen ab-

stracts, two "President's Messages " (AMLI) and two grad-

uate theses over a twenty-seven year period (1967-1993).

Of the research papers, thirty-eight (68%) treated mem-

bers of the Cerithioidea (twenty-five specifically on the

Cerithiidae), thirteen (23%) were on other prosobranch

gastropod families, and five (9%) were on general topics.

As for publication venue, the following statistics were

obtained. Thirty-one (55%) of his research papers were

published in malacological serials, twenty (36%) were

published in non-malacological serials (primarily the

Proceedings of the Biological Society of Washington and

the Smithsonian Contributions to Zoology), and the re-

maining five (9%) were published as book chapters.

Note: his papers for 1967-1971 were authored as "JR.

Houbrick"; those for 1972-1974b as either "R.S. Houb-

rick or "R.S. (JR.) Houbrick", and the remainder as

■R.S. Houbrick".

Abbreviations

AMU American Malacological Union

WSM Western Society of Malacologists

1967 [24 April]. Notes on Cyctostremiscus .whramnni The

Nautilus 80(4):131-133.

1968a [1 Jan ] New record of Conu4c/?rae2i,s in Costa Rica

The Veliger 10(3):292.

1968b [1 July]. A survey of the littoral marine mollusks of

the Caribbean coast of Costa Rica. The Veliger 11(1):4-

23.

Page 64

THE NAUTILUS, Vol. 108, No. 3

1969 [.^ug.] Houbriok, J R and Fretter, V Some a.spects of

the functional anatomy and biology of Cymatium and

Bursa. Proceedings of llie Malacological Society of London

38(5):415-429.

1970 [June]. Houbrick, J.R. and Fretter, V. Some aspects of

the functional anatomy and biology of Cymatium and

Bursa. In: (Kay, E.A., ed.). The Biology of Molluscs. A

collection of abstracts from the National Science Foun-

dation Graduate Research Training Program June - Sep-

tember 1968, pages 12-13. University of Hawaii, Hawaii

Institute of Marine Biology, Technical Report 18, 30 pages.

[Abstract]

1971a "1970 [18 Feb ] Reproduction and development in

Florida Cerithium AMU, Key West, Florida, 16-20 July

1970. Annual Report of the American Malacological L'nion

for 1970, page 74. [Abstract]

1971b [26 Oct.]. Some aspects of the anatomy, reproduction,

and early development of Cerithium nodulosum (Bru-

guiere) (Gastropoda, Prosobranchia) Pacific Science 25(4):

560-565.

1972 [Nov.]. The Smithsonian Institution - Great Lakes role.

Great Lakes Basin Commission, Communicator 3(5):3-4.

1973 [10 Oct! CcnV/iium Bruguiere, [1789], (Gastropoda);

Proposed preservation by designation of a type-species

under the plenary powers. Bulletin of Zoological Nomen-

clature 30(2)404-107.

1974a [29 Jan ]. Growth studies on the genus Cerithium

(Gastropoda: Prosobranchia) with notes on ecology and

microhabitats. The Nautilus 88(1); 14-27.

1974b "1973" [7 May]. Studies on the reproductive biology

of the genus Cerithium (Gastropoda: Prosobranchia) in

the Western Atlantic. Bulletin of Marine Science 23(4);

875-904.

1974c [29 July]. The genus Cerithium in the Western At-

lantic (Cerithiidae; Prosobranchia) Johnsonia 5(50)33-84.

1975 [29 Oct.]. Clavocerithium (Indocerithium) taeniatum,

a little-known and unusual cerithiid from New Guinea

The Nautilus 89(4);99-105.

1976 "1975" [30 Jan.]. Preliminary revision of supraspecific

taxa in the Cerithiinae Fleming, 1882 (Cerithiidae: Pros-

obranchia) Bulletin of the American Malacological Union,

Inc. for 1975, pages 14-18, 18a.

1977 [1 Oct.]. Reevaluation and new description of the genus

Bittium (Cerithiidae). The Veliger 20(2); 101-106.

1978a [31 Jan ] Redescription of Bittium proteum (Jous-

seaume, 1930) with comments on its generic placement.

The Nautilus 92( I ):9-ll,

1978b [April]. Form, function, and adaptive radiation in

snails of the family Cerithiidae. AMU, Naples, Florida,

10-15 July 1977. Bulletin of the American Malacological

Union, Inc. for 1977, page 84. [Abstract]

1978c [6 Dec.]. Reassignment of Batillaria sordida (Gmelin)

from the Cerithiidae to the Potamididae (Gastropoda:

Prosobranchia). Proceedings of the Biological Society of

Washington 91(3);642-649

1978d [15 Dec ] The family Cerithiidae m the Indo-Pacific

Part 1; The genera Rhinoclavis. Pseudovertagus and Chiv-

ocerithium. Monographs of Marine Mollusca 1:1-130

1979 [5 June]. Classification and systematic relationships of

the Abyssochrysidae, a relict family of bathyal snails (Pro.s-

obranchia; (Jastropoda) Smith.sonian Ointributions to Zo-

ology 290:1-21.

1980a [24 March]. Reproductive biology of Modttlu.s mod-

ulus Linnaeus. 1758 (Prosobranchia: Ceritliiacea). AML'

and WSM, (;orpus Christi, Texas, 5-11 August 1979. Bul-

letin of the American Malacological L'nion, Inc for 1979,

page 71 [title only]; The Western Society of Malacologists

Annual Report 12:10. [Abstract]

1980b [30 June] Form, function and adaptive radiation in

the Cerithiidae. Symposium on the biology and evolution

of Mollusca, Australian Museum, Sydney, 21-25 May 1979.

Journal of the Malacological Society of Australia 4(4):230-

231. [Abstract]

1980c [23 July] Review of the deep-sea genus Argyropeza

(Gastropoda: Prosobranchia: Cerithiidae). Smithsonian

Contributions to Zoology 321:1-30.

1980d [22 Aug ]. Observations on the anatomy and life his-

tory of Moduhis modulus (Prosobranchia: Modulidae).

Malacologia 20(1):1 17-142.

1980e [Aug.]. Anatomy and biology of Campanile symhul-

icum Iredale with reference to adaptive radiation of the

Cerithiacea. Unitas Malacologica, Perpignan, France, 31

Aug. - 7 Sept. 1980. In: (Gaillard, J. M., ed). [AbstractsJ

Septieme Congres International de Malacologie. Haliotis

10(2): 168. [Abstract]

1980f [6 Nov,]. Reapprai.sal of the gastropod genus Varico-

peza Griindel (Cerithiidae: Prosobranchia). Proceedings

of the Biological Society of Washington 93(3):525-535.

1980g Systematics, anatomy and biology of Campanile sym-

bolicum. a relict cerithiacean snail. AMU, Louisville, Ken-

tucky, 19-25 July 1980 Bulletin of the American Mala-

cological Union for 1980, page 72. [Abstract]

1980h Presidents Message. The .American Malacological

Union, Inc., Newsletter, lI(2-A):l-2.

1981a [29 Jan ]. .Anatomy and systematics of Gourmya gour-

mi/i (Prosobranchia: Cerithiidae), a Tethyan relict from

the Southwest Pacific. The Nautilus 95(1):2-1I.

1981b [20 July] Anatomy of Dias/oma nip/anioirfes (Reeve,

1849) with remarks on the systematic position of the family

Diastomatidae (Prosobranchia: Gastropoda). Proceedings

of the Biological Society of Washington 94(2):598-621.

1981c [16 Oct ]. Systematic position of the genus C.lyptozaria

Iredale (Prosobranchia: Gastropoda) Proceedings of the

Biological Society of Washington 94(3):838-847.

1981d [8 Dec ]. .Anatomy, biology and systematics of Cam-

panile symbolicum with reference to adaptive radiation

of the Cerithiacea (Gastropoda: Prosobranchia). Malaco-

logia 21(l-2):263-289.

1981e What is Diastema? Systematic position of the Dias-

tomatidae AMU. Fort Lauderdale, Florida, 19-25 July

1981 Bulletin of the American Malacological L'nion for

198L page 31 [Abstract]

198 If President's Message The American Malacological Union,

Inc., Newsletter 12(1):1.

1983 [31 Oct.]. A new Strombina species (Gastropoda: Pros-

obranchia) from the tropical Western Atlantic. Proceed-

ings of the Biological Societ\ of Washington. 96(3):349-

354.

1984a [Feb.] Revision of higher taxa in genus Cerithidea

(Mesogastropoda: Potamididae) based on comparative

morphology and biological data American Malacological

Bulletin 2:1-20, 95.

1984b [29 March]. Functional morpholog) and ontogeny of

Mollusca as related to higher category systematics; Intro-

duction. Malacologia 25(I):1.

1984c [April], (Joing collecting? Look for a live Royella si-

mm Hawaiian Shell News 32(4); 12.

1984d [31 Ma\ ] Ke\ision of higher taxa in genus CVn7/!i(ipa

based on comparative morphology and biological data.

AMLI and WSM, I'niv. of Washington, Seattle, 7-13 Aug.

M. G. Harasewvch and A. R Kabat, 1995

Page 65

1983 The Western Society of \4alacologists. Annual Re-

port 16:23. [Abstract]

l*)84-e [6 Jub ] A new "Mctula" species from the Indo-West

Pacific (Prosobranchia; Bucciniclae). Proceedings of the

Biological Society of Washington 97(2)420-424.

1984f [19 July]. The Giant Creeper, Campanile symbolicum

Iredale, an Australian relict marine snail. In: (Eidredge,

N. and Stanley, S.M eds. ), Living Fossils, pages 232-235

New York, Springer Verlag, xii + 291 pages.

I'JBtg [19 Jub] Diastoma mclaniuidcs (Reeve), a relict snail

Iriiui South Australia. In: (Eidredge, N. and Stanley, S.M.

eds.), Living Fossils, pages 236-239. New York, Springer

Verlag, xii + 291 pages.

1081-h [19 July] The relict cerithiid prosobraiich, Gourmya

gutirniyi (Crosse). In: (Eidredge. N. and Stanley, S.M. eds.).

Living Fossils, pages 240-242. New York, Springer Verlag,

xii + 291 pages.

IQSSa "1984" [Feb.]. Comparative reproductive biology of

P/aiiflxis species. AMU, Norfolk, Virginia, 22-27 July 1984.

American Malacological Bulletin 3(1)96. [Abstract]

1985b [3 April] Genus CUjpcomorus Jousseaume (Cerithi-

idae: Prosobranchia). Smithsonian Contributions to Zool-

ogy 403:1-131

1986a [2 Jan.]. Cerithidea reidi. spec, nov., from Western

Australia. The Veliger 28(3):280-286.

1986b [Feb.] C/i/pfomoru.s, a genus of littorinid-like cerith-

ids. AMU, Univ. of Rhode Island, Kingston, 28 July - 2

Aug. 1985. American Malacological Bulletin 4( 1 ) 109, [Ab-

stract]

1986c [1 April]. The systematic position of Royella sinon

(Bavle) (Prosobranchia: Cerithiidae). The Veliger 28(4):

429-435.

1986d [4 June]. Discovery of a new living Cerithioclava

species in the Caribbean (Mollusca: Prosobranchia: Ceri-

thiidae). Proceedings of the Biological Society of Wash-

ington 99(2):257-260.

1986e [4 June] Transfer of Quadrasia from the Planaxidae

to the Buccinidae (Mollusca: Gastropoda: Prosobranchia).

Proceedings of the Biological Society of Washington 99(2):

359-362.

1986f [June] Observations on the anatomy and life histories

of cerithiacean prosobranchs. In: (Rice, M., ed). Abstracts,

Symposium on Current Research at the Smithsonian Ma-

rine Station at Link Port, June 24 - 25, 1986. Fort Pierce

(Florida), Smithsonian Marine Station [unpaginated]. [Ab-

stract]

1986g [Aug ] Functional anatomy and systematicsof Litiopa

and Alaba (Prosobranchia: Orithiacea). AML' and WSM,

Monterey, California, 1-6 July 1986, American Malaco-

logical Bulletin 4(2):235. [Abstract]

1986h [Aug ]. Phylogenetic relationships in the superfamily

Cerithiacea. Unitas Malacologica, Edinburgh, Scotland, 31

Aug. - 6 Sept 1986 In: (Heppell, D , ed). Abstracts, Unitas

Malacologia. Ninth International Malacological Congress,

page 38. Edinburgh, National Museums of Scotland, 104

pages. [Abstract]

1986i Prosobranch Mollusca of Carrie Bow Cay In: (Riitzler,

K , ed), Caribbean Coral Reef Ecosystems, Progress Re-

port 1986, page 42. Washington, DC, National Museum

of Natural History, Smithsonian Institution, 49 pages. [Ab-

stract]

1987a [30 Jan.]. Anatomy of Alaba and Litiopa (Prosobran-

chia: Litiopidae): systematic implications. The Nautilus

101(1):9-18.

1987b [18 March]. Anatomy, reproductive biology, and phy-

logeny of the Planaxidae (Cerithiacea: Prosobranchia).

Smithsonian C^jntributions to Zoology 445:1-57.

1987c [28 .'Vpril] Transfer of Cerithiopsifi crystallina Dall

to the genus Varicupeza Griindel, family Cerithiidae

(Prosobranchia: Gastropoda). The Nautilus 101(2):80-85.

1987d [31 July]. Houbrick, R.S., Robertson, R , and Abbott,

R T. Anatomy and systematic position of Fasligiella car-

inata Reeve (Cerithiidae: Prosobranchia) The Nautilus

101(3):10I-110

1987e [6 Nov.], Description of a new, giant Ataxoccrithium

species from Australia with remarks on the systematic

placement of the genus (Prosobranchia: Cerithiopsidae).

The Nautilus 101(4):155-161.

1988a [15 Oct.]. Houbrick, R S., Stiirmer, W. and Yochelson,

EL. Rare Mollusca from the Lower Devonian Hunsriick

Slate of southern Germany. Lethaia 21(4):395-402.

1988b [20 Dec]. Cerithioidean phylogeny In: (Ponder, WF.,

et ai, eds ), Prosobranch Phylogeny. Malacological Re-

view, Supplement 488-128.

1989a [April] Campanile revisited: implications for ceri-

thioidean phylogeny. American Malacological Bulletin 7(1):

1-6.

1989b [Aug.]. Functional morphology and systematics of

mangrove snails Terebralia and Telescopium (Potamidi-

dae; Prosobranchia). Unitas Malacologica, Universitat Tii-

bingen. West Germany, 27 Aug. - 3 Sept. 1989 In: (Meier-

Brook, C, ed). Abstracts, Unitas Malacologica, Tenth In-

ternational Malacological Congress, page 106 Tiibingen,

Tropenmedizinischen Institijt der Universitat Tubingen,

li + 287 pages. [Abstract]

1989c [29 Sept ]. Comments on the proposed conservation

of Iphinoe Bate, 1856 (Crustacea, Cumacea). Bulletin of

Zoological Nomenclature 46(3):191.

1 990a [ 1 1 June]. Anatomy, reproductive biology and system-

atic position of Fossarus ambiguus (Linne) (Fossaridae:

Prosobranchia). AML' and WSM, Univ. of Southern Cal-

ifornia, Los Angeles, 25-30 June 1989 Western Society of

Malacologists Annual Report 2222. [Abstract]

1990b [6 Sept ] Review of the genus Co/ina H and A. Ad-

ams, 1854 (Cerithiidae: Prosobranchia). The Nautilus

104(2):35-52.

1990c [Oct.] Anatomy, reproductive biolog) and systematic

position of Fossarus ambiguus (Linne) (Fossarinae: Plan-

axidae; Prosobranchia) In: (Frias Martins, A.M., ed). The

Marine Fauna and Flora of the Azores Proceedings of the

First International Workshop of Malacology, Sao Miguel,

Azores, 11-24 July 1988, pages 59-73. A^oreana, Suple-

niento, 173 pages.

1990d [31 Dec.]. Aspects of the anatomy of Plesiotrochus

(Plesiotrochidae, fam. n.) and its systematic position in

Cerithioidea (Prosobranchia, Caenogastropoda). In: (Wells,

F.E., Walker, D.I., Kirkman, H & Lethbridge, R., eds).

Proceedings of the Third International Marine Biological

Workshop: The Marine Flora and Fauna of Albany, West-

ern Australia, pages 237-249. Perth, Western Australian

Museum, volume 1 (of 2), x -t- 437 pages.

1991a [15 July] Functional inference from gastropod shell

morphology - some caveats Lethaia 24(3):265-270.

1991b [6 Sept.], Systematic review and functional morphol-

ogy of the mangrove snails Terebralia and Telescopium

(Potamididae; Prosobranchia). Malacologia 33(l-2):289-

338.

1991c [16 Dec.]. Anatomy and systematic placement of Fai/-

nus Montfort 1810 (Prosobranchia: Melanopsinae). Mal-

acological Review 24(l-2):35-54.

Page 66

THE NAUTILUS, Vol. 108, No. 3

1992a [2 Jan ]. Sinitilathcna pajmcnsis. a new planaxid genus

antl species from the Imlo-West Pacific. The Vehger 35(1):

64-69.

1992b [8 June]. Anatomy and prehminary generic review of

the Bittium group. AMLI and WSM, Univ. of California,

Berkeley, 30 June - 5 July 1991. Western Society of Ma-

iacologists, Aniuial Report 24:27. [Abstract]

1992c [29 Sept.]. Monograph of the genus Cerithiuiu Bru-

guiere in the Indo-Pacific (derithiidae: Prosobranchia)

Smithsonian C'ontributions to Zoology 510, iv + 211 pages.

1993a [24 March]. Two confusing Indo-Pacific cerithiids.

The Nautilus 107(l):14-23.

1993b [2 Dec.]. Phylogenetic relationships and generic re-

view of the Bittiinae (Prosobranchia: Cerithioidea).

Malacologia 35(2):261-313.

NOTE: Dr. Winston F. Ponder (Australian Museum) is

editing and revising several uncompleted manuscripts

left by R.S. Houbrick for eventual publication.

Graduate Theses

1967 [May]. A survey of the littoral marine mollusks of East-

ern Costa Rica. Master of Science, University of Miami,

Coral Gables, Florida, iv + 90 pages, 5 plates. See Houb-

rick, 1968b.

1971 [May]. Taxonomy, anatomy and life history studies on

the genus Cerithium (Gastropoda: Prosobranchia) in the

Western Atlantic. Doctor of Philosophy, University of South

Florida, Tampa, Florida xii -I- 182 pages. See Houbrick,

1974a, 1974b and 1974c.

OTHER LITERATURE CITED

Healy, J M. 1993. Transfer of the gastropod family Plesi-

otrochidae to the C^ampaniloidea based on sperm ultra-

structural evidence Journal ol Moiluscan Studies 59(2):

135-146.

THE NAUTILUS 108(3):67-79, 1995

Page 67

New Data on the Distribution and Morphology of some Western

Indian Ocean Sepiid Cuttlefishes (Cephalopoda: Sepiida)

Yu. A. Filippova

D. N. Khroniov

Russian Research Institute of Marine

Fisheries & Oceanography (VNIRO)

1071-10 Moscow RUSSIA

K. N. Nesis

I. V. Nikilina

P F. Shirsliov Institute of Oceanology

Russian Academy of Sciences

117218 Moscow RUSSIA

ABSTRACT

New data are presented on the distribution and/or morphology

of 21 species of the family Sepiidae, based on the collections

of various expeditions during 1956-1989 in the Western Indian

Ocean from the Northern Arabian Sea to Southern Mozam-

bique, Agulhas Bank and Cargados-Carajos Islands Previously

neglected or inadequately described characters are provided

for Sepia joulnni, S. confusa. S. typica and others. New dis-

tribution records are provided for S. ornani off Pakistan, S.

acuminata off Somalia, Kenya and Madagascar, S. hieronis and

S. simoniana off Kenya, S. joubini, S. incerta, S. confusa. S.

burnupi, S. hieronis, S. simoniana, S. officinalis vermiculaia

and S. zanzibarica on Saya-de-MaIha Bank The Eastern At-

lantic species S. elegans was recorded from the SW Indian

Ocean on the Agulhas Bank Four species were found at shal-

lower depths and seven at greater depths than recorded pre-

viously Five species are recorded at depths greater than 300 m

Key words Sepiida, Sepia, cuttlefish, species, morphology,

distribution, new records

INTRODUCTION

The cuttlefish (Sepiidae) fauna of the Indian Ocean is

incompletely known The area has been relatively well

studied in the northern part of the ocean (Adam, 1959;

Adam & Rees, 1966; Silas ct ai, 1986), in the south-

western part off South Africa (Voss, 1962; Roeleveld,

1972) and to some extent in the southeastern part off

Western Australia (Adam, 1979). However, the Western

Indian Ocean between the Gulf of Aden and Southern

Mozambique, including many islands, archipelagoes and

shallow banks (Socotra, Seychelles, Madagascar, Saya-de-

Malha, Cargados-Carajos) remain almost unexplored. This

broad area was studied during the last several years by

various Soviet expeditions organized by many USSR in-

stitutions; All-Union (now Russian) Research institute of

Marine Fisheries and Oceanography (Moscow), Southern

Research Institute of Fisheries and Oceanography

(Kerch), Institute of Oceanology of the LiSSR (now Rus-

sian) Academy of Sciences (Moscow), Institute of the

Biology of the Southern Seas, Ukrainian Academy of

Sciences (Sevastopol), Moscow State University, "Hy-

dronaut' Base (Sevastopol), etc. As a result of these ex-

peditions large amounts of data were gathered, partic-

ularly on the taxonomy, distribution and biology of cut-

tlefishes (Zuev ct al., 1968; Zuev, 1971; Druzhinin &

Filippova, 1974; Homenko, 1976, 1983, 1989; Nesis, 1980,

1993; Homenko & Khromov, 1984; Khromov, 1982, 1987,

1988a; Filippova & Khromov, 1991; Khromov et ai,

1991). As a result of these expeditions, the sepiid fauna

of the Western Indian Ocean was found to be richer and

more diverse than was previously supposed. Ten new

species have been described to date, and others are await-

ing description. Some species are common and may have

commercial potential. The data gathered permit a fresh

examination of the composition, distribution and rela-

tionships of the sepiid fauna of the Indian Ocean as a

whole. We present new data on the distribution of several

known species, and on the morphology of some species

described incompletely or inadequately. All but one are

inhabitants of the Indian Ocean — Sepia elegans is an

Atlantic species recorded for the first time in the Indian

Ocean off South Africa.

This work is based primarily on the catches made in

1976 by the R/V "Professor Mesyatzev" off East Africa

between 2°S and 25°S, off the Cargados-Carajos Islands

and on Saya-de-Malha Bank, and by the R/V "Vityaz"

in 1988-89 oft Socotra, Madagascar, Mozambique and

on Saya-de-Malha Bank. We have also studied collections

of many other expeditions, and are grateful to the col-

lectors of the following material: G. M. Belyaev ("Ob",

1956), V. N. Semenov ("Akademik Knipovich ', 1965-

66), the late V. I. Chekunova ("Akademik Knipovich",

1967), the late S. V. Mikhailin ("Gizhiga", 1975), B. G.

Ivanov ("Professor Mesyatzev", 1975-76), L. S. Hom-

enko ("Nauka", 1976, "Zheleznyakov", 1977, "Yelsk",

1977-78), N. S. Khromov ("Fiolent", 1976, "Sabah",

1979), V. A. Spiridonov ("Odissey", 1980), M. G. Kar-

pinsky ("Professor Mesyatzev", 1983), D. L. Ivanov

("Akademik Petrovsky", 1984), Yu. S. Korzun ("Skif",

1986), V. A. Bizikov and D. O. Alekseyev ("Akademik

Knipovich", 1984, "Odissey", 1985, "Gidrobiolog", 1987).

All specimens are either in the Russian Research Institute

THE NAUTILUS, Vol. 108, No. 3

^f^c:

-"ii

Yii. A. Filippova et al , 1995

Page 69

of Marine Fisheries and Oceanography (VNIRO), the

Zoological Museum of Moscow llniversity (ZMML'), or

in the Institute of Oceanology of the Russian Academy

of Sciences (lOAS), Moscow.

SYSTEMATICS

Family Sepiidae

1. Sepia (Doratuscpion) jotibini Massv, 1927

(Figs. 1, 2, 20)

Material examined: "Vitvaz", 25/11/1988, sta. 2634a,

bottom trawl, 90-92 m, 25°05'S, 34°50'-34°44'E: 1 fe-

male ML 42 mm, lOAS; "Professor Mesyatzev", 24/01/

1976, sta. 64, bottom trawl, 167-170 m, 24°23'S, 35°29'E:

1 female ML 64 mm, ZMMU 327.

Description and remarks: This species was described

by Massy (1927), redescribed (syntypes) by Adam and

Rees (1966), and again redescribed (topotypes) by Roe-

leveld (1972), In spite of the large number of described

specimens of both se.xes, the sepions of S. joubini remain

nearly unknown because they were damaged and de-

calcified in both the syntypes and the topotvpes.

Adam and Rees (1966) doubted the validity of S. joub-

ini. According to Roeleveld (1972) the cuttlebones of the

topotypes were narrow and elongated, pointed anteri-

orly, with a median longitudinal ridge dorsally and groove

ventrally. The striated zone is convex on both sides of

the median groove, angular anteriorly, becoming flat-

tened posteriorly, with convex and wavy striae posteri-

orly. The inner cone has narrow limbs that form a ledge

posteriorly. The outer cone has posterior wings. Roelev-

eld (1972) also mentioned a peculiar structure at the base

of the spine, but it is not known whether it occurs in all

specimens. Roeleveld's figure of S. joubini illustrates an

expansion of the posterior part of the imier cone that is

not described in the text With this discrepancy and her

description of a decalcified sepion, unambiguous iden-

tification of this species is difficult.

There are two females of S. joubini in our material,

one (sta. 2634a) properly fixed and in good condition.

Therefore we are able to add several important details

to the description of this species.

The dorsal surface of the mantle of the living animal

has small red spots and interrupted light-colored stripes

along the fins that form a series of elongate, raised bolsters

as in S. (D.) peterscni Appelloef, 1886, and S. (D.) viet-

namica Khromov, 1987. The fins are narrow and slightly

expanded posteriorlv The characteristic arms of the fe-

male differ in length (lateral arms longer than the dorsal

and ventral), have attenuated tips and well developed

protective membranes (Fig. 1), and correspond to Roe-

leveld s (1972) description. Longitudinal red stripes are

found on the dorsal side of the head and on dorsal and

lateral arms of living animals, but are not evident on

fixed specimens. The sepion is dark-cream colored, nar-

rowly elongate, width 24*^(, thickness 9% of length. A

metlian rib, not limited b> grooves, lies on the dorsal

surface of the sepion. Only the median third of the dorsal

shield is calcified and bordered by wide, chitinous mar-

gins. The sepion is semi-rounded in cross-section, with a

flattened middle part of phragmocone. The median

groove, shallow on the striated zone and deep on the last

loculus, is clearly marked along the whole length of the

phragmocone. The striated zone occupies the entire width

on the ventral side, and is bordered by smooth marginal

zones. Anterior striae are reversed V-shaped. The inner

cone forms a ledge that is slightly expanded posteriorly,

but without any blade-like structure, as in Roeleveld s

figure. Inner cone limbs are close to the chitinous mar-

gins. The outer cone forms a regular, cup-like, decalcified

structure posteriorly. The spine is long, narrow at its base,

slightly expanded in the middle part, and spindle-like in

lateral view.

We confirm Roeleveld s opinion on the validity of S.

joubini and consider this species to be fully described

now.

A female (ML 64 mm) in these collections (ZMMU)

is the largest known specimen.

Distribution: Sepia joubini was recorded off South-

eastern Africa, from the Tugela River to Cape Natal

(Roeleveld, 1972). We foimd this species off Southern

Mozambique, 24-25°S. The depth range of 66-134 m

(Roeleveld, 1972), is expanded to 66-170 m

2. Sepia (Doratosepion) incerta Smith, 1916

(Figs. 3, 4, 20)

Material examined: "Professor Mesyatzev , 18/01/1976,

sta. 41, bottom trawl, 100 m, 18°29'S, 37°17'E: 3 males,

ML 44, 115, 118 mm, 2 females. ML 82, 88 mm; "Pro-

fessor Mesyatzev ', 19/01/1976, sta. 44, bottom trawl,

152-172 m', 19°07'S, 36°59'E: 1 male, ML 123 mm; "Pro-

fessor Mesyatzev", 24/01/1976, sta. 64, bottom trawl,

167-170 m', 24°23'S, 35°29'E: 2 males, ML 53, 111 mm,

all specimens are in the collections of ZMMU, No 185,

212, 230, 232, 334. "Professor Mesyatzev", 31/01/1976,

sta. 77, bottom trawl, 330-345 m, 25°21'S, 34°19'E: 1

female, ML 82 mm, VNIRO. "Vityaz", 25/11/1988, sta.

Figures L 2. Sepia (Doratosepion) jotihini Massy, 1927. MI. 42 mm, 2.5°05'S. 34°.50'-34°44'E, 90-92 mm, lOAS, uncatalogued 1.

Head of female. 2. Sepion of female. Figures 3, 4. Sepia {Duratusepion) incerta Smith, 1916, ML 11.5 mm, 1S°29'S, 37°17'E, 100

m, ZMMU No. 212. 3. Dorsal and 4. lateral views of sepion of male. Figures 5, 6. Sepia (Doratosepion) confusa Smith, 1916. ML

103 mm, 24°23'S, .35°29'E, 167-170 m, VNIRO, uncatalogued 5. Sepion of male 6. Posterior part of male sepion Figure 7. Sepia

(Doratosepion) trygonina Rochebrune, 1884. ML 65 mm, Saya-de-MaIha Bank, VNIRO, uncatalogued Sepion of male. All scale

bars = 1 cm. as, anterior striae; ds, dorsal shield; ic, inner cone; lie, limbs of inner cone; II, last loculus; mg, median groove (or

furrow); oc, outer cone; p, phragmocone; s, spine; smz, smooth marginal zone; sz, striated zone; s, spine; w, wings of outer cone

Page 70

THE NAUTILUS, Vol. 108, No. 3

cm

Yu. A. Filippova et al., 1995

Page 71

2634a, bottom trawl, 90-92 m, 25°05'S, 34°50'-34°44'E:

1 male ML 45 mm. 1 female, ML 91 mm, lO.^S.

Description and remarks: All our specimens are quite

typical and agree completely with the descriptions of

Adam and Rees (1966) and Roeieveld (1972) Large males

are characterized by a peculiar transformation of the

dorsal arms. Males and females differ from all other

species in having a furrow on the posterior ventral part

of the sepion inner cone. In the largest female (ML 91

mm) the eggs from the ovary are very elongate (8.9 x

3.2 mm).

Distribution: Sepia inccrta was recorded off eastern

Cape Province and Natal, East London to Durban (Massy,

1925). Sepions were also found from Port Elizabeth to

Southern Mozambique (Roeieveld, 1972). We found the

animals from North to South Mozambiciue, 1S°29' to

25''2rS, at depths of 90-345 m, significantly deeper than

previous records for this species (70-79 m; Roeieveld,

1972).

3. Sepia (Doratosepion) confusa Smith, 1916

(Figs. 5, 6, 20)

Material examined: "Professor Mesyatzev", 06/01 / 1976,

sta. 25, bottom trawl, 335 m, 05°19'S, 39°09'E: 3 females,

ML 64, 69, 69 mm; "Professor Mesyatzev". 22/01/1976,

sta. 57, bottom trawl, 150-170 m. 21°27'S, 35°32'E: 1

male, ML 70 mm; "Professor Mesyatzev ', 24/01/1976,

sta. 64, bottom trawl, 167-170 m, 24°23'S, 35°29'E: 5

males, ML 91-148 mm; ""Professor Mesvatzev ', 25/01/

1976, sta. 65, bottom trawl, 256-258 m, 24°43'S, 35°26'E:

5 males, ML 62-117 mm; "Professor Mesyatzev ", 25/

01/1976, sta. 66, bottom trawl, 200-210'm, 24°57'S,

35°07'E: 90 males, ML 48-135 mm; ""Professor Mesy-

atzev", 25/01/1976, sta. 67, bottom trawl, 56-53 m,

24°51'S, 35°05'E: 1 male, ML 120 mm, some of the spec-

imens are in the collection of ZMMU, Nos. 206, 215.

""Vityaz", 25/11/1988, sta. 2635, bottom trawl, 225-228

m, 25°05'-25°06'S, 35°15'E: 5 females ML 63-68 mm,

30 males 75-115 mm; some of the specimens are in the

collection of lOAS, (there were nearly 600 specimens of

both sexes of this species in this catch, but males greatly

outnumbered females).

Description and remarks: Sepia confusa males differ

from all other species in having a long "tail" (Adam &

Rees, 1966; Roeieveld, 1972). The length of this structure.

formed by fins fused or not fused posteriorly, depends

on the stage of maturity. Mature males have long tails

that are absent in immature males and in females. The

tail first appears at the 2nd maturity stage, and begins

to grow exponentially at ML about 90 mm, at which size

males change from the 3rd to the 4th maturity stage. In

females, the dorsal arms have expanded membranes and

two rows of suckers distally as in S. joubini. The lateral

arms have the same two rows of suckers distally, but the

protective membranes do not expand. Dorso-lateral arms

are not attenuated and are subequal in length. The sepion

differs from other species in having a dorsal posterior

keel and a ventral margin of the outer cone that is in-

verted to the outer side, triangular and thickened me-

dially.

Egg size in a female (ML 72 mm) was 5.0-5,1 x 2.6-

2.7 mm.

Distribution: Sepia confusa occurs off eastern and

southeastern Africa (5° to 30°S). The northernmost re-

cords are off Zanzibar (Adam & Rees, 1966) and Pemba

(our data). We found it to be very common along the

coast of Mozambique, 21°30'-25°S. The record of this

species from the Arabian Sea (Zuev, 1971), is erroneous,

while a record from Saya-de-Malha Bank (Nesis, 1982)

was not confirmed by more recent expeditions (Nesis,

1993). This species was recorded at depths of 53-352 m

(Roeieveld, 1972; our data).

4. Sepia {Doratosepion) burnupi Hoyle, 1904

(=Sepia exsignata Barnard, 1962)

(Fig. 20)

Material examined: ""Professor Mesvatzev", 23/01/1976,

sta. 62, bottom trawl, 235-240 m,' 23°29'S, 35°33'E: 2

males, ML 54, 56 mm, ZMMU 221.

Description and remarks: This species was hitherto

known from three sepions and three specimens taken off

Natal (Massy, 1925; Barnard, 1962; Roeieveld, 1972). The

two males of S. Inirntipi in our collection are practically

identical with Roeieveld s (1972) description and figures

of this species.

Distribution: Sepia burnupi is a very rare species. Our

collections enlarge its known range to Southern Mozam-

bique, and known depth distribution from 40-48 m to

235-240 m.

Figure 8. Sepia (Anomalosepia) omani Adam, 1939 Posterior part of female sepion. Redrawn from Adam & Rees, 1966, pi. 22,

fig. 144. Figures 9-1 L Scpia (Rhomhosepion) acuminata Smith, 1916. 9. ML 84 mm. 05°19'S. 39°09'E, 335 m, ZMMU No. 204.

Atypical sepion of female with furrow 10-1 1. ML 87 mm, 24°25'S. 3.5°29'E. 170 m, VNIRO uncatalogued 10. Ventral and 11.

dorsal views of typical sepion of female with rib. Figures 12, 13. Sepia (Rhonthosepion) hieronis Robson, 1924 ML 62 mm,

04°47'S. 39°24'E, 180-185 m, ZMMU No. 227. 12. Ventral and 13. lateral views of sepion of a male. Figure 14. Sepia (Acanthosepion)

zanzilmrica Pfeffer, 1884. ML 92 mm, 12°22'S,54°28'E, 50 m, ZMMU No, 317, Sepion of male. Figure 15. Sepia (Acanthosepion)

prashadi Winckworth, 1936, ML 90 mm, 04°47'S, 39°24'E, 180-185 m, ZMMU No. 249, Sepion of male. Figures 16, 17. Sepiella

cyanea Robson, 1924. ML 60 mm, 17°13'S, 38°39'E, 14 m, ZMMU No. 207. 16. Ventral and 17. lateral views of sepion of male.

Figures 18, 19. Sepiella inermis (Ferussac & d'Orbigny, 1835). ML 66 mm, Kuwait territorial waters, 0-20 m, ZMMU No. 198.

18. Ventral and 19. lateral views of sepion of male. Scale bars = 1 cm. cm, chitinous margin of sepion; ds, dorsal shield; gr, grooves

beside median ridge; mr, median ridge (or rib).

Page 72

THE NAUTILUS, Vol. 108, No. 3

40°E

60°E

20°

I I I I I I I I -r-

r I I 1 I 1 II

20"

40-E

60°E

f'igure 20. Distribution of Sepia (Doratoscpion) in the western

Indian Ocean. Triangles = Sepia (Doratosepiun) jouhini, cir-

cles = S. (D.) incerta, crosses = S. (D.)co7ifusa, stiuare = S.(D.)

burnupt.

5. Sepia (Doratosepion) trygonina Rochebrune, 1884

(Figs. 7, 21)

Material examined: "Ob", 08/06/1956, sta. 150, bot-

tom trawl, 50 m, 11°50'N, SPIO'E: 8 females, ML 53-

85 mm; "Ob", 10/06/1956, sta. 152, bottom trawl, 30

m, 12°05'N, 44°12'E: 1 male, ML 48 mm, 1 female, ML

36 mm; "Professor Mesyatzev", 19/12/1975, sta. 2, bot-

tom trawl, 67-77 m, 2°'24'S, 40°56'E: 2 males, ML 110,

41 mm; "Yelsk", December 1977 - January 1978, Saya-

de-Malha Bank, 205 m, some sepions; "Professor Mesy-

atzev", 27/03/1983, sta. 14, bottom trawl, 200 m, 1 r30'S,

60°49'E: 2 females, ML 46, 53 mm; "Akademik Petrov-

sky", 20/01/1984, sta. 67, Sigsbee trawl, 55 m, 15°07'N,

41°41'E: 1 female, ML 28 mm; "Odissey", 27/05/1984,

sta. 9, bottom trawl, 28 m, 12°19'N, 53°22'E: 2 males,

ML 51, 59 mm, 8 females, ML 51-74 mm; "Odissey",

27/05/1984, sta. 10, bottom trawl, 27 m, 12°14'N, 53°46'E:

1 female, ML 31 mm; "Odissey", 19/04/1985, sta. 95,

Sigsbee trawl, 80 m, 12°21'N, 54°28'E: 1 male, ML 30

mm; "Odissey", 19/04/1985, sta. 96, bottom trawl, 100

m, 12°19'N, 54°25'E: 2 males, ML 47-60 mm; "Skif",

01/11/1986, sta. 140, bottom trawl, 20 m, 15°35'N,

52°32'E: I male, ML 47 mm, 1 female, ML 45 mm, 1

juv, ML 25 mm; "Gidrobiolog", 01/05/1987, sta. 2, bot-

tom trawl, 50 m, 13°37'N, 47°23'E: 3 femaie.s, ML 47,

49, 61 mm, 5 juv, ML 30-20 mm; "Gidrobiolog", 16/

05/1987, sta. 6, bottom trawl, 33 m, 14°47'N, 50°01'E:

2 females, ML 52, 62 mm; "Gidrobiolog", 29/05/1987,

sta. 10, bottom trawl, 50 m, 13°55'N, 48°29'E: 5 females,

ML 52-64 mm; "(iidrobiolog", 10/05/1987, sta. 18, bot-

tom trawl, 37-40 m, 11°56'N, 53°44'E: 2 females, ML

71, 74 mm; Gidrobiolog", 28/05/1987, sta. 18, bottom

trawl, 40 m, I3°59'N, 47°57'E: 6 females, ML 37-63

mm, collections of VNIRO and ZMMU, Nos. 184, 197,

304, 310, 314, 316, 319, 324, 331.

Remarks: Sepions of S. trygonina are very similar to

those of . Imn^iipi, but the soft bodies of these species

are distinctly different. InS. trygonina, tliearmsof males

are not transformed as in S.burnupi and the lateral arms

of females have two widely spaced rows of suckers in

the distal portions.

Distribution: Scpia trygonina was first described from

the Red Sea and later reported from different areas of

the northwestern Indian Ocean: from the Gulf of Aden

and Kuria-Muria Islands to the Gulf of Mannar (Adam

& Rees, 1966; Druzhinin & Filippova, 1971; Sarvesan,

1976). We found this species south of the ecjuator on the

Saya-de-Maiha Bank, at 15°35'N to 1 1°30'S, at depths of

20 to 205 m. Mantle length reaches 110 mm in males,

85 mm in females, and is much larger than reported

previously.

6. Sepia (Doratosepion) niascaren.sis Filippova and

Khromov, 1991

(Fig. 21)

Material examined: "Professor Mesyatzev", 04/04/1976,

sta. 128, bottom trawl, Kargados-Karajos Islands: 1 male,

ML 105 mm; "Professor Mesyatzev" 05/04/1976, sta.

129, bottom trawl, 316-325 m,'l5°45'S, 61°15'E: 1 male,

ML 90 mm; "Professor Mesvatzev", 27/03/1983, sta. 14,

bottom trawl, 200 m, 11°36'S, 60°49'E: 20 males ML ?

(in poor state); "Professor Mesyatzev", 27/03/1983, sta.

15, bottom trawl, 195 m, 11°31'S, 6r00'E: 15 males, ML

67-124 mm; "Yunaya smena", 1985, Saya-de-Malha

Bank: dry and broken sepions, collections of VNIRO and

ZMMU, Nos. 233, 234, 349, 350. "Vitvaz", 07/01/1989,

sta. 2803, bottom trawl, 87-110 m, 11°21'-11°22'S,

61°47'E: 7 females, ML 50-63 mm, lOAS.

Description and remarks: This species is fully described

from males (Filijipova & Kliromov, 1991) and females

(Khromov et al.. 1991 ). Our specimens agree completely

with these descriptions. The males of ii. mascarensis are

characterized by their untransformed arms, the absence

of a hectocotylus, and a narrow and relatively thick se-

pion that has a rib on its ventral side and reversed,

W-shaped, ventral striae. The females of this species have

long second arms with two rows of suckers distally and

narrow sepions with reversed, W-shaped ventral striae

as well as a furrow (not a rib) on the ventral side.

Distribution: Scpia mascarensis is reported from the

Mascarenc Ridge, the Sa\a-de-Mailia Bank and shoals

around the Kargados-Karajos Islands, at depths of 87-

325 m.

7. Sepia (Dorato.sepiim) arahica Ma.ssv, 1916

(Fig. 21)

Material examined: "Skif", 01/1 1/1986, sta 10, bottom

Yu. A. Filippova et al., 199.5

Page 73

trawl, 20 m, 15°.35'N, 52°32'E: 1 male, ML 88 mm. col-

lection of VNIRO.

Description and remarks: This specimen agrees with

the description and illustrations of Adam and Rees (1966).

This mature male with a developed hectocotylus is the

largest recorded specimen of this species.

Distribution: Red Sea, Gulf of Aden to southern India.

8. Sepia (Doratoscpion) sp.

(Fig. 21)

Material examined: "Vitvaz", 07/01/1989, sta. 2804,

bottom trawl, 230-235 m, ll°06'-ir09'S, 62°14'-62°13'E:

1 female ML 65 mm, lOAS.

Description and remarks: Mantle elongate oval, ante-

rior margin somewhat produced dorsally, emarginate

ventrally. Arms subequal in length, lateral arms slightly

longer. Arms pairs 1, II, and III with well developed,

spade-like protective membranes covering the suckers

on the distal third of arms. The suckers are cjuadriserial

proximally, becoming biserial and widely spaced distalK .

The fourth arms have quadriserial suckers. Tentacular

club with 5 rows of suckers, differing slightly in size,

with the suckers of the central row the largest. Sepion is

cream colored, narrow, elongate, with a rib, delimited

by furrows dorsally and a median groove ventrally, and

slightly W-shaped in cross-section. The striated zone oc-

cupies the entire width of the ventral side, with the

anterior striae semi-rounded, slightly emarginated in the

ventral groove. The inner cone forms a ledge that is

slightly raised posteriorly, but not expanded. The outer

cone is cup-like and calcified, forming a toothed structure

on the ventral margin similar to that of S. incerta (Roe-

leveld, 1972, Fig.6,d). The spine is long, lacks a base and

spindle-like laterally.

This female is similar to one of S. conjusa. but the

sepion differs distinctly by the semi-rounded anterior

striae, the absence of an inverted, triangle-shaped ventral

margin on the inner cone, and a posterior dorsal keel on

the sepion. It also resembles S. joubini, but the arms of

the latter species are markedly longer and attenuated at

their ends. Moreover, the striae of S. joubini are reversed

V-shaped, its sepion is semi-rounded in cross-section, its

inner cone is slightly expanded posteriorly, and its outer

cone is decalcified. The sepion of Sepia sp. can be easily

confused with that of S. incerta, but it has no groove on

the posterior portion of the inner cone.

We could not identify Sepia sp. in spite of the good

condition of the specimen. Possibly it is a new species,

but the description of a new species in so complex a

subgenus as Doratosepion would not be appropriate in

the absence of a male specimen.

9. Sepia (Anomalosepia) omani Adam and Rees, 1966

(Figs. 8, 22)

Material examined: "Akademik Knipovich", 19/01/

1966, sta. 310, bottom trawl, 210 m, 23°42'N, 66°14'E:

2 males, ML 78, 80 mm, 1 female, ML 85 mm, 3 juv,

ML 30, 31, 35 mm, ZMMU 214.

Figure 21. Distribution of Sepia (Doratosepion) in the western

Indian Ocean. Circles = S. {D.) trygonina, triangles = S. (D.j

mascarensis, square = S. (D.) arabica, cross = S.(D.) sp.

Description and remarks: Sepia omani is the only spe-

cies with two pairs of outer cone wings on the sepion.

Thus, the specimens referred to S. omani by Voss and

Williamson, 1971 are certainly not correctly identified.

The opinion (Khromov, 1988b) that these animals are S.

rex is also erroneous. The Hong Kong specimens probably

belong to a new species.

Distribution: Sepia omani was known only from the

Gulf of Oman, at a depth of 201 m. We found it in the

northern part of the Arabian Sea, off Pakistan, at nearly

the same depth, 210 m.

10. Sepia (Rhombosepion) acuminata Smith, 1916

(Figs. 9-11, 22)

Material examined: "Professor Mesyatzev", 19/12/1975,

sta. 1, bottom trawl, off North Kenya: 1 male, ML 93

mm, 1 female, ML 75 mm; "Professor Mesyatzev ', 21/

12/1975, sta. 3, bottom trawl, 290-295 m, 3°02'S, 40°26'E:

1 male, ML 84 mm, 1 broken sepion; "Professor Mesy-

atzev", 04/01/1976, sta. 23, bottom trawl, 44 m, 04°03'S,

40°00'E: 1 male, ML 73 mm, 1 female, ML 75 mm;

"Professor Mesyatzev", 06/01/1976, sta. 25, bottom trawl,

p.... . - 1

THE NAUTILUS, Vol. 108, No. 3

Figure 22. Distribution of Sepia (Anomalosepia) and Sepia

(Rhombosepion) in the western Indian Ocean Triangle = Sepia

(Anomalosepia) omani, circles = Sepia (Rhombosepion) acu-

minata, squares = S.(R.) hieronis, cross = S. (R.) elegans.

335 m. 05°19'S, 39°09'E: 1 male, ML 102 mm, 3 females,

ML 56, 100, 111 mm; "Professor Mesyatzev", 24/01/

1976, sta. 64, bottom trawl, 170 m, 24°25'S, 35°29'E: 1

sepion 84 mm; "Zheleznyakov ', 07/03/1977, bottom

trawl, 230-235 m, 0r31'N', 44°39'E: 1 male, ML 55 mm,

2 broken sepions, collections of VNIRO and ZMMU, Nos.

191, 192, 204, 225, 242. "Vityaz", sta. 2635, 25/11/1988,

bottom trawl, 225-228 m, 25°05'- 25°06'S, 35°15'E: 11

females, ML 84-119 mm, 14 males, ML 52-91 mm, 1

juv ML 31 mm, collection of lOAS, 66 unsexed and

unmeasured specimens of the same species; "Vityaz' ,

sta. 2644, 02/12/1988, bottom trawl, 325-332 m, 22°19'-

22''23'S, 43°06'E: 4 females ML 89-115 mm. I sepion

110 nmi, collection of lOAS.

Description and remarks: We note an interesting vari-

ability of sepions in this species. Typical sepions of S.

acuminata caught in South African waters (Port Eliza-

beth) show a distinct median rib and a faint indication

of two lateral ribs on the dorsal surface (Adam & Bees,

1966). These authors also reported on three ailditional

sepions from Mombasa that had a deep, narrow groove

instead of a rib Rocleveld (1972) described specimens

from Mozambic]ue that had a median ridge sometimes

sunken below the level of rest of dorsal surface.

Having examined our material, we noted a clear rib

on the sepions of specimens from southern Mozambique,

a flat dorsal side or one slightly sunken below the rib on

sepions from northern Mozambique and Tanzania, and

a groove on the sepions of animals from Kenya and

southern Somalia. Females of S. acuminata from south-

ern Madagascar have a flat dorsal surface on the sepion.

Small males look alike, but the largest male has a sepion

with a slightly marked groove. We therefore suggest that

the presence of a rib or groove may be a geographical

variation that is more evident in large animals.

Distribution: Sepia acuminata has not been previously

reported from off Madagascar, northern Kenya, or So-

malia. This species has a wide range along all eastern

Africa, from 0r30'N to 30°S, including Madagascar. It

was recorded at depths of 44-369 ni (Roeleveld, 1972;

our data). Maximum size is presently a mantle length of

up to 100 mm in males and 120 mm in females. Egg

size in a female of 110 mm ML was 5.0-5.2 x 4. 5-4. 7

mm, while in smaller females it ranged between 4.2-4.9

X 3.3-4.5 mm. Females mature at about 85-90 mm ML.

11. Sepia (Rhombosepion) hieronis Robson, 1924

(Fig. 12, 13, 22)

Material examined: "Professor Mesyatzev", 05/01/1976,

sta. 24, bottom trawl, 180-185 m, 04°47'S, 39°24'E: 1

male, ML 62 mm; "Professor Mesyatzev", 27/01/1976,

sta. 73, bottom trawl, 430-440 m, 25°28'S, 33°32'E: 2

males, ML 42, 48 mm, ZMMU 227, 228.

Description and remarks: Our specimens agree com-

pletely with the descriptions of Adam and Rees (1966)

and Roeleveld (1972).

Distribution: Sepia hieronis has been reported in the

south-eastern Atlantic, from northern Namibia to Slang-

kop and in the western Indian Ocean off southern Mo-

zambique and on the Agulhas Bank (Voss, 1962, 1967;

Roeleveld, 1972; Okutani & Ilasegava, 1979; Sanchez &

Moli, 1984; Sanchez, 1988; Sanchez & Villanueva, 1988).

We collected it off Mozambique and Kenya, so S. hi-

eronis is quite widely distributed south ot Angola and

Zanzibar, but is not reported off South .\frica between

the Cape of Good Hope and St. Francis Ba\. The depth

range is 43-457 m (Roeleveld, 1972).

12. Sepia (Rhombosepion) elegans d'Orbigny, 1826

(Fig. 22)

Material examined: "Ob", 18/03/1957, sta. 263, bot-

tom trawl, 110 m, 37°12'S, 22°30'E: 4 males, ML 39-45

mm; "Gizhiga", 15/02/1975, sta. 266, bottom trawl, 180-

200 m, 21°10'S, 13°20'E: 1 female, ML59 mm; "Fiolent",

.sta. 136, 12/03/1976, 250 m, 06''26'S, 1 1°36'E: 4 females,

ML 73-89 mm; "Odissey" 06/12/1980, bottom trawl,

200 m, 31°25'N, 2riO'\V: 1 male, ML 55 nmi; "Aka-

demik Knipovich", sta. 8, 05/01/1967, 180-210 m. West

Africa: I male, ML 50 mm; "Akademik Knipnvich", 07/

Yu. A. Filippova et al., 1995

Page 75

06/1969, 80-95 m, 21°50'N, 17°20'W; 1 male, ML 50, 1

female, ML 53 mm; "Akademik Knipovich", sta. 1, 30/

01/1984, 140-200 m, 23°46'N, 16°51'W; 3 females, ML

36, 39, 46 mm, collections of VNIRO and ZMMU, Nos.

186. 193 222, 226, 307.

Description and remarks: Our specimens agree com-

pletely with the description of Adam and Rees (1966).

Distribution: Sepia clegans is a widely distributed east-

ern Atlantic species, reported along the coasts of Europe

and West Africa from Ireland to Namibia (Adam & Rees,

1966; Sanchez & Moli, 1984; Sanchez, 1988). We re-

corded mature specimens from the Agulhas Bank, so the

range of S. elegans extends into the southwestern Indian

Ocean.

13. Sepia (Sepia) papillata Quoy and Gaimard, 1832

(Fig. 23)

Material examined: "Professor Mesyatzev", 06/04/1976,

sta. 131, bottom trawl, 180-210 m. 15°48'S, 59°58'E: 1

male, ML 123 mm, 1 female, ML 107 mm; "Odissey",

12/06/1984, sta. 14, bottom trawl, 35 m, 09°42'S, 61°08'E:

2 males, ML 50, 64 mm, ZMMU 254, 330; "Vitvaz", 08/

01/1988, sta. 2808, bottom trawl, 58-61 m,' 11°05'S,

62°02'-62°04'E: 5 males ML 37-58 mm, 10 females ML

28-51 mm; "Vityaz", 08/01/1988, sta. 2810, bottom

trawl, 57-70 m, 10°15'-10°16'S, 61°09'E: 2 males, ML

63, 65 mm, 1 female, ML 75 mm, lOAS.

Description and remarks: This species differs from the

previous ones in having a broadly oval sepion with a

deep and wide median ventral groove and a considerably

narrowed and flat inner cone According to Adam and

Rees (1966), there is no spine, but only a blunt knob on

the sepion posterior. Roeleveld (1972) described a short

spine not exceeding the posterior margins of the shell.

Our specimens from Saya-de-MaIha Bank are not typical,

as some have a short spine visible from the ventral side

of the sepion. Thus, the length of spine of S. papillata

is variable.

Living animals of S. papillata from "Vityaz" station

2808 were very brightly colored, scarlet and gold, with-

out stripes or spots, while specimens from station 2810

were brown with zebra-like stripes and red spots. The

color of the animals probably reflects the color of the

substrate (crustose red algae at sta. 2808). Fixed animals

are not brightly colored and can be easily confused with

S. o. vermiculata and S. simoniana. Females mature at

about 50 mm ML.

Distribution: This species was previously reported along

the coast of South Africa from Luederitz Ba\ (Atlantic)

to the Tugela River (Indian Ocean), and especially near

Cape Town (Roeleveld, 1972; Okutani & Hasegawa,

1979). As we have caught it far from this region, on the

Mascarene Ridge, the range of S. papillata is greatly ex-

tended. The depth range for our samples is 35-210 m,

somewhat deeper than 26-127 m range recorded pre-

viously by Roeleveld (1972).

Figure 23. Distribution of Sepia (Sepia) in the western Indian

Ocean. Triangles = Sepia (Sepia) papillata, squares = S. (S)

simoniana, circles = S. (S.) officinalis vermiculata

14. Sepia (Sepia) airnoniana Thiele, 1921

(Fig. 23)

Material examined: "Professor Mesyatzev", 05/01/1976,

sta. 24, bottom trawl, 180-185 m, 04°47'S, 39°24'E: 1

male, ML 91 mm. 1 female, ML 95 mm; "Professor

Mesyatzev". 24/01/1976, sta. 64, bottom trawl, 170 m,

24°23'S, 35°29'E: 5 females, ML 58-99 mm. ZMMU 200,

250.

Description and remarks: Sepia simoniana is closely

related to S. papillata, but differs in having equal-sized

suckers on the long club, and a long spine on the sepion.

Distribution: This species was first described from Si-

mons Bay. South Africa, and repeatedly reported from

the same region, near Cape Town and along the east

coast of South Africa to the Tugela River (Roeleveld,

1972). Two specimens were caught on the Agulhas Bank

(Adam, 1983). Our material extends the range of S. si-

moniana to the area trom northern Kenya to southern

Mozambique, at depths 170-185 m (14-134 m reported

by Roeleveld, 1972).

15. Sepia (Sepia) officinalis vermiculata Quoy and Gai-

mard, 1832

(Fig. 23)

Material examined:

sta. 46, bottom trawl

"Professor Mesyatzev", 19/01/1976,

30-40 m, 19°17"S, 36°22'E: 2 males,

ML 82, 87 mm, 1 female, ML 82 mm; "Professor Mes-

yatzev", 20/01/1976, sta. 47, bottom trawl, 70-75 m,

i9°45'S, 36''22'E: 1 male, ML 98 mm; "Professor Mes-

yatzev", 20/01/1976, sta. 50, bottom trawl, 30 m, 19°37'S,

35°43'E: 4 females, ML 87-133 mm; "Professor Mesy-

atzev", 26/01/1976, sta. 68, bottom trawl, 44-50 m,

24°54'S, 34°53'E: 1 sepion 93 mm; "Professor Mesy-

Page 76

THE NAUTILUS, Vol. 108, No. 3

Figure 24. Distribution of Sepia (Acanthosepion) and Sepia

(Hemisepius) in the western Indian Ocean. Circles = Sepia

(Acanthosepion) zanzibarica, squares = S. (A.) prashadi, cross

= S. (A.) savignyi. triangle = S. (H.) typica.

atzev", 13/04/1976, sta. 148, bottom trawl, 25 m, 09°52'S,

60°57'E; 2 males, ML 64, 69 mm, collections of VNIRO

and ZMMU, Nos. 224, 251, 252, 253, ■'Vityaz", 25/11/

1988, sta. 2634a, bottom trawl, 90-92 m, 25°05'S, 34°50'-

34°44'E: 1 male, ML 100 mm, 1 female, ML 135 mm;

"Vityaz", 25/11/1988, sta. 2634b, bottom trawl, 95-102

m, 25°05'- 25°06'S, 34°45'E: 1 female, ML 1 18, collection

of lOAS.

Description and remarks: Our specimens agree com-

pletely with the description of Adam ands Rees (1966).

Distribution: Sepia officinalis vcrmictilata has been

found along southern Africa from 30°S in the southeast-

ern Atlantic Ocean to southern Mozambique in the south-

western Indian Ocean (Voss, 1962; Adam & Ree.s, 1966;

Roeleveld, 1972) and reported off Namibia (Sanchez &

Moll, 1981; Sanchez, 1988). According to our data, S. o.

vermiculata ranges northward to central Mozambique,

19°S and on the Saya-de-Malha Bank at depths of 25-

102 m. The largest of our specimens are immature (male

100 mm ML and females 1 18 anil 135 mm ML were in

the second stage of maturity).

Figure 25. Distribution of Sepiella in the western Indian Ocean.

Triangles = Sepiella cyanea. circles = S. inermis.

16. Sepia (Acanthosepion) zanzibarica Pfeffer, 1884

(Fig. 14, 24)

Material examined: "Odissey", 27/05/1984, sta. 9, bot-

tom trawl, 28 m, 12°19'N, 53°22'E: 1 female, ML 104

mm; "Odissey", 12/06/1984, sta. 14, bottom trawl, 35

m, 09°42'S, 61°08'E: 1 female, ML 58 mm; "Odissey",

17/04/1985, sta. 95, bottom trawl, 50 m, 12°22'N, 54°28'E:

9 males, ML 71-139 mm, 4 females, ML 88-115 mm;

"Odissey", 19/04/1985, sta. 95, bottom trawl, 40 m,

12°16'N, 53°59'E: 1 male, ML 172 mm; "Odissey", 17/

04/1985, sta. 94, Iwttom trawl, 40 m, 12°22'N, 54°21'E:

1 female, ML 73 mm, collections of VNIRO and ZMMU,

Nos. 317, 326. "Vitvaz", 04/12/1988, sta. 2657, bottom

trawl, 40 m, 22°10'- 22°15'S, 43°09'-4.3°10'E: 1 female,

ML 204 mm; "Vityaz", 08/01/1988, sta. 2810, shrimp

trawl, 57-70 m, l6°16'-10°15'S, 61°09'E: 6 males, ML

57-122 mm, 2 females, ML 108, 178 mm, 1 juv ML 41

mm; Toliara (Madagascar), beach: 1 sepion 118 mm,

collection of lOAS.

Description and remarks: Our specimens agree \\itli

the description of Adam and Rees (1966).

Distributi<m: Sepia zanzibarica was formerly known

Yu A Filippova et al., 1995

Page 77

from sepions found from Natal to Kenya and Madagascar

and by two specimens with soft parts (Adam & Rees,

1966). According to L. S Homenko (personal commu-

nication), S. zanziharica isoneol the common cuttlefishes

of the northwestern Indian Ocean and Saya-de-Malha

Bank, at depths of about 20-125 m. Our data confirm

this conmiunication. Sepia zanziharica is common in the

open regions of the Indian Ocean, near islands (Socotra,

Madagascar) and banks (Saya-de-Malha).

17. Sepia (.-\canthoscpion) prasliadi Winckworth, 1936

(Fig. 15, 24)

Material examined: "Professor Mesyatzev", 05/01/1976,

sta. 24, bottom trawl, 180-185 m, 04°47'S, 39°24'E: 1

male, ML 90 mm, 2 females, ML 80, 65 mm; "Akademik

Knipovich", 13/01/1966, sta. 305, bottom trawl, 55 m,

17°41'N, 56°47'E: 3 males, ML 33, 35, 40 mm, collection

of VNIRO and ZMMU, No. 249 "Vityaz", 25/11/1988,

sta. 2634a, bottom trawl, 90-92 m,' 25°05'S, 34°50'-

34''44'E: 2 males, ML 50, 53 mm, collection of lOAS.

Description and remarks: Our specimens agree with

the description of Adam and Rees (1966).

Distribution: Sepia prashadi is a common northern and

western Indian Ocean species, recorded from the Gulf

of Suez to the Bay of Bengal and southward to southern

Mozambique, Madagascar and the Mauritius. Depth re-

cords span the entire continental shelf.

18. Sepia (Acanthosepion) savignyi Blainville, 1827

(Fig. 24)

Material examined: "Odissey", 27/05/1984, sta. 10,

bottom trawl, 27 m, 12°14'N, 53°46'E: 3 males, ML 94,

104, 108 mm, 2 females, ML 83, 90 mm, collection of

VNIRO.

Description and remarks: Sepia savignyi can be con-

fused with ii. plathyconchalis Filippova and Khromov,

1991, from which it differs in having a ventral median

groove on the sepion and a normal striated zone, which

is very narrow in S. plathyconchalis

Distribution: Sepia savignyi is known from the Red

Sea, the Gulf of Aden and the Persian Gulf, but was

never recorded south of Socotra Island. This species is

replaced by S. plathyconchalis off the coast of East Af-

rica.

19. Sepia (Heniisepius) typica Steenstrup, 1875

(Fig. 24)

Material e.xamined: "Vityaz", 22/11/1988, sta. 2626,

Sigsbee trawl, 290 m, 24°39'S, 35°31'E: 3 males, ML 9,

11, 12 mm, 6 females, ML 5-8 mm, collection of lOAS.

Description and remarks: Sepia typica is the type spe-

cies of the subgenus Hemisepius, and distinctly differs

from the other species of Heniisepius in having ventral

mantle pores (12 on each side) (Khromov, 1987). Sepia

typica is a rather rare species, known from several spec-

imens caught from Saldahna Bay to Cape Natal (Roe-

leveld, 1972). It was suggested that there is a difference

between western (Atlantic) and eastern (Indian Ocean)

forms of the species, and the latter was named S.(H.)

typica var. chuni (Thore, 1945). Roeleveld (1972:264)

questioned the validity of these forms and wrote: "A

decision must await the collection of further specimens

from the eastern coast of South Africa". We think such

a decision cannot be final in the absence of a special

population stud\ of S. typica.

We confirm Thore s report of the differences in size

between western and eastern forms. The latter (Massy,

1927 and our material) are much smaller (males at 5th

stage of maturity are only 11-12 mm); in fact, our spec-

imens are the smallest mature Heniisepius ever reported.

We cannot distinguish any morphological differences

other than size. Our specimens have a variable number

of enlarged suckers in the 7- 10th rows of suckers on the

dorso-lateral arms. Usually two pairs of suckers occur on

the first-second arms and one pair on the third arms. The

suckers of the right ventral arm decrease in size distally

except for the suckers in approximately the 12th row,

which are enlarged. On the hectocotylus there are 8-9

pairs of widely spaced reduced biserial suckers (starting

from the proximal pair) and about 10-14 minute suckers

distally. These characters are the same as in the Atlantic

form as described by Roeleveld (1972).

It is interesting that the eyelids of our specimens look

very similar to those of S. (D.) bathyalis (Khromov et

al., 1991) with two narrow triangular outgrowths on the

upper margins. The fins of our specimens extend forward

from the anterior margin of the mantle. They are widest

in the middle part of the mantle, not in the posterior

area as in other species of Sepia. The sepion occupies

only the anterior half of the mantle, as is characteristic

of the subgenus Hemisepius. On the other hand, the

sepions of our specimens though very thin and fragile,

are calcified, which has not previously been reported.

We believe that the decalcification of sepions in previ-

ously reported specimens of this species (Adam & Rees,

1966; Roeleveld, 1972; etc. ) is an artifact due to improper

fixation. This supports our opinion that S. (H.) robsoni.

S. (H.)faurei. S. (H.) dubia and S. (H.) pulchra belong

to the subgenus Hemisepius of the genus Sepia because

the decalcification of the sepion per se cannot be con-

sidered as a character of the subgenus (Khromov, 1987).

The phragmocone, inner and outer cones are present but

strongly reduced in our specimens

Distribution: Sepia (H.) typica was not previousK re-

corded north of Cape Natal in the Indian Ocean. We

found this species off southern Mozambique far north of

the known range and at a much greater depth (290 m

versus 2-156 m) (Roeleveld, 1972). The other four species

of this subgeims are known oidy from the area off the

Cape of Good Hope at depths of 15-168 m.

20. Sepiella cyanea Robson, 1924

(Figs. 16, 17, 25)

Material examined: "Professor Mesyatzev", 16/01/1976,

sta. 33, bottom trawl, 14 m, 17°13'S, 38°39'E: 2 males,

ML 36, 60 mm, 2 females, ML 30, 42 mm; 'Akademik

Page 78

THE NAUTILUS, Vol. 108, No. 3

Knipovich", 29/09/1977, sta. 375, bottom trawl, 20-25

m, 18°33'S, 36°48'E: 2 males, ML 54, 67 mm, 1 female,

ML 68 mm, ZMMU 187, 207. Sepiella, which can be

only S. cyanea were also noted by B. G. Ivanov (log book)

off Mozambique between 17°37' and 18°17' S at the depths

of 17-40 m. "Vitvaz", 19/11/1988, sta. 2620, bottom

trawl, 13-14 m, 17°1 l'-17°12'S, 38°34'-38°30'E: 14 males,

ML 23-50 mm, 16 females, ML 40-65 mm, lOAS.

Description and remarks: Our specimens agree with

the descriptions of Adam and Rees (1966), and Roeleveld

(1972). Egg size in a female of 65 mm ML ("'Vityaz' )

was 3-5 mm.

Distribution: Sepiella ctjanea has been reported in South

African waters from Durban to Port Elizabeth and off

Madagascar (Roeleveld, 1972). We found it near the coast

of Central Mozambique at depths of 13-40 m, shallower

than previously reported (51-73 m) (Roeleveld, 1972).

21. Sepiella inermis (Ferussac and d'Orbigny, 1835)

(Figs. 18, 19, 25)

Material examined: "Nauka", 30/03/1976, sta. 61,

Hindustan: 3 females, ML 54, 55, 56 mm; "Nauka", 06/

04/1976, sta. 98, Gulf of Kutch: 3 males, ML 27, 33, 42

mm, 1 female, ML 51 mm, 1 juv ML 23 mm; "Akademik

Knipovich", 21/01/1966, sta. 317, bottom trawl, 20-25

m, 22°55'N, 68°22'E: 2 juv, ML 22, 35 mm; "Akademik

Knipovich", 09/03/1966, sta. 436, bottom trawl, 40 m,

14°42'N, 97''26'E: 2 females, ML 57, 60 mm: "Sabah",

March 1979, 0-20 m, Kuwait territorial waters: 8 males,

ML 51-78 mm. 22 females, ML 51-105 mm, collections

of VNIRO and ZMMU, Nos. 198, 201, 311.

Description and remarks: According to Adam and Rees

(1966), the number of sucker rows on the tentacular club

of S. inermis is very variable, ranging from 13 to 24

depending on sex and geographical area. Moreover, we

must consider the possibility of changes in sucker number

during club regeneration. For example, one of our spec-

imens (female, ML 83 mm) has one normal club with

about 18 rows of suckers and another (regenerated?) with

only 9 rows of suckers. In this case S. inermis can hardly

be separated from S. cijanea, because the latter species

is characterized by a similar sepion and 12 rows of club

suckers.

Distribution: Sepiella inermis is a wide ranging species

inhabiting the nortfiern Indian Ocean and adjacent west-

ern Pacific from the Persian Gulf and Red Sea to the

Gulf of Tonkin and Indonesia.

DISCUSSION

The analysis of material gathered by the Soviet research

vessels in African waters and the Mascarene Ridge region

between 24°N and 37°S sigiiihcantly broadens our knowl-

edge of the .sepiid fauna ol this poorly known area. Sev-

eral species, S. (A.) zanzibarica, S. (D.) trygonina, S. (S.)

officinalis vermiculata, S, (S.) papillata, were caught for

the first time in the open waters of the Mascarene Ridge,

which are separated from Africa and Madagascar by a

large area of deep water, the Amirante Trench and the

Mascarene Basin. Three new species of Sepia, S. (D.)

mascarensis, S. (S.) plathyconchalis, and S. (D.) saya,

described in previous papers (Filippova & Khromov, 1991;

Khromovefa/., 1991) are present in this area. The species

list of this region may expand further, because we have

in our collections several poorly preserved specimens,

including at least two more new species.

This survey also revealed the occurrence of several

species near the coast of Madagascar: S. (R.) acuminata

in this paper and S. (D.) tala and S. (D.) hathyalis re-

ported in Khromov et al., (1991).

The distribution ranges of many South African sepiids

are broader than previously believed. Sepia (D.) joubini

and S.(D.)burniipi, known previously from Natal, are

recorded from southern Mozambique and S. (D.) incerta

from northern Mozambique. Sepia (S.)simoniana, which

was considered endemic to Cape Province, as well as S.

(R.) hieronis occur along the East African coast to north-

ern Kenya, while S.(R.) acuminata ranges to Somalia

(and off Madagascar). The Cape-Natal species S. (H.)

typica was found off northern Mozambique. Thus the

endemism of the South African fauna was hitherto great-

ly exaggerated (Roeleveld, 1972) because of a poor

knowledge of the East African offshore and open sea

faunas.

It is very significant that the southern and eastern

African sepiid faunas, as well as the entire fauna of the

Western Indian Ocean, are represented mainly by the

species of subgenus Doratosepion. it was noted (Nesis,

1980; Khromov, 1987) that the subtropical waters off

South Africa were one of the centers of species diversi-

fication of Doratosepion. In summarizing the data pre-

sented in this paper and in previous ones (Khromov, 1982,

1988; Filippova & Khromov, 1991; Khromov et al., 1991),

there are 14-16 species of Doratosepion in the Western

Indian Ocean. This subgenus comprises more than half

of the Sepia species in this area. Sepia s. str. is the second,

and Acanthosepion the third most diverse subgenus in

the region.

An intriguing and puzzling fact is that, despite the

large number of trawls, no cuttlefish were recorded from

the Seychelles Islands or the Seychelles Plateau, an an-

cient granitic microcontinent. This is in striking contrast

with the great richness and diversity of the cuttlefish

fauna of Saya-de-Malha Bank and Madagascar, two other

ancient areas with continental t\pe crust (Nur & Ben-

Avraham, 1982), which have not been in contact with

the African continent or Indian subcontinent since at

least the Late Cretaceous (Kennett, 1982). The geological

history of the Seychelles Islands and the Mascarene Ridge

is obviously very different (Kennett, 1982). We believe

that the distribution of sepiids was governed by dispersal

rather than by vicariant events.

LITERATURE CITED

Adam, W 1959 LesGephalopodcsde la mer Rouge. Resuitats

Yu. A. Filipfxiva et al., 1995

Page 79

scientifiques Mission R. P. Dollfus en Egypte (dec 1927 -

mars 1929)3(28)125-193.

Adam, W. 1979. The Sepiidae (Cephalopoda, Decapoda) in

the collection of the Western Australian Museum. Records

of the Western Australian Mu.seiun 7(I2);109-212.

Adam, W, 1983. Ck'phalopoda from West and South Africa

Atlantide Reports 13151-180

Adam, W and W J Rees. 1966. A review of the cephalnpotl

famil) Sepiidae Scientific Reports of the John Murra\

E.xpedition 1933-34 ll(l):l-165.

Barnard, K. H. 1962. New species and records of South .Af-

rican marine Mollusca from Natal, Zululand and Mozam-

bique. Annals of the Natal Museum 15:2-47-254.

Druzhinin, A. D. and Yu. A. Filippova 1974. Distribution

and fishery of cuttlefish off the southeastern coast of the

Arabian Peninsula Proceedings of the All-Union Research

Institute of Marine Fisheries and Oceanography (VNIRO)

99: 117-122. [Russian].

Filippova, Yu A. and D. N. Khromov 1991. New data on

the sepiid fauna (Sepiidae, Cephalopoda) of the Western

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sian, English summary].

Homenko, L. P. 1976. Some data on the biology and distri-

bution of Sepia pharaonis Ehrenberg and S. hrevimana

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Homenko, L. P. 1983. Distribution of cephalopods off the

western coast of Hindustan. In: Ya. I. Starobogatov and

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Khromov, D. N. 1982. A new species of the genus Sepia

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Khromov, D. N. 1987. Systematics and phylogeny of the

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Khromov, D. N. 1988a. Two new species of the genus Sepia

(Cephalopoda, Sepiida) from the Socotra Island (People s

Democratic Republic of Yemen) waters. Zoologichesky

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Khromov, D. N 1988b. Cuttlefishes of the family Sepiidae

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Institute, Systematics and fauna of Gastropoda, Bivalvia

and Cephalopoda 187:174-195. [Russian, English sum-

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Khromov, D. N., 1. V. Nikitina and K. N. Nesis. 1991. Three

new species of cuttlefish Sepia (Doratosepion) from the

western Indian Ocean, with new data on S. (D.) mascar-

ensis Filippova et Khromov (Ophalopoda, Sepiidae).

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mary].

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Annals of the Natal Museum 5201-229.

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the World Ocean Legkaya i Pishchevaya Promyshlennost.

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Indian Ocean Transactions of the P. P. Shirsov Institute

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42 [Russian].

THE NAUTILUS 108(3):80-82, 1995

Page 80

Three Additional New Genera and two Replacement Names for

Northeastern Pacific Prosobranch Gastropods

James H. McLean

Los Angeles County Museum of

Natural History

900 Exposition Blvd

Los Angeles, CA 90007 USA

ABSTRACT

Three new genera or subgenera and two replacement names

for species fiomonyms are proposed. New genus level taxa;

Trochidae: Costomargarites. new subgenus of MargariJfs; type

species Trochus costalis Gould. 1841. Calyptraeidae: Grandi-

crepidula, new subgenus of Crepidula. type species Crepidula

grandis Middendorff, 1849. Turridae, Crassispirinae: Pseudo-

taranis new genus; type species Mangelia (Tarants) strongi

Arnold, 1903. New names for homonyms: Trochidae: Marga-

rites (Costomargarites) baxteri, new name for Margarites

(Pupillaria) rudis Dall, 1919, not Margarita groenlandica var.

rudis Morch, 1869 Turridae: Leucosyrinx hantori, new name

for Antiplanes aniycus Dall, 1919, not Leucosyrinx amycus

Dall, 1919.

Key words: Mollusca; Prosobranch gastropods; Trochidae; Ca-

lyptraeidae; Turridae; new species; new names

INTRODUCTION

This paper follows a previous paper (McLean, 1995) with

the same objective — to make some generic level names

available for other work in progress. Additionally, the

opportunity is taken to provide replacement names for

two homonvms.

NEW GENUS LEVEL TAXA

Family TROCHIDAE Rafinesque, 1815

Subfamily MARGARITINAE Stoliczka, 1868

Costomargarites McLean, new subgenus

Type species: Trochus costalis (iould, 1841.

Included species: Type species [Arctic, North .Atlantic

and North Pacific] and Trochus ochotensis Philippi, 184(i

[Northwestern Pacific]. The complex synonymy for the

type species was outlined by Abbott (1974;36). The ju-

venile shell was illustrated by Pilsbry (1889, pi. 60, figs

27, 28).

Diagnosis: Subgenus of Margarites in which early tc-

leoconch .sculpture is axial rather than spiral.

Remarks: Sculpture of typical species of Margarites and

species of other available subgenera of Margarites is

spiral, to the exclusion of axial sculpture altogether except

for growth increments. The strong axial sculpture of ju-

venile shells of Margarites (Costomargarites) costalis

suggests a closer resemblance to Solariclla obscure

(Couthouy, 1838) than to other species of Margarites.

The species in which axial sculpture dominates the mor-

phology of the early teleoconch (and may or may not be

expressed in the mature shell) need to be distinguished

at least at the subgeneric level. For many years I have

been surprised that this common Arctic species has not

been made the type species of a subgenus, a gap that I

hereby fill.

Family CALYPTRAEIDAE Lamarck, 1809

Grandicrepidula McLean, new subgenus

Type species: Crepidula grandis Middendorff, 1849.

Included species: Type species [Boreal Pacific]; Crepi-

dula princeps Conrad, 1856 [early Miocene through

middle Pliocene of southern California]; C. excavate

(Broderip, 1834) [Panamic],C. maculosa Conrad, 1846

[Western Atlantic]; C. convexa Say, 1822 [Western At-

lantic].

Diagnosis: Beak excavated, projecting posteriorly on right

side; septal margin nearly straight, extending farther for-

ward on left side; single muscle scar on right side, rect-

angular or chevron-shaped.

Remarks: Hoagland (1977) reviewed the Recent and

fossil species of Crepidula. treating them in alphabetical

order, while not using the available subgenera. Surpris-

ingly, none of the 14 available taxa (see Hoagland, 1977:

399) has a type species in which the septum projects on

the right siiie. It is evident that this condition is not

ambiguous — all species illustrated b\ Hoagland can be

assigned on the basis ol this character. These diagnostic

characters were mentioned and show clearly in Hoag-

land's drawings of septal configuration (figure 2B, C.

excavate: figure 2F, C. gratidis). Moreover, the five spe-

cies assigned above key out together in Hoaglaml s key

J. H. McLean, 1995

Page 81

to the species (page 363, couplets 16-18). Contrary to

Hoagland s assertion that the division of Crepidula into

subgenera is not warranted, I find it useful to base su-

praspecific distinctions on the morphology of the septum,

particular!} when considering fossil species. In my opin-

ion the remaining species can also be assigned to the

available taxa on shell characters.

Family TURRIDAE Swainson, 1840

Subfamily CRASSISPIRINAE Morrison, 1966

Pseudotaranis McLean, new genus

Type species: Mangelia (Taranis) strongi Arnold, 1903.

Included species: Type species and the more slender

Antiplanes hijperia Dail, 1919a (p. 35, pi. 9, fig. 6). The

type species was described from the Lower Pleistocene

of San Pedro, California; it was also illustrated by Grant

& Gale (1931:572, pi. 26. fig. 37); Borsonia inculta Mood) ,

1919 (p. 54, pi. 1, figs. 2a, 2b) is a s>non\ni. The two

species (Pseitdotaranis strongi and P. hyperia) live off-

shore in moderately deep water (100-400 m).

Diagnosis: Shell small (length to 17 nun), spire high,

anterior canal short, whorls 6. Axial sculpture lacking,

spiral sculpture of two cords emerging on teleoconch and

three cords on base; anal sinus shallow, at periphery and

coinciding with uppermost cord; lip not projecting. Pro-

toconch paucispiral, of 1.2 low, rounded whorls. Radula

of marginal teeth attached to membrane; teeth of long,

flat type.

Remarks: Authors prior to 1971 used Taranis Jeffreys,

1870 for the type species, but the presence of a radula

precludes placement in that genus, in which the radula

is lacking (see Powell, 1966:55). Earlier (McLean, 1971;

120, fig 40), I illustrated the radula of strongi and as-

signed it to Antiplanes (Rectisulciis), a genus now in the

subfamily Cochlespirinae, according to Ta\lor et al.

(1993). Pseudotaranis differs in having the sinus more

shallow, the lip less protracted, the anterior canal shorter,

and the protoconch more compressed. The radula lacks

the vestigial rachidian tooth and the marginals are not

of the bifurcated type indicated for Antiplanes by Kan-

tor and Sysoev (1991:122). The flat morphology of the

lateral teeth agrees with the subfamily Crassispirinae, as

defined by Taylor et al. (1993). Thus, the radular dif-

ferences place Antiplanes and Pseudotaranis in different

turrid subfamilies. However, on shell characters, Pseu-

dotaranis is atypical of Crassispirinae in not having pro-

nounced callus developed near the anal notch.

NEW NAMES FOR SPECIES LEVEL HOMONYMS

Family TROCHIDAE Rafinesque, 1815

Subfamily MARGARITINAE Stoliczka, 1868

Margatites (Costomargarites) baxteri McLean, new-

name for Margarites (Pupillaria) rudis Dall, 1919b:364,

not Margarita groenlandica var. rudis Morch, 1869:23.

Remarks: Margarites (Costomargarites) baxteri was fig-

ured by Dall (1921:179, pi. 18, fig. 13, 14) and by Kosuge

(1972, pi. 2, fig. 5), but has otherwise been ignored in

the literature. It will be treated by me in work in prep-

aration as a geographic subspecies (in the Gulf of Alaska)

of the boreal M. (Costomargarites) costalis (Gould, 1841).

The new name honors the late Rae Baxter for his efforts

at reviewing Alaskan mollusks, cut short by his untimely

death in 1991.

Family TURRIDAE Swainson, 1840

Subfamily COCHLESPIRINAE Powell, 1942

Leucosyrinx kantori McLean, new name for Antiplanes

anujeus Dall, 1919a (p. 36, pi. 11, fig. 5), not Leucosyrinx

amyctis Dall, 1919a (p. 5, pi. 3, fig. 7).

Remarks: The proposal of Leucosyrinx kantori remedies

a problem of secondary homonymy that was initiated

when Dall (1919a) described a species in Antiplanes that

I now consider to be a true member of Leucosyrinx Dall,

1889. Dall (1919a) also proposed a species with the name

Leucosyrinx amycus. Note that both species were de-

scribed in the same paper. Previously (McLean in Keen,

1971:713), I placed Leucosyrinx amycus Dall, 1919 in

the synonymy of Aforia goodei (Dall, 1890). The new

name honors Yuri Kantor for his recent work on turri-

form gastropods.

LITERATURE CITED

Abbott, R. T. 1974. American Seashells. Second Edition. New

York: Van Nostrand Reinhold, 663 pp.

Dall, W. H. 1919a. Descriptions of new species of mollusks

of the family Turritidae from the west coast of America

and adjacent regions. Proceedings of the United States

National Museum 56(2288): 1-86, pis. 1-24.

Dall, W. H 1919b Description of new species of Mollusca

from the North Pacific Ocean in the collection of the

United States National Museum. Proceedings of the L'nited

States National Museum 56(2295):29;3-371.

Dall, W H, 1921. Summary of the marine shell-bearing mol-

lusks of the northwest coast of America, from San Diego,

California, to the Polar Sea, mostly contained in the col-

lection of the United States National Museum, with illus-

trations of hitherto unfigured species. United States Na-

tional Museum, Bulletin 112, 217 pp., 22 pis.

Grant, U. S, IV and H. R Gale 1931. Catalogue of the marine

Pliocene and Pleistocene Mollusca of California and ad-

jacent regions. Memoirs of the San Diego Society of Nat-

ural History 1:1-1036, pls.1-32.

Hoagland, K. E. 1977. Systematic review of the fossil and

Recent Crepidula and discussion of evolution of the Ca-

lyptraeidae. Malacologia 16(2):353-420.

Kantor, Y. I, and A. V. Sysoev. 1991. Mollusks of the genus

Antiplanes (Gastropoda: Turridae) of the northwestern

Pacific Ocean. The Nautilus 105(4):1 19-146

Keen, A, M. 1971. Sea shells of tropical west America. Second

Edition. Stanford Universitv Press, Stanford, 1064 pp.,

22 pis.

Kosuge, S. 1972. Illustrations of type specimens of Molluscs

described by William Healey Dall. National Science Mu-

seum, Tokyo, 29 pis. and captions.

THE NAUTILUS 108(3):82, 1995

Page 82

McLean, J H. 1971. A revised classification of the family

Turridae, wilh the proposal of new subfamilies, genera,

and subgenera from the Eastern Pacific. The Veliger 14(1):

11-4-130.

McLean, J. H. 1995. Four new genera for northeastern Pacific

prosobranch gastropods. The Nautilus 108(2):39-41.

Moody, C L. 1916. Fauna of the Fernando of Los Angeles.

University of California Publications, Bulletin, Depart-

ment of Geology 10(-4):39-62, pis. 1-2.

Morch, 0. A. L. 1869 Catalogue des mollusqnes du Spitzberg

recueillis par le Dr. H Kroyer pendant le voyage de la

corvette La Recherche en juin 1838. Memoires de la So-

ciete Malacologique de Belgique 4:7-32

Pilsbry, H. A. 1889-90. Manual of Conchology. Vol. 11 Tro-

chidae. Philadelphia, 519 pp., 67 pis,

Powell, A. W. B. 1966. The molluscan families Speightiidae

and Turridae. Bulletin of the Auckland Institute and Mu-

seum, no. 5, 184 pp., 23 pis.

Taylor, J. D., Y. \. Kantor, and A. V'. Sysoev 1993. Foregut

anatomy, feeding mechanisms, relationships and classifi-

cation of the Conoidea ( = Toxoglossa) (Gastropoda). Bul-

letin of the Natural History Museum London (Zoology)

59(2): 125- 170.

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TH EtyNAUTI LUS

Volume 108, Number 4

May 11, 1995

ISSN 0028-1344

CONTENTS

Bruce A. Marshall A Revision of the Recent Calliostoma Species of

New Zealand (Mollusca: Gastropoda; Trochoidea) 83

THE NAUTILUS 108(4):83-127, 1995

Page 83

A Revision of the Recent Calliostoma Species of New Zealand

(Mollusca: Gastropoda: Trochoidea)

Bruce A. IVIarshall

Museum of New Zealand Te Papa

Tongarewa

P.O Box 467

Wellington, New Zealand

ABSTRACT

Thirty-three Recent calliostomatids are recorded from New

Zealand, ten of which are described as new Calliostoma (Otu-

haia) blacki (Dell, 1956) is renamed due to homonymy, and

the following taxa are newly synonymized: Venustas tigris

chat liamensis Dell, 1950 with Calliostoma (Maurea) tigris

(Gmelin, 1791); Maurea punctulata ampla Powell, 1939 and

V. punctulata multigemmata Powell, 1952 with C. granti

(Powell, 1931); V. cuuperi Vella, 1954 and Thoristclla chath-

amensis profunda Dell, 1956 with C. blacki (Powell, 1950),

which is resurrected from synonymy under C. joveauxanum

(Dell, 1950); Zizijphinus hodgci Hutton, 1875, Z. ponderosus

Hutton, 1885, and C carnicolor Preston, f907 with C. sctectum

(Dillwyn, 1817); C. undulatum Finlay, 1923 and C. pellucidum

spiratum Oliver, 1926 with C. pellucidum (Valenciennes, 1846).

Lectotypes are designated for Trochus selectus Dillwyn, 1817,

T. pellucidus Valenciennes, 1846, and Calliostoma onustum

Odhner, 1924.

Key words: Mollusca; prosobranch gastropods, Trochoidea;

Calliostomatidae; Calliostoma, systematics.

INTRODUCTION

The family Calliostomatidae comprises about 250 living

species. They occur in all oceans from the intertidal zone

to about 3000 meters depth, mostly on rocky ground. All

known species are carnivores, most feeding on cnidaria,

and sometimes carion, though a few feed exclusively on

sponges. The group is particidarK well represented in

the New Zealand region, with 33 living endemic species,

including some of the worlds largest, notably Callios-

toma (Maurea) tigris (Gmelin, 1791), which may e.xceed

100mm in shell height.

Besides the 33 Recent species recorded herein, at least

double this number are known from the New Zealand

Cenozoic (Early Eocene — Early Pleistocene), of which

more than half are undescribed. Several of the living

species have excellent fossil records in the extensive Plio-

Pleistocene deposits of the southern North Island, and

fossil material is recorded and discussed where appro-

priate.

Two of the recorded Recent species, both of which

are type species of new genera, together with a new

genus for Calliostoma onustum Odhner, 1924 are named

elsewhere (Marshall, 1995). Five additional species are

known from off the Kermadec Islands (Appendix).

Attention is drawn to the fact that calliostoinatid shell

morphology tends to become more variable with increas-

ing size/age, so that species with dissimilar early teleo-

conchs can be superficially similar at maturity and vice

versa. For accurate discrimination of species (and for

objective descriptions), it is thus essential to trace and

compare the development of individual sculptural ele-

ments from the earliest teleoconch whorls following the

system used by Ikebe (1942) and Marshall (1988, 1995)

(i^igure 28).

ABBREVIATIONS AND TEXT CONVENTIONS

AUG Geology Department, Auckland University

BMNH The Natural History Museum, London

MNHN Museum National d'Histoire Naturelle, Paris

MNZ Museum of New Zealand, Wellington

NZGS Institute of Geological and Nuclear Sciences,

Lower Hutt

NZOI National Institute of Water and Atmospheric

Research, Wellington

sa spire angle (see below)

ZMA Zoological Museum, Amsterdam

Spire angle measurements for individual shells were av-

eraged (mean spire angle) to reduce bias induced by

cyrtoeonoid spire profile and expanded or narrowed last

adult whorl. In other words, for a shell with a cyrtoeonoid

spire outline, which becomes more narrowly conical with

increasing shell size, the given spire angle is the mean

of the maximum and minimum spire angles measured

from that specimen. Vice versa for a shell with an evenly

conical or coeloconoid contour in which the last adult

whorl expands more rapidly than the previous ones. In

illustrations of shells, height dimension precetles diam-

eter.

Readers requiring more detailed information on New

Zealand stratigraphy should refer to Fleming (1953), Beu

and Maxwell (1990), and Abbott and Carter (1994).

Page 84

THE NAUTILUS, Vol. 108, No. 4

SYSTEMATICS

Order Vetigastropoda Salvini-Plawen, 1980

Siiperfamily Trochoidea Rafinesqiie, 1815

Family Calliosiomalidae Thiele, 1924

Genus Calliostoma Swainson, 1840

Calliostoma Swainson, 1840:218, 351. Type species (by sub-

sequent designation of Herrmannsen, 1846:154): Trochus

conulus Linnaeus, 1758; Recent, north-eastern Atlantic

and Mediterranean For further discussion and synonymy,

see Marshall (1995).

Subgenus Maurea Oliver, 1926

Maiirea Oliver, 1926:108. Type species (by original designa-

tion): Trochus tigris Gmelin, 1791; Recent, New Zealand

[20th December 1926-see below].

Maitriella Oliver, 1926:109. Type species (by original desig-

nation): Trochus punctulatus Martyn, 1784; Recent, New

Zealand [20th December 1926].

Calliotropis Oliver, 1926:110 Type species (by original des-

ignation): Trochus cunninghami Gray, 1834 = Trochus

selectus Dillwyn, 1817; Recent, New Zealand Not Cal-

liotropis Seguenza, 1903 [20th December 1926]

Mucrinops Finlay, 1926:360. Type species (by original desig-

nation): Ziziphinus spectabilis A. Adams, 1855); Recent,

New Zealand [23 December 1926].

Venustas Finlay, 1927:360. Type species (by original desig-

nation): Trochus tigris Gmelin, 1791; Recent, New Zea-

land [10 March 1927]. Officially rejected name (ICZN

Opinion 479). Not Venustas Allan, 1926.

Calotropis Thiele, 1929:49. Replacement name for Calliotropis

Oliver not Seguenza

Remarks: As indicated by Ben et al. (1969), Venustas

Allan, 1926 (Allan, 1926) was published on 7th December

1926 (ICZN opinion 479) and has priority over Maurea

Oliver, 1926 (and the officially rejected name Venustas

Finlay, 1927). Fortunately the type species of Venustas

Allan (Calliostoma fragile Finlay, 1923; Early Miocene,

New Zealand) seems unlikely to be consubgeneric with

Calliostoma {Maurea) tigris (Gmelin, 1791), the type

species of Maurea. The paper in which Oliver (1926)

introduced Maurea was published in Parts II / III of

Volume 17 of the Proceedings of the Malacological So-

ciety of London, the date of publication of which was

interpreted as 20th December 1926 by Dell (ICZN Opin-

ion 479) on the basis of a letter from L. R. Cox to M. K

Mestayer bound into the MNZ volume of the journal.

This letter is dated 19th April 1927. The title page to

Volume 17, issued on 30th December 1927, states that

Parts II and III were issued on 30th December 1926.

Whereas I have been unable to trace any firmer evidence

that Parts II and 111 were published prior to 30th De-

cember, for the sake of noinenclatural stability it is ap-

propriate to follow ICZN acceptance (right or wrong) of

20th December 1926, otherwise Maurea would fall as a

junior synonym of Mucrinops Finlay, 1926 (23 Decem-

ber). Maurea is not endangered by Venusta.s Finlay,

which has been deemed to have been published on lOth

March 1927 and officially rejected (ICZN Opinion 479).

The type species of Maurea, Calotropis, Mauriella,

and Mucrinops are similar to the type species of Cal-

liostoma in radular morphology and external anatomy.

Although I am unable to justify genus-level status for

Maurea. it would be inappropriate to treat it as a syn-

onym of Calliostoma because all of the New Zealand

species are strongly dissimilar to the type species of Cal-

liostoma in shell morphology. On the other hand, there

is strong mosaic overlap between the type species of

Maurea. Calotropis, Mauriella, and Mucrinops via the

other Recent species herein referred to Maurea, and I

am unable to justify segregation of these genus-group

taxa from each other. Accordingly, they are all inter-

preted as synonyms of Maurea, which in turn is inter-

preted as a subgenus of Calliostoma. Although the origins

and relationships of the Recent species are obscure, it is

nevertheless clear that Maurea as here limited is poly-

phyletic. In whatever way the Recent species are ar-

ranged in groups, most or all contain species of excep-

tionally large size for the family. With the exception of

"Trochus" mutus Finlay, 1924 and " Benthastelena" su-

sanae Maxwell, 1992 (Late Eocene — Early Miocene),

which are not closely related to any taxa living in the

New Zealand region, none of the numerous pre-Pliocene

species from New Zealand exceeds much more than about

20 mm in maximum shell dimension, so it would seem

that gigantism occurred independently in several species

groups after the Miocene. Many of the pre-Pliocene spe-

cies are similar to the type species of Fautor Iredale,

1924 (Ziziphinus compttis A. Adams, 1854; Recent,

southern Australia) and other species referred there (as

a subgenus of Calliostoma) by Marshall (1995) from the

New Caledonia area. Calliostoma regale new species and

C. aupourianum new species from northern New Zea-

land are similar to these small-shelled New Caledonian

Fautor species and to many of the New Zealand pre-

Pliocene taxa, and because (independently derived) gi-

gantism alone is no criterion for genus-group discrimi-

nation, it is thus difficult to justif\ segregation of Maurea

even from the prior Fautor. EvidentK highlv conser-

vative external anatomy and gross shell and radular mor-

phology are inadequate to construct objectively definable

supraspecific groupings, or rather, real phylogenetic

groups are rendered nebulous and are obfuscated due to

conservation, convergence, gigantism, and uncertain

character-state polarity. Molecular cladistic techniques

would seem to be a promising source of data for reso-

lution of these problems. For more detailed discussion

see Marshall (1995).

Calliostoma (Maurea) tigris (Gmelin, 1791)

(Figures 1-9, 110, 127)

Chemnitz, 1781:100, pi 170, figs 1654, 16,55

Trochus tigris Martyn, 1784, fig 75, (Officially rejected name —

ICZN Opinion 479).

Trochus granatuvi Gmelin, 1791:3584 (refers to Chemnitz,

1781. pi. 170, figs. 1654, 1655); Lamarck, 1822:26; Fischer,

1875:69, pi 15, fig. 1.

Trochus tigris Gmelin, 1791:3585 (refers to Martyn, 1784, fig.

Bruce A. Marshall. 1995

Page 85

75) (Officially accepted name — ICZN Opinion 479); Plii-

lippi, 1848;50, pi. 10, figs. 16, 17.

Turbo granalum — Rixling, 1798:88.

Ziziphmus tigris — Gray, 1843:237.

Ziziiphmus ligris— Reeve, 1863, fig 4; Hutton, 1873:38.

Zizyphimis graimtum—HiMon. 1880:98; Hutton, 1884:359.

Cailiostoma gratxatum—?ihhry, 1888:313, pi. 41, fig. 30.

Calliostoma ( igris— Pilsbry, 1889:333; Suter, 1897:280; Suter,

1913:148, pi. 40, fig. 6.

Calliostoma (Maurea) (igrii— Oliver, 1926:108; Wenz, 1938:

282, fig. 600; Shikama, 1964:106, fig. 185.

Venustas (Venustas) tigris — Finlay, 1926:360, 371.

Maitrea tigris — Powell, 1937:64, pi. 1, fig. 12; Matsukunia,

Okutani & Habe, 1991, pi 17, fig 11.

Venustas tigris tigris— Dell. 1950:41, figs. 22, 23, 24.

Venustas tigris chathamensis — Dell, 1950:43, figs. 26, 27. New

synonym.

Maurea tigris tigris — Powell, 1957:88, pl.l, fig. 12; Powell,

1979:60, pi. 10, fig 1.

Maurea tigris chathamensis — Powell, 1957:88; Powell, 1979:

61, pi 19, fig 3

NOT Zizyphinus granatum — Reeve, 1863:pl.l, fig. 2 (C. pel-

lucidum)

Type Data: Trochus tigris: MarUn, 1784, fig. 75, "New

Zealand"; Trochus granatum: Chemnitz, 1781, pi. 170,

figs. 1654, 1655, "Neuseeland"; Venustas tigris chath-

amensis : Holotype MNZ M.2128, Mangare Island, Chat-

ham Islands.

Other IVIaterial Examined: Fossil — Boulder at head of

largest bend in Wainui Stock Road, Ohope (map ref.

W15/638502), B.A. Marshall, 1965 (late Castlecliffian,

Late Pleistocene) (1 MNZ); Banks of Ohinekoao Stream,

coastal clifl's, Matata, B.A. Marshall, 1969 (late Castle-

cliffian, Late Pleistocene) (1 MNZ). Recent — 478 speci-

mens in 198 lots MNZ, 13 specimens in 8 lots NZOI.

Distribution (figure 9): Mid-Pleistocene (late Castleclif-

fian) to Recent, off Three Kings, North, South, Stewart

and Chatham Islands, living at 0-211 m on rocky sub-

strata.

Diet: All guts examined contained thecate hydroids (Cni-

daria) and indeterminate tissue.

Remarks: This well-known species, the largest known

calliostomatid, is characterised by its large size, relatively

thin shell, coeloconoid spire, very narrowly conical and

distinctively sculptured early teleoconch (figure 108), and

by the (usual) colour pattern of yellowish or reddish

brown wavy axial bands. Sculptural development in nor-

mal specimens (see below) proceeds through a distinctive

intermediate stage in which the spiral cords become

weakly nodular or smooth and much broader than the

interspaces, then reverts to strongly nodular with wider

interspaces on the last adult whorl. Mature specimens

have a gently sloping shoulder on the last part of the last

whorl, a pronounced thickening within the outer lip, and

typically a slight abapical descent of the apertural rim.

Shells having adult characteristics may be as small as

27.5 mm in height (MNZ M. 84227, Cape Runaway),

although most are about twice this size. The species at-

tains exceptional size off the Three Kings Islands (height

up to 100 mm, figure 3).

Most specimens from Parengarenga Harbour (figure

6) and many specimens from off eastern Northland (fig-

ure 2), including Whangamata and the Aldermen Islands

in the northern Bay of Plenty, have appreciably narrower

spire angles than specimens from north of Cape Reinga,

from Hauraki Gulf, and from south of Mayor Island, Bay

of Plenty (sa = 61-72°, mean 66°, S.D. 3.26, n = 20 as

against sa = 68-87°, mean 77°, S.D. 3.91, n = 58). There

is complete intergradation between narrowly and broad-

ly conical forms, however, both within and between pop-

ulations, and there are no other differences between them.

Because narrowly conical forms occur only in the warm

waters off the north-eastern North Island, it would seem

that differences in spire angle are linked in some way to

sea temperature (see C. hlacki below). The occurrence

of broadly conical specimens from further north, off the

Three kings Islands, presumably reflects local cooling due

to upwelling (Marshall, 1981).

Most specimens seen from off the C^hatham Islands,

including the holotype of Venustas tigris chathamensis

Dell, 1950, differ from mainland specimens in lacking

discrete axial bands and are instead predominantly red-

dish or yellowish brown with scattered, irregular white

spots and streaks (figure 5). Some Chatham Islands shells

(e.g., NZOI Q23), however, have axial bands and are

essentially similar to specimens from off the South Island

and Stewart Island, which commonly have broader bands

than northern shells (figure 7). One specimen from the

Three Kings Islands (figure 4) is indistinguishable from

typical Chatham Islands shells in colour and in density

of colour pattern. Although Dell (1950) considered that

Chatham Islands specimens differed from mainland shells

in having stronger, more persistent nodules, examination

of much additional material reveals that sculpture is vari-

able and that the Chatham Islands form cannot be dis-

tinguished using this criterion. Accordingly I am unable

to justify continued recognition of V . tigris chathamensis

as a geographic subspecies. Unless C. tigris intermittently

reaches the Chatham Islands as drifting eggs or larvae

from the mainland, it may have reached there along the

summit of the Chatham Rise during periods of low sea

level, presumably during Pleistocene glaciations.

Three specimens are known from the north-eastern

North Island on which the spiral cords remain strong,

widely spaced, and nodular throughout (figures 6, 8).

The two from Paua, Parengarenga Harbour (figure 6)

are also more deeply pigmented and more densely pat-

terned than others living beside them. The third example,

from off Cape Karikari (figure 8), differs from all other

material examined in that the spiral cords are alternately

spotted yellowish brown and white. To some extent the

Parengarenga Harbour shells resemble C. pellucidum

(Valenciennes) in adult fades, whereas that of the Cape

Karikari specimen approaches those of both C. punctu-

latum (Martyn) and C. osbornei Powell, w ith all of which

they respectively occurred. Although we cannot entirely

preclude the possibility, they seem unlikely to be hybrids

Page 86

THE NAUTILUS, Vol. 108, No. 4

Bruce A. Marshall, 1995

Page 87

because all have the characteristic and highly stable early

teleoconch morphology of C. tigri.s (figure 110).

The epithet tigris is from the Latin tigris ("tiger",

third declension feminine) and is a noun in apposition

to the generic name Calliostoma, hence it is not declin-

able.

Calliostonia (Maurea) punctulatum (Mart\n,

(Figures 10-18, 28, 32, 128)

784)

Spengler, 1776: 152, pi. 5, figs. 2a, b.

Chemnitz, 1781:26, pi. 161, figs. 1520, 1521.

Trochus punctulatus Martvn, 1784, fig. 36; Philippi, 1855, pi.

15, fig. 7; Reeve, 1862, pi 16, figs. 95a-d.

Trochus diaphanus Gmelin, 1791:3580 (refers to Spengler, 1776:

152, pi. 5, figs. 2a, b; and C:hemnitz, 1781, pi 161, figs.

1520, 1521); Wood, 1825, pi. 29, fig. 99; Quoy & Gaimard,

1834:254, pi. 64, figs. 1-5; Gray, 1842, pi. 40*, figs. 1, la;

Philippi, 1846:8, pi, 2, figs, 5, 6; Fischer, 1873:43, Turbo

pi. 10, fig 2.

Turbo diaphaiius — Lamarck, 1822:45.

Ziziphinu.s punctulatus — Gray 1843:237.

Turbo grandineus Valenciennes, 1846, pi. 4, figs. 4, 4ab; Kiener,

1847, Turbo pi. 10, fig. 2.

lizyphinuspiinctidatus^Huiion, 1873:39; Hiitton, 1880:98 (in

part = C. granti): Hutton, 1882:165, pi 7, fig 11; Hutton,

1884:360.

Calliostoma punctulatum — Pilsbry, 1889:334, pi. 65, fig. 75;

Suter 1897:280; Suter, 1913:146, pi. 8, fig. 11 (in part =

C. granti), Buclcnill, 1924:32, pi. 3, fig. 4 (in part = C.

granti).

Calliostoma (Mauriclla) punctulatum punctulatum — Oliver,

1926:109.

Calliostoma (Mauriella) punctulatum stcwartianum Oliver,

1926:109, pi. 10, fig. 1.

Calliostoma (Mauriella) wanganuicum Oliver, 1926:109, pi.

10, fig. 2. INew synonym.

Venustas (Mucrinops) punctulata punciulata — Finiay. 1926:

361, 371.

Venustas (Mucrinops) punctulata urbanior P'inlay, 1926:361,

pi. 18, fig. 27; Cernohorsivy, 1972:243, fig. 17'.

Maurea (Mucrinops) punctulata punctulata — Powell, 1937:64,

pi. 13, fig. 3 (in part = C. granti)-

Calliostoma (Mauriella) punctulatum — Wenz, 1938:282, fig,

601.

Maurea (Mucrinops) punctulata urbanior — Powell, 1937:64

Venustas punctulata punctulata — Dell, 1950:46, fig. 19, 20,

Venustas punctulata urbanior — Deii, 1950:47.

Maurea punctulata punciulata — Powell, 1957:88, pi. 13, fig.

3 (in part = C. granti).

Maurea punctulata stewartiana — Powell, 1957:88.

Calliostoma (Maurea) punctulata stewartiana — Shikama, 1964,

pi. 59, fig. 6.

Figure 9. Map of New Zealand region showing distribution of

Calliostoma (Maurea) tigris. 200 and 1000 meter contours in-

dicated.

Maurea (Mauriella) punctulata punctulata — Fleming, 1966:

38.

Maurea (Mauriella) punctulata stewartiana — Fiemmg, 1966:

38.

Maurea punctulata— PoweW. 1976:84, pi. 20, fig. 3; Powell,

1979:62, pi. 10, fig. 4 (both in part = C. granti).

Calliostoma (sensulato) punctulatum — Beu & Maxwell, 1990:

404

Calliostoma (sensu lato) wanganuicum — Beu & Maxwell, 1990:

404.

NOT Venustas punctulata urbanior — Powell, 1955:55 (C.

granti).

NOT Maurea punctulata— DeW, 1956:46 (C. granti).

Figures 1-8. Calliostoma (Maurea) tigris (Gmelin, 1791). 1. Off Southwest Island, Three Kings Islands. 22-23 m, MNZ M. 75143

(65.5 X 65.0 mm). 2. Off Poor Knights Islands, 20-30 m, MNZ M. 75150 (56.2 x 45.7 mm). 3. Off Three Kings Islands, craypot,

MNZ M. 75191, natural size (96.0 x 85.3 mm). 4. Reef between Great Island and Farmer Rocks, Three Kings Islands, 33 m, MNZ

M. 84239 (54.0 x 52.3 mm). 5. Off Southeast Island, Chatham Islands, MNZ M. 118120 (54.0 x 51 7 mm) 6. Paua wharf,

Parengarenga Harbour, low tide, possibly hybridized with C. (M.) pellucidum (Valenciennes, 1846), MNZ Ml 18337 (57.0 x 45.0

mm). 7. Off Ruggedy Island, Stewart Island, 37 m, MNZ M. 18996 (700 x 68 0 mm) 8. Off Cape Karikari, 80-120 m, possibly

hybridised with C. (M.) punctidatum (Martyn, 1784) or C. (M.) osbornei Powell, 1926, MNZ M 87026 (.36.6 x ;52 8 mm).

Page 88

THE NAUTILUS, Vol. 108, No. 4

Figures 10-18. Calliostoma (Maurea) punctulatum (Martyn. 1784). 10. Houhora Heads, low tide, MNZ M.2133 (34.6 x 33.6

mm). II. Paua, Parengarenga Harbour, low tide, MNZ M. 80482 (32.8 x 29.0 mm). 12. Northern Pania Reef, Napier, 20m, MNZ

M.86724 (34 0 x 3,5.3 mm). 13, 14. Barretts Reef, Wellington Harbour, MNZ M 45062 (29.0 x ,30.4 mm, and 29.0 x 31.2 mm).

15. Simp.son's Rock, Hauraki Gulf, 15-18 m, MNZ M.89971 (31.0 x 29.0 mm). 16. Timaru breakwater, low tide, MNZ M 5324

(42.4 X 408 mm).17. Cook Strait, 256-274 m, MNZ M.54911 (32.6 x 38.6 nun). 18. Masons Bay, Stewart Island, beach, MNZ

M.7200 (51.5 X 47.3 mm).

Type Data: Trochufi puncttdatus: Martyn, 1784; fig. 36,

"New Zealand"; Truchus diaphanus. Spongier, 1776: pi.

5, figs. 2a, b, "Siidsee"; Turbo grandineus. Lectotype

(here selected) and 1 paralectotype MNHN, New Zea-

land; Calliostoma {Matiriella) punctulatum stewartian-

um. Holotype MNZ M.879, Stewart Island; Calliostoma

(Mauriclla) tvanganuicum: Holotype NZGS TM 4999,

mouth ot Okehu Stream, Nukumarii Beach, near Wang-

Bruce A. Marshall, 1995

Page 89

anui (Castiecliffian, Middle Pleistocene); Venustas (Mu-

crinops) punctulata urbanior. Holotype AIM 70824,

Foveau.x Strait, 37 m.

Other Material Examined: Fossil — 56 Early-Middle

Pleistocene (Castlecliffian) specimens in 22 lots AUG,

MNZ, NZGS; Recent— 1169 specimens in 268 lots MNZ.

Distribution (figure 32): Early Pleistocene to Recent.

North, South, and Stewart Islands, living intertidally to

274 m on hard substrata.

Diet: The intestinal tracts of most of the 22 specimens

examined (10 localities) contained thecate hydroids (Cni-

daria), sand, and sometimes soft tissue of unknown origin

(cnidarianP). Two of three specimens examined from

Cornwallis, Manukau Harbour contained thecate hy-

droids, soft tissue, and sand, whereas the intestinal tract

of the third animal was packed with calcareous octocoral

spicules.

Remarks: This-well known, common species is extremely

variable in shape, size, thickness, colour, colour pattern,

and sculpture (figures 10-18). Recent specimens, how-

ever, are readily distinguishable from closely related taxa

(see below) by features of the early teleoconch, notably

the pronounced whorl angulation at P2, the relatively

weak axial costae, and the slow enlargement of S3 (figure

28). Shells range in shape from narrowly to broadly con-

ical (height/diameter ratio 0.87-1.13; sa 65-96°, mean

84.7°, SD 5.83, n = 74). Specimens from Parengarenga

Harbour are the most narrowly conical, with spire angles

ranging from 65° to 72° (figure 11). Specimens taken

alive from Cook Strait at 256-274 m (figure 17), the

deepest record for the species, are the most broadly con-

ical (sa 86-96°). The latter specimens, and some from off

East Otago, Stewart Island, and Foveaux Strait, are un-

usual in having a callous-filled umbilical depression, the

umbilicus being completely invaded by the inner lip in

other Recent material. Mature specimens are usualK

characterised by contraction and descent of the last part

of the last whorl and range from 18 to 51 (est.) mm in

height, the smallest adult specimens examined occurring

off Cape Maria van Diemen, and the largest at Stewart

Island in shallow depths, mainly as beach shells (figure

18). Shell thickness varies by a factor of 2 or more. Thick-

er specimens tend to predominate in exposed situations,

and most are more darkly pigmented with fewer and

coarser spiral cords than shells from deeper water. Colour

ranges from dark reddish to pale yellowish brown, the

spirals darker, the nodules either predominantly white

or roughly alternating brown and white. Specimens from

the northern North Island tend to be more darkh pig-

mented than material from the southern South Island.

Spiral cords are multiplied by repeated intercalation of

secondaries and tertiaries at variable stages of growth

and range in number from 6 to 25 on the second-to-last

whorl of mature specimens There is a strong correlation

between the strength of sculpture and the number of

spiral cords on later whorls. Coarsely sculptured speci-

mens with strong primary and secondary spirals tend to

develop fewer secondary spirals, whereas more finely

sculptured specimens tend to develop more numerous

tertiary spirals that enlarge more rapidly to resemble the

adjacent spirals Coarsely sculptured specimens tend to

predominate in the northern North Island, finely sculp-

tured forms in the southern South Island and in deep-

living populations. A notable exception is the occurrence

of a pale, thin, finely sculptured form at low tide in

Manukau Harbour, presumably in response to some local

environmental factor. In most specimens from north of

Cape Egmont and north of East Cape, including Ma-

nukau Harbour, S3 commences about midway between

P3 and P4, whereas in most specimens from the south,

especially the southern South Island and Stewart Island,

S3 commences close beside P4. Although samples from

the southern North Island and northern South Island

often comprise one form or the other, both forms fre-

quently occur together and completely intergrade in sev-

eral large samples (e.g., MNZ M.32569, M.44051,

M. 45062, M.54911). Southern forms have been treated

as geographic subspecies [stewartiana Oliver, urbanior

Finlay) of the coarser nominate northern forms, but in

fact there is complete mosaic intergradation in all ex-

tremes of shell morphology both within and between

populations, and it is quite impossible to define regional

subspecies.

In the Wanganui basin, C. punctulatum first appears

in the Butlers Shell Conglomerate (early Castlecliffian,

Early Pleistocene) (AU1047, AUG; GS4l69, NZGS) (Tur-

ner & Kamp, 1990). An imperfectly preserved shell (MNZ

M. 95387) from an Early Pleistocene (late Nukumaruan)

horizon exposed in a cutting on White Rock Road, south

of Hautotara Bridge, Martinborough appears to repre-

sent this species. A worn, incomplete specimen from a

Middle Pliocene (Waipipian) horizon in the Waipara

Gorge (GS4946, NZGS) is also similar but cannot be iden-

tified with certainty. All specimens from the Butlers Shell

Conglomerate and from between it and the younger

Kaikokopu Shell Grit (GS4064, 4075. 4134, 4163, 4168,

NZGS) (late Castlecliffian, Early Pleistocene), including

the holotype of C. wanganuicum, differ from Recent

specimens in having stronger axial costae on the early

teleoconch and in the earlier appearance and more rapid

enlargement of S3 (appearing at shell diameter of 1.9-

3.4 mm, mean 2.6 mm, n = 11 as against 3.1-8.0 mm,

mean 4.72 mm, n = 24), which is highly conspicuous on

early whorls as a suprasutural cord close beside P4. In

this character, these specimens bear some resemblance

to the Waipipian-Recent species C. granti (Powell) but

differ in being more broadly conical and by having more

strongly convex whorls as in Recent specimens of C.

punctulatum. S3 appears midway between P3 and P4

in most Recent specimens of C. punctulatum, though as

already stated, its origination position may vary from

median to submedian in some populations. Specimens

from the Kupe Formation (late Castlecliffian, Middle

Pleistocene) and overlying formations (GS4041, 4052,

4120, 4121, 4122, 4175, 4186, NZGS) are morphologi-

cally intermediate between earlier and Recent forms.

Page 90

THE NAUTILUS, Vol. 108, No. 4

Moreover, the fossils are as variable in shape, thickness,

nodule size, and strength and luiinber of spiral cords as

in Recent material, and there is intergradation between

extremes within horizons. Accordingly it seems impos-

sible to justify recognition of C. wanganuicutn as a dis-

tinct species or chronosubspecies. Three specimens from

the Butlers Shell Conglomerate (GS4109, NZGS; AU1047,

AUG) are unusual in having the umbilicus wide open

instead of fully closed through invasion by the inner lip.

One of the two umbilicate specimens from GS4109 has

the umbilicus considerably narrower than in the other

and is thus intermediate between umbilicate and an-

omphalous forms from the same sample, which are oth-

erwise indistinguishable.

Calliostonia punctulatiim is notable for its absence

from the Chatham Islands and the subantarctic islands,

including The Snares, and all specimens hitherto so iden-

tified from there are C. granti (see below). Of yet greater

interest is its absence from the Three Kings Islands, some

60 km north of Cape Maria van Diemen, where there

are two superficially similar species with closer relation-

ship to C. granti (see below). Its absence from this island

group may be due to a locally unfavourable environment,

either at present or when sea levels were low enough for

the species potentially to have reached them, for example

during Pleistocene glaciations. As with other local cal-

liostomatids that show pronounced clinal variation, it is

likely that C. punctulatuni has a drifting larval stage of

short duration.

Type Data: Maurea granti: Holotype AIM 70449, Waihi

Beach, Hawera (Waipipian, Middle Pliocene); Maurea

punctulata ampla. Holotype AIM 70450, Masons Bay,

Stewart Island; Vcnustas punctulata multigemmata:

Holotype AIM 71 185, off East Otago, 91-128 m.

Other Material Examined: Fossil — GS4253, Upper Wai-

pipi Shellbed, Waverley Beach, S. Taranaki (Waipipian,

Middle Pliocene) (3 NZGS); GS4949, c. 200 m SE of

north end of Greenwood's Bridge, left bank, lower Wai-

para Gorge, map ref. N34/922867 (f6180) (Waipipian)

(1 NZGS); GS5237, 724 m N of Trig Turangatairoa, on

main Taihape/Waiouru Road, map ref. T21/429771

(f8503) (probably Waipipian; A.G. Beu, per.comm);

GS40I3, Tainui Shellbed, Wanganui (late Castlecliffian,

Late Pleistocene) (1 AUG); GS4025, Pinnacle Sand,

Wanganui (late Castlecliffian) (1 NZGS). Recent— 508

specimens in 124 lots MNZ, and 4 specimens in 2 lots

NZOI.

Distribution (figure 33): Middle Pliocene (Waipipian) to

Recent. North Island south of East Cape and Cape Eg-

mont, and South, Stewart, Snares, Auckland, Campbell,

and Chatham islands, living from low-tide level to 220

m on hard substrata.

Diet: From gut contents (pers.obs. ) and field observations

(M.H.B. O'Neill, pers. comm.), C. granti feeds princi-

pally on Cnidaria, including thecate hydroids and anem-

ones, as well as ascidians.

Calliostonia (Maurea) granti (Powell, 1931)

(Figures 19-27, 29, 33, 129)

Zizyphinus punrtulatus — Hutton, 1880:98 (in part not Martyn,

1784).

Calliostoma puncttilatum — Suter, 1913:146 (in part).

Maurea (Mucrinops) granti Powell, 1931:97, pi. 13, figs. 34,35.

Maurea (Mucrinops) punctulata punctulata — Powell, 1937:64

(in part)

Maurea (Mucrinops) punctulata ampla Powell, 1939:229, pi.

50, fig. 6. New §ynonyni

Venustas punctulata ampla — Dell, 1950:47

Venustas punctulata (?) n.subsp. Dell, 1950:47.

Venustas punctulata multigemmata Powell, 1952:173, pi. 35,

figs 2,3. New synonym

Venustas punctulata urhanior — Powell, 1955:55 (not Finlay,

1926).

Maurea punctulata — Dell, 1956:46 (not Martyn, 1784).

Maurea punctulata punctulata — Powell, 1957:88 (in part).

Maurea punctulata ampla — Powell, 1957:88.

Maurea punctulata multigemmata — Powell, 1957:88.

Maurea (Mauriella) granti — Fleming, 1966:38.

Maurea (Mauriella) osbornei — Fleming. 1966:38 (not Powell,

1926).

Maurea punctulata — Powell, 1976:84; Powell, 197962 (in part)

Maurea nmlligemmata—WmeW. 197681; Powell, 197962, pi

10, fig. 6, pi. 19, fig. 8.

Maurea hlackii [sic] — Horikoshi, 1989, pi. 4, fig. 12 (not Powell,

1950).

Calliostoma (sensu lato) granti^ncn & Maxwell, 1990:404

Remarks: I am unable to detect any constant differences

between Waipipian (Middle Pliocene) specimens of Cal-

liostoma granti and the Recent forms named Maurea

punctulata ampla and Venustas punctulata multigem-

mata. Maurea punctulata ampla was originally sepa-

rated from the sympatric(!) "subspecies " M. punctulata

urhanior Finlay, 1926 [i.e., C. punctulatiim Martyn,

1784) on the basis of the larger, more elevated shell, that

has 9-12 instead of 12 equally developed spiral cords.

Venustus punctulata multigemmata was separated from

M. punctulata ampla because of its more lightly built

shell and finer, more numerous spiral cords. Study of

many times the number of specimens available when

these taxa were proposed, however, reveals that M. punc-

tulata ampla and V. punctulata multigemmata are based

on forms of a single polymorphic species (i.e., C. granti)

that occurs sympatricalK with C. punctulatum from

East Cape southward to Stewart Island. It transpires that

neither M. punctulata ampla nor V. punctulata mul-

tigemmata can be distinguished from C. punctulatum

using the characters cited in the original descriptions.

Like C. punctulatum. C. granti has considerable vari-

ation in shell size and shape, both within populations and

clinally, and in the strongly correlated number and rel-

ative sizes of the spiral cords on the last two adult whorls.

Beach shells and specimens collected intertidally (figure

22) tend to be thick and darkly pigmented with few,

strong, strongly nodular spiral cords, whereas specimens

from deeper water (figures 20,23) tend to be thin and

Bruce A. Marshall, 1995

Page 91

Figures 19-27. Calhoatuina (Maurca) granti (Powell, 1931). 19. Boil Reel, Napier, 10-20 m, MNZ M. 84224 ( 32.0 x 28.5 mm).

20. Off Wanganui, 82 m, MNZ M. 50181 (21.4 x 22.0 mm). 21. Westhaven Inlet, low tide, MNZ M. 81,531 (38.4 x ,35.1 mm). 22.

Kaliurangi Point, north-west Nelson, beach, MNZ M. 23101 (28.0 x 27.4 mm). 23. Off Oamaru, c. 100 m, MNZ M. 102602 (44.6

X 41.4 mm), 24. Off Se>mour Island, Dusky Sound, 24 m, MNZ M. 80476 (36.0 x 34.8 mm). 25. Paterson Inlet, Stewart Island,

beach, MNZ M. 19119 (45.6 x 42.0 mm) 26. Point Munning, Chatham Islands, 12 m, MNZ M.111927 (41 0 x ,38.1 mm) 27. Off

Auckland Islands, 104 m, NZOI D80 (24 0 x 2.3 8 mm).

Page 92

THE NAUTILUS, Vol. 108, No. 4

lightK pigmented (or white), with finer, more finely nod-

ular, more numerous spiral cords: the former include the

type material of M. puuciulata atupla, the latter V.

punctulata rnultigemrnata and C granti. There is com-

plete mosaic intergradation between these extremes in

the Recent material.

Although C. granti and C. punctulatum are similar

in gross facies, there are nevertheless marked and con-

stant differences in the shape and sculpture of the early

teleoconch whorls (figures 28, 29). Calliostoma granti

differs from C. punctulatum in the following features:

(1) the more rapid development of S3, which enlarges

to resemble the adjacent primaries at least a full whorl

earlier; (2) the stronger a.xial riblets, which persist over

one or two additional whorls; (3) the more weakly convex

and more slowly expanding early teleoconch whorls {i.e.,

narrower early spire angle); and (4) by comparison with

C. punctulata from within its range, the early teleoconch

is evenly conical in shape instead of coeloconoid, al-

though in allopatric C. punctulatum from the far north

of its range, the spire is typically evenly but yet more

broadly conical. Adult teleoconch whorls are typically

more weakly convex than in C. punctulatum, although

they tend to become as strongly convex in deep-living

populations (figures 20, 23). The sides of the foot are

considerably more finely pustulose in C. granti than in

C. punctulatum.

The holotype of C. granti (Powell, 1931, pi. 13, figs.

34, 35) is more narrowly and evenly conical than any

known Recent specimen (sa 67°, instead of 70-87°, mean

78°, n = 31), though all other fossils resemble Recent

shells in shape. A coeval specimen (GS4949, NZGS) is

substantially larger than the largest known Recent shell

(diameter 61 mm, as against 49 mm). All of the fossils

are indistinguishable from Recent specimens in sculp-

ture. Because I am unable to detect any character or

character state, single or combined, that would enable

separation of Recent and fossil specimens, they are all

interpreted as forms of a single polymorphic species.

Should the Recent form prove to be specifically distinct

when additional, better-preserved Waipipian material is

available, Powell's amplum will be available for the Re-

cent form.

All specimens seen from the Chatham Islands (figure

26) and beach shells from the Auckland Islands and

Campbell Island are particularly thick and heavily sculp-

tured. Two specimens from 104 m depth off the Auck-

land Islands (NZOI D80, figure 27) are thin and finely

sculptured and resemble topotypes of the multigemmata

form. Unless the species has a planktonic larval stage of

sufficient duration for transportation to these islands from

the mainland by ocean currents, it probably had a more

or less continuous distribution between them during pe-

riods of lowered sea level, presumably during the Plio-

cene or Pleistocene. That sea temperature may be a fac-

tor restricting its northernmost limit to East Cape is sug-

gested by the occurrence of two closely related species

(C. jamic.soni new species and C. gihhsorum new species)

off the Three Kings Islands where sea temperature is

cooler than off adjacent Northland due to local upwelling

(see C. tigris).

Calliostoma granti resembles C. osbornei Powell, 1926

in the rapid enlargement of S3 relative to SI and S2, but

the latter two spirals enlarge more slowly in C. osbornei,

and C. granti has a weak but distinct angulation at P2

on the early teleoconch (figures 29, 31). By comparison

with specimens of C. osbornei from north of Cape Eg-

mont, the spiral cords on the early teleoconch whorls in

C. granti are yellowish brown with white nodules instead

of being predominantly uniform white, whereas the spi-

ral interspaces are pale butt instead of orange or yellow-

ish to reddish brown Calliostoma granti differs further

from C osbornei by having stronger, more numerous

axial costae on the early teleoconch whorls, especially on

the 4th and early 5th whorls, which are also more strongly

convex.

Specimens of C. osbornei from off the south-western

North Island (figure 41) differ from most northern shells

in being as broadly conical as C. granti (sa 75-82°, mean

78°), and in that the spiral cords on the early teleoconch

whorls are reddish brown between white nodules. Spec-

imens of C. osbornei from these southern populations

differ from coexisting C. granti by having more weakly

convex 4th and 5th teleoconch whorls and b\ having

much more deeply and evenly pigmented spiral cords

on the early teleoconch whorls.

Calliostoma (Maurea) benthicola (Dell, 1950)

(Figures 33, 34)

Venmtas benthicola Dell, 1950:47, fig 21.

Maurea benthicola— PoweW, 1957:88; Powell, 1979:62.

Type Daia: Holotype MNZ M.4728, 41°21'S, 175°00'E,

Mernoo Bank, western Chatham Rise, 95 m.

Other Material Examined: 22 specimens in 5 lots MNZ.

Distribution (figure 33): Endemic to Mernoo Bank, west-

ern Chatham Rise, 75-129 m, taken alive at 95-129 m

on shell substratum.

Remarks: Calliostoma benthicola differs from C granti

by having weaker nodules, aiul by being white with

broad, reddish brown bands on the spiral cords after the

fifth teleoconch whorl. It is otherwise extremely similar,

particularly to specimens from equivalent depths off Ota-

go, and there can be little doubt that the two forms are

very closely related.

Calliostoma benthicola appears to be endemic to the

Mernoo Bank and is absent from hundreds of dredge

and trawl stations from the adjacent (Chatham Rise and

off Banks Peninsula and Kaikoura. Mernoo Bank rises

Irom depths exceeding 450 m, which are considerably

greater than the known lower limit for living C. granti

(220 m). It seems likely that C. benthicola diverged from

C. granti stock that was isolated on the Mernoo Bank

(and the Chatham Islands), perhaps following a period

of lowered sea level during the Pleistocene when Mernoo

Bank was an island and the Chatham Rise was sufficiently

Bruce A. Marshall, 1995

Page 93

close to the surface for icebergs to strand on its summit

(Culien, 1962). Calliostoma bcnthicola appears to be an

example of a species that arose from a small, isolated

population that carried only a fraction of the total genetic

material of the parent population (founder effect of Mayr,

1963). This explanation probably accounts for the origin

of the distinctive buccinid Cominella olsoni (Dell, 1956),

which is also endemic to the Mernoo Bank and is prob-

ably derived from Cominella nassoides (Reeve, 1846),

forms of which occur to the east off Kaikoura and to the

west off the Chatham Islands.

Calliostoma (Maurea) jamiesoni Marshall, new species

(Figures 30, 33, 36)

Description: Shell up to 33 mm high, higher than broad,

spire gently cyrtoconoid, spire angle 69-78° (mean 73°,

n = 20), stout, glossy, anomphalous. Colour of protoconch

and 1st 4 teleoconch whorls uniform yellowish to orange

brown. Subsequent whorls yellowish to orange brown,

spiral cords reddish brown, nodules white or buff white.

Protoconch 400 ^m wide, sculptured with fine threads

that enclose hexagonal spaces. Teleoconch of up to 8

whorls, last adult whorl contracted at maturity. Spire

whorls convex, periphery rounded, base weakly convex.

First 0.25 whorl delineated by a growth scar, with 2 axial

riblets and fine spiral threads. Subsequent whorls sculp-

tured with spiral cords, rounded nodules, and axial rib-

lets; axials strong on 1st 3 whorls, weakening on 4th

whorl, becoming obsolete on 5th whorl. Spiral cords

rounded, narrow, with broad interspaces on 1st 3 whorls,

cords broadening and interspaces narrowing on 4th whorl

Spiral cords numbering 7 or 8 on penultimate whorl and

8 or 9 on last adult whorl. Pl-4 commencing immedi-

ately, P4 partly covered by succeeding whorls, becoming

fully exposed on penultimate whorl by descent of last

adult whorl. Sl-3 appearing late on 2nd or on 3rd whorl,

gradually enlarging to resemble adjacent primaries (S3

absent in 1 specimen). A subsutural spiral appears on

penultimate or last whorl in most mature specimens and

gradually enlarges to resemble primaries; 1 or more ad-

ditional spirals intercalate on last adult whorl. Spiral in-

terspaces with collabral growth lines, most interspaces

on later spire whorls with up to 4 fine spiral threads.

Columella thick. Aperture subcircular. Outer lip thin at

rim, thickened within, inner lip spreading onto colu-

mella, parietal glaze very thin.

Type Data: Holotype MNZ M. 75141 (height 28.5 mm,

width 26.0 mm), off Prince s Rocks, Three Kings Is., alive,

15 m, 2 December 1983, scuba, G.S. Hardy and A.L.

Stewart. Paratypes (26 MNZ), all from off Three Kings

Is.: Off Three Kings Is., alive, craypot, AD. Howell (1);

Three Kings Is., alive, 5 m, March 1982, scuba, K. Burch

(2); Tasman Bay, Great King I., alive, 9 m, 19 February

1974, scuba, A.N. Baker & J. Moreland (1); South West

I., alive, 22-23 m, 2 December 1983, scuba, G.S. Hardy

& A.L. Stewart (3); S side South West I., alive, 27 m, 12

February 1986, scuba, G.S. Hardy (1); reef between Great

King 1. and Farmer Rocks, dead, 33 m, 17 February

1986, scuba, G.S. Hardy (15); North West Bay, Great

King I., alive, 30 m, 14 February 1986, scuba, G.S. Hardy

(1); N. face Hinemoa I., 24 m, 11 February 1986, alive,

3-5 m, 12 February 1986, scuba, G.S. Hardy (2).

Other Material Examined: 129 specimens from 6 stations

off Three Kings Is., MNZ, mostly immature.

Distribution (figure 33): Endemic to Three Kings Islands,

5-128 m, taken alive at 5-55 m on rock.

Diet: Intestinal tracts contained thecate hydroids (Cni-

daria) and much indeterminate organic matter.

Remarks: CMlliostonia jamiesoni is superficially similar

to C. punctulatum, C. granii, C. benthicola, and broad

forms of C. osbornei, but most closely resembles C. granti

and C. benthicola in development of the teleoconch

sculpture. It differs from all of them, however, in the

uniform coloration of the protoconch and early teleo-

conch whorls and in the low relief of the spiral cords

and nodules on the fourth to sixth teleoconch whorls

(figures 28-31). The spire tends to be more strongly cyr-

toconoid and the body whorl more contracted at ma-

turity. It differs further from C. punctulatum, C. granti,

and C. bcnthicola in the early appearance and more

rapid enlargement of the secondary spirals, especially S3,

and from C. osbornei in the roughly simultaneous ap-

pearance and rather even rate of enlargement of S2 and

S3 and the more strongly convex whorls.

Judging from development of the teleoconch sculp-

ture, C. jamiesoni is more closely related to C. granti,

C. benthicola, and C. gibbsorum new species (see below)

than to the superficially similar species C. punctulatum

and C. osbornei.

Etymology: After Peter Jamieson (Wellington), who

sought and provided much material for this revision, and

as an appreciation for his fostering of New Zealand mal-

acology in general, both amateur and professional.

Calliostoma (Maurea) gibbsorum Marshall, new species

(Figures 33, 35, 109, 130)

Description: Shell up to 28.5 mm (est.) high, higher than

broad, spire angle 53-76°, stout, glossy, anomphalous.

Colour of protoconch and 1st teleoconch whorl buff or

white. Next 2 whorls buff with yellowish brown spirals

and white nodules and axials. Subsequent whorls yellow-

ish to orange brown, nodules more lightly pigmented or

white, spirals more darkly pigmented. Protoconch 370-

400 ^im wide, sculptured with fine threads that enclose

hexagonal spaces. Teleoconch of up to 8.2 whorls. Spire

whorls weakly convex or almost flat, periphery rounded

or subangulate, base weakly convex. First 0.25 whorl

delineated by a growth scar, with 2 axial riblets and fine

spiral threads. Subsequent whorls sculptured with spiral

cords, rounded nodules, and axial riblets, the axials strong

on 1st 4 whorls, weakening and becoming obsolete on

5th whorl. Spiral cords rounded, narrow with broader

Page 94

THE NAUTILUS, Vol. 108, No. 4

Bruce A. Marshall, 1995

Page 95

Figure 32. Map of New Zealand region showing distribution

of Calliostoma (Maurca) ptinctulatum 200 and 1000 meter

contours indicated

interspaces on 1st 4 whorls, spirals broadening and in-

terspaces narrowing on 5th whorl. Spiral cords number-

ing 7 on adult penultimate whorl and 11 on base. PI- 4

commencing immediately, P4 almost or entirely covered

by succeeding whorls throughout or (as in holotype) be-

coming entirely exposed at suture from as early as 5th

whorl. SI -S3 gradually enlarging to resemble adjacent

primaries. Additional spirals intercalated on penultimate

and last adult whorls of an exceptionally large specimen.

Spiral interspaces with collabral growth lines, a few spiral

threads in some specimens. Aperture subquadrate to sub-

Figure 33. Map of New Zealand region showing distribution

of Calliostoma (Maurea) granti ( circle), C. (M.) benthicola

(star), and collectively C. (M.) gibbsorum, C. (M.) jamiesoni,

and C. (M.) regale (triangle) 200 and 1000 meter contours

indicated.

circular. Outer lip thin at rim, thickened within, inner

lip spreading onto thick columella, parietal glaze thin.

Type Data: Holotype MNZ M.35456 (height 16.8 mm,

width 12.5 mm), BS 389, between Palmer Rocks and

South East I., Three Kings Is., dead, 82 m, 18 Februar\

1974, r.v. Acheron. Paratvpes (3 MNZ): BS 893 (0639),

33°59.9'S, 171°45.3'E, Middlesex Bank, NW of Three

Kings Is., dead. 186-196 m, 31 January 1981, r.v. Tan-

Figures 28-31. Early teleoconch whorls. Primary (?) and secondary (S) spiral cords indicated. Figure 28. Calliostorrm (Maurea)

puiictulatum (Martyn, 1784). Cook Strait, 256-274 m, MNZ M.54911. Note late appearance and slow enlargement of S3, weak

axial sculpture, and broad, strongly convex .5th teleoconch whorl. Figure 29. Calliostoma (Maurea) granti (Powell, 1931). Off

Stephens Island, Cook Strait, 183-187 m, MNZ M. 50266. Note early appearance and rapid enlargement of S3 (arrowed), strong

axial sculpture, and narrow, weakly convex 5th teleoconch whorl. Figure 30. Calliostoma (Maurea) jamiesoni Marshall, new-

species. Off West Island, Three Kings Islands, 37 m, MNZ M. 80656. Note closer similarit> of sculpture to that of C, (M.) granti

(29) than to C. (M.) punclulatum (28), S3 arrowed. Figure 31. Calliostoma (Maurea) osbornei Powell, 1926, Off Cape Maria Van

Diemen, 38-43 m, MNZ M. 74665 Note late appearance of SI and S2, early appearance and very rapid enlargement of S3 (arrowed),

evenly conical outline, and flattened whorls. Scale bars = 1 mm.

Page 96

THE NAUTILUS, Vol. 108, No. 4

garoa (1); RS 921, Elingamite wreck, off West Island,

Three Kings Is., dead, 37 m, 16 March 1981, suction

dredge, K. Tarlton (1); off Three Kings Is., aUve, craypot,

AD. Howell (1).

Other Material Examined: 60 specimens from 13 stations

off Three Kings Islands, MNZ, mostly juveniles.

Distribution (figure 33): Off Three Kings Islands, 33-805

m, taken alive at 102 m and probably living as shallow

as about 30 m on rugged bryozoan/shell substratum with

corals, sponges, and gorgonians. Probably endemic.

Diet: Unknown.

Remarks: Calliostoma gibbsoriim is superficially similar

to C. jamicsoni, C. granti, C. benthicola, and to a lesser

degree, C. punctiilatxim, and C. osbornei. It closely re-

sembles C. jarniesoni. C. granti, and C. benthicola in

development of the spiral teleoconch sculpture, specifi-

cally in the more or less simultaneous origin and even

rate of development of Sl-3. Compared with the sym-

patric species C. jarniesoni, it differs constantly in colour

and colour pattern (especially when immature — see de-

scriptions); in having stronger, more crisply defined spi-

rals and nodules after the third teleoconch whorl; in

having narrower, more numerous spiral cords on the base

(11 instead of 8 or 9); and in the shape of teleoconch

whorls, which expand more slow ly and are more weakly

conve.x after the fourth whorl. Compared with the al-

lopatric species C. granti and C. benthicola, develop-

ment of teleoconch sculpture is entirely more rapid, S2

and S3 appearing and enlarging to resemble the pri-

maries a half to a full whorl earlier. Moreover, the nod-

ules are more rounded, and the axial sculpture persists

for longer and is stronger, especially on the fourth and

early fifth whorls. It differs from the allopatric species

C. punctulatum in the earlier appearance and more rap-

id enlargement of the secondary spirals (especially S3),

the stronger, more persistent axial costae, and the more

weakly convex whorls. From the sympatric species C.

osbornei it differs in numerous details of colour, colour

pattern, and sculpture, most obviously in the more or

less simultaneous appearance of S2 and S3. For further

remarks see C. jarniesoni (above).

Etymology: After David and Sharon Gibbs (Auckland)

who provided much valuable material and as an appre-

ciation for their fostering of amateur and professional

malacology in New Zealand.

Calliostoma (Maurea) osbornei Powell, 1926

(Figures 31, 37-39, 41, 43, 131)

Calliostoma osbornei Powell, 1920 591, pi 102, fig I, 2.

Calliostoma (Mauriclla) osbornei — Oliver, I92(j:l 10.

Venustas (Murrinops) osbornei — Firilay, 1926;361, .371.

Maurea (Mucrinops) osiwrnei — Powell, 19.37:64, pi. 13. fig. 4.

yentistas osbornei — Dell, 1950:47.

Maurea osbornei— ?ov,e\\, 1957:88, pi. 13, fig. 4; Powell, 1979:

62, pi. 10, fig. 5.

Maurea (Mauriclla) osbornei — Fleming, 196():38

Type Data: Holotype AIM 72037, off Cape Barrier, Great

Barrier I., c. 49 m, from fish stomach.

Other Material Examined: Fossil — GS4253, Upper Wai-

pipi Shellbed, Waverley Beach, south Taranaki (Wai-

pipian. Middle Pliocene) (1 NZGS). Recetit — 167 spec-

imens in 45 lots MNZ.

Distribution (figure 43): Middle Pliocene (Waipipian) to

Recent, Three Kings Islands southward to off Kapiti Is-

land, southern North Island (34°08.5'S-40°50'S), 0-102 m,

living at 12-93 m on the sponge Ancorina alata Dendy,

1924 on rocky ground.

Diet: Ancorina alata Dendy, 1924 (Porifera : Stelletti-

dae).

Remarks: Calliostoma osbornei bears a strong superficial

resemblance to C. punctulatum and C. granti. From C.

punctulatum it differs in numerous details, most notably

the rapid enlargement of S3 and the relatively slow en-

largement of SI and S2 (figure 31). Other differences

include the more weakly convex whorls, stronger nodules

on the early teleoconch, and the lack of a prominent

angulation at P2 on the third-fifth teleoconch whorls.

Among differences in colour and colour pattern, the most

notable are on the first four whorls, which are typically

orange with white spiral cords and nodules instead of

being predominantly white with reddish or yellowish

brown spiral cords and white or alternately spotted nod-

ules. The only known exceptions are specimens from oft

New Plymouth (figure 41), Wanganui and Kapiti Island

at the extreme southern limit of its range, which resemble

C. punctulatum in colour pattern. Most specimens of C.

osbornei are more narrowly conical than C. punctula-

tum. though some shells may be as broadly conical, es-

pecially those from the extreme south of its range (sa in

southern material 50-82°, mean 66°, S.D. 7.29, n = 52,

as against 65-96°, mean 85°, S.D. 5.83, n = 64). Despite

the similarity in shape, broad specimens of C. osbornei

are readily separable from C. punctulatum by the char-

acteristic teleoconch sculpture. The Middle Pliocene

(Waipipian) specimen lacks the early spire whorls but is

otherwise well preserved and indistinguishable from Re-

cent material. Judging from its present restricted north-

ern distribution (figure 43) and rarity south of Cape

Egmont, the lack of fossils from overlying horizons in

the Wanganui section may be related to sea tempera-

tures, which vverecooler after the Waipipian (Beu, 1966).

The present distribution of C. osbornei is not entirely

determined by that of its food, the grey sponge, An-

chorina alata, which ranges at least as far south as Banks

Peninsula (M.H.B. O'Neill, pers. comm).

Calliostoma (Maurea) regale Marshall, new species

(Figures 33, 40, 111, 132)

Description: Shell up to 12.6 mm high, glossy, of mod-

erate thickness; spire narrowly and rather evenly conical,

1.50-1.87 X higher than aperture in adults; spire angle

54-63°, anomphalous. Colour of tip of apical fold yel-

Bruce A. Marshall, 1995

Page 97

lowish brown, rest of protoconch white Most fresh spec-

imens pale yellowish brown between nodules on P2 and

P3 on 2nd and 3rd teleoconch whorls, some specimens

with addition of a sub- and/or suprasutural band of dull

olive that persists onto 4th whorl First 2 \\ horls t\ picalK

with a pinkish flush. Fresh juveniles with pale vellowish

brown spots on spiral cords on innermost third of base.

Shell elsewhere rather uniform pale buff white. Proto-

conch 370 /um wide, sculptured with a network of fine

threads that enclose hexagonal spaces, terminal vari.x

strong, rounded. Teleoconch of up to 7.25 whorls. First

few whorls rather strongly convex, angulated at P2 and

P3, subsequent whorls very weakly convex. Periphery

rounded at maturity, base more or less flat First quarter

whorl delineated by a growth scar, with a strong, rounded

axial varix and fine spiral threads. Subsequent spire whorls

encircled by prominent spiral cords with rounded nod-

ules, the spirals multiplying by intercalation of second-

aries and tertiaries that enlarge to resemble primaries.

Nodules evenly developed on each spiral, occasionally

becoming very weak after 5th whorl on spirals other

than PI. PI commencing late on 2nd or early on 3rd

whorl, gradually enlarging to resemble P2 and P3, which

commence immediately after growth scar on 1st whorl.

P4 covered by succeeding whorls, becoming fully ex-

posed on penultimate whorl by descent of last adult whorl,

nodular on last adult whorl, smooth before it SI com-

mencing from early on 3rd whorl to start of 6th whorl,

S2 commencing from late on 2nd whorl to midway

through 3rd whorl, S3 commencing from late on 3rd

whorl to midway through 4th whorl In adults a tertiary

spiral intercalates on penultimate whorl, where the total

of 8 spirals includes emergent P4. Additional tertiaries

intercalate on last adult whorl. Axial riblets strong on 1st

3 whorls, weakening on 4th and becoming obsolete on

5th whorl. Interspace of P3 and P4 on 1st 4 whorls with

numerous fine, close, crisp, secondary axial riblets that

become obsolete on succeeding whorl. Basal spirals mul-

tiplying by intercalation of a few secondaries to number

about 10 in adults; interspaces about as wide as each

spiral or narrower, sculptured with fine collabral growth

lines; spirals weak and smooth in immature specimens,

becoming stronger and nodular with increasing size, rarely

more or less smooth in adults. Outer basal spirals resem-

bling spire spirals, inner spirals stronger and more strong-

ly nodular. Aperture subquadrate, strongly thickened

within and immediately behind apertural rim in adults,

especially at base and columella.

Type Data: Holotype M.86730 (height 12.5 mm, width

9.30 mm) and paratype MNZ, BS 902 (0648), 34°10.5'S,

172°I1 4'E, off Three Kings Is., dead, 153 m, 1 February

1981, r.v. Tangaroa. Paratypes (14, all from off Three

Kings Is): BS 900 (0646), 33°57.0'S, 171°45.4'E, alive,

98-103 m, 31 January 1981, r.v. Tangaroa (3 MNZ); BS

905 (0651), 33°57.4'S, 172°19.4'E, alive, 128-123 m, 1

February 1981, r.v. Tangaroa (1 MNZ); BS 894 (0640),

34°00.9'S, 171°44.7'E, alive, 201-216 m, 31 January 1981,

r.v. Tangaroa (1 MNZ); BS 898 (0644), 34°01.2'S,

17r45.8'E, alive, 221-206 m, 31 January 1981, r.v. Tan-

garoa (4 MNZ); E 846, 34°07.5'S, 17r57.5'E, dead, 417

m, 16 March 1968, m.v. \'iti (1 NZOI); BS 902 (0648),

34°10 5'S. 172°11.4'E, dead, 1.53 m, 1 February 1981,

r.v. Tangaroa (2 MNZ); BS 906 (0652), 34°14.8'S,

172°13.6'E, dead, 173-178 m, 2 February 1981, r.v. Tan-

garoa (1 MNZ).

Other IHaterial Examined: 189 specimens from 21 sta-

tions oH Three Kings Is., 53-805 m, MNZ, mostly im-

mature.

Distribution (figure 33): Off Three Kings Islands

(33°57.0'S-34°22.8'S), 53-805 m, taken alive at 98-221 m

on rugged, comminuted bryozoan/shell substratum with

sponges, hydroids, gorgonians, corals, etc.

Diet: l^nknown

Remarks: Calliostoma regale resembles the sympatric

species C. osbornei in shape but differs in details of colour

and colour pattern and in development of teleoconch

sculpture, including appearance of S2 before S3 and slow

enlargement of PI on the first two whorls. Among pre-

viously described New Zealand Recent calliostomatids,

it is rendered highly distinctive by the presence of fine

axial riblets between P3 and P4 on the early teleoconch

whorls, which facilitates recognition of even very im-

mature specimens (figure 109). Calliostoma regale seems

to be closely related to C. simplex Schepman, 1908 from

the Banda Sea, from which it differs principally in having

more markedly convex spire whorls and stronger nodules

on the base. Moreover, PI develops later, and P2 and P3

are relatively larger on the early teleoconch whorls. Al-

though the protoconch and first teleoconch whorl of the

holotype of C. simplex (ZMA) are eroded, enough re-

mains to show that PI is present almost immediately

after the protoconch and that it is as large as P2 midway

through the second whorl. In C. regale, PI commences

late on the second or early on the third whorl and does

not rival P2 in size until midway through the third or

fourth whorl Calliostoma simplex is otherwise similar

in the order of appearance of the secondary spirals and

in having axial riblets between P3 and P4.

Etymology: Royal (Latin). Alluding to the Three Kings

Islands.

Calliostoma (Maurea) aupourianum Marshall, new

species

(Figures 42, ,50, 112, 133)

Description: Shell up to 8 60 mm high, glossy, of mod-

erate thickness; spire narrowly and rather evenly conical,

up to 1.65 X higher than aperture; mean spire angle 58-

65°, anomphalous Colour of extreme tip of protoconch

yellowish brown, elsewhere white. Subsequent whorls

yellowish or pale yellowish brown, spire irregularly ax-

ially mottled in a darker shade, each dark band followed

by a narrow white band, base in adults irregularly axially

mottled in yet darker shades Protoconch 370-380 ^lm

Page 98

THE NAUTILUS, Vol. 108, No. 4

Menioo Bank, Clhathaiii Hise, 9.5 m, MNZ M. 23626

29.0 mm).

VifHirp :i4: Culliostoma {Maurea) hcnthicola (DeW, 19.56). Mernoo Hank, (..Mat .-x,-, .. or <kc; ; ic s

Figure .-{5. Calliostoma (Maurca) gibhsorum Marshall, new species. Hololype, off Three Kings Islands, 82 m, ^NZ M.35456 (16.S

X 12.5 mm) Figure 36. Calliostoma (Maurea) jamiesoni Marshall, new species. Holotype, off Three Kings Islands, 15 m, MNZ

M 75141(28.5 x 26.0 mm). Figures 37-39, 41. Calliostoma (Maurea) oshornei Powell, 1926. 37. North Ahipara Bank, western

Northland .53-56 m MNZ M.74590 (25.0 x 19.5 mm). 38. Between Pandora Bank and C^ape Maria Van Diemen, 38-4.3 m, MNZ,

M 70955 (22 7 x 20 8 mm). .39. Astrolabe Reef, off Motiti Island, 20-23 m, MNZ M. 117841 (32.7 x 26.0 mm).

41. Seal Rock,

Bruce A. Marshall. 1995

Page 99

wide, sculptured with network of fine threads that en-

close roughly hexagonal spaces, terminal varix strong,

rounded. Teleoconch of up to 6.4 whorls. First few whorls

rather strongly convex, later whorls weakly convex, early

or all whorls distinctly angulated at P2, periphery su-

bangulate at maturit\ , base more or less flat. F"irst quarter

whorl delineated by a fine growth scar, sculptured with

2 rounded axial varices and fine spiral threads. Subse-

quent whorls encircled by prominent spiral cords with

conical nodules, spirals multiplying by intercalation from

3 (P2-P4) to 7 (P1-P4, Sl-So), secondaries enlarging to

resemble primaries. P4 smooth throughout, S.'3 finely nod-

ular or smooth, other spire spirals nodular. On 3rd and

4th whorls, or all whorls other than last adult whorl, a

single nodule on P2 is considerably enlarged and white

immediately following each dark mottling; interval at

about each 3rd nodule on early whorls, intervals variable

and up to 7 nodules apart on later whorls. PI com-

mencing early on or midway through 2nd whorl, grad-

ually enlarging to resemble P3; P2 and P3 commencing

immediately after growth scar on 1st whorl. P4 almost

or entirely covered by succeeding whorls, becoming fully

exposed on penultimate whorl by descent of last adult

whorl, smooth throughout. Si commencing late on 5th

or 6th whorl or entirely absent in adults, S2 commencing

late on 3rd to midway through 4th whorl, S3 commenc-

ing early on 4th or 5th whorl. Up to 3 tertiary spirals

may intercalate on last adult whorl. Axial riblets strong

on 1st 3 whorls, weakening and becoming obsolete on

4th whorl. Interspace of P3 and P4 on 3rd-5th whorls

with numerous fine, close, crisp, secondary axial riblets

that become obsolete on succeeding whorl. Basal spirals

smooth throughout, nimibering 11 or 12. Fine collabral

growth lines throughout, stronger on base. Aperture

subquadrate, strongly thickened immediately within in

adults, especially at base and columella.

Type Dala: Holotvpe M86731 (height 7.15 mm, width

6.00 mm) and paratype MNZ: BS 395, 34°10'S, 172°12'E,

off Three Kings Is., dead, 252 m, 19 February 1974, r v.

Acheron. Paratypes (5 MNZ); BS 905 (0651)', 33°57.4'S.

I72°19.4'E, off Three Kings Is., alive, 128-123 m, 1 Feb-

ruarv 1981, r.v. Tangaroa (1); BS 901 (0647), 34°14.1'S,

172°b9.0'E, off Three Kings Is., dead, 192-202 m, 1 Feb-

ruary 1981, r.v. Tangaroa (1); BS 833 (0578), 37°38.5'S,

178°56.4'E, SE slope of Ranfurly Bank, East Cape, dead,

153-143 m, 22 January 1981, r.v. Tangaroa (3).

Other IVIaterial Examined: 38 specimens from 13 stations

off Three Kings Is., dead, 102-805 m, MNZ, mostly im-

mature; BS 747 (R105), 37°16.7'S, 176°17.5'E, off E side

Mayor I., dead, 104-109 m, 22 January 1979, r.v. Tan-

garoa (2 immature).

Figure 43. Map of New Zealand region showing distribution

of Calliostoma (Maurea) osbornei. 200 and 1000 meter con-

tours indicated.

Distribution (figure 50): Off Three Kings Islands. Mayor

Island, and East Cape, 102-805 m, taken alive at 123-

128 m on rugged comminuted bryozoan/shell substra-

tum with sponges, hydroids, gorgonians, corals, etc.

Diet: LInknown.

Remarks: Compared with the s\ mpatric species Cal-

liostoma regale, which it most closely resembles, C. au-

pourianum differs in the later appearance and slower

enlargement of the secondary spirals, in having more

sharply pointed nodules, in the irregular size of the nod-

ules on P2, and in details of colour pattern as described

above.

Etymology: .\lluding to the distribution within the

boundaries of the former Aupourian marine province.

New Plymouth, 12 m, MNZ M. 80156 (28.0 x 27.0 mm). Figure 40. Calliostoma (Maurea) regale Marshall, new species Holotvpe.

off Three Kings Islands. 153 m, MNZ M. 86730 (12.5 x 9.30 mm) Figure 42. Calliostoma (Maurea) aupourianum Marshall, new

species. Holotype. off Three Kings Islands, 252 m. MNZ M. 86731 ( 7.15 x 6.00 mm).

Page 100

THE NAUTILUS, Vol. 108, No. 4

Calliostoma (Maurea) spectabile (A. Adams, 1855)

(Figures 44-47, 50, 113, 134)

Zizyphintis spectabilis A. Adams, 1855:37, pi. 27, fig. 7; Reeve,

1863, pi. 1, fig. 5a,b; Hutton, 1873:38; Hutton, 1880:98;

Hiitton, 1884:360.

Calliostoma spectabilis— Pihbry, 1889:332, pi. 16, fig, 12.

Calliostoma spectabile— Suter, 1897:280; Suter, 1913:147, pi.

40, fig. 5.

Calliostoma (Mauriella) spectabile — Oliver, 1926:110.

Venustas (Mucrinops) spectabilis — Finlay, 1926:360.

Venustas (Mticrinups) spectabilis — Finlay, 1926:360, 371.

Maurea (Mucrinops) spectabilis — Powell, 1937:64

Venustas spectabile— DeW, 1950:45, figs. 16-18; Powell, 1955:

55.

Maurea spectabile — Powell, 1957:88.

Calliostonm (Maurea) spectabilis — Cemohorsky, 1977:93, fig. 9.

Maurea spectabilis — Powell, 1979:63, pi 19, fig. 1.

NOT Venustas (Mucrinops) spectabilis— Finlay, 1926:362, pi.

18, fig. 26 (C. foveauxanum).

Type Data: Hoiotype BMNH 1968150 (Cernohorsky,

1977, fig. 9).

Other Material Examined: 55 specimens in 27 lots MNZ,

4 specimens in 2 lots NZOI.

Distribution (figure 50): Off Auckland and Campbell

islands, 0-146 m, taken alive at 9-146 m from hard sub-

strata.

Diet: Guts examined contained thecate hydroids (Cni-

daria) and much indeterminate organic matter.

Remarks: This species is characterised by a large, heavy,

strongly sculptured shell; cyrtoconoid early spire whorls;

and convex, rapidly expanding early teleoconch whorls.

Late teleoconch whorls range from strongly convex to

almost flat, and the spire is broadly to narrowly conical

(sa 52-80°, mean 61°, SD 7.59, ;i = 29), the most broadly

conical specimens tending to have the most strongly con-

vex whorls. Specimens washed ashore and living as deep

as 9 m at the Auckland Islands (figure 45) have the most

broadly conical spires (sa 60-80°, mean 70°, SD 5.83, n

= 9). Auckland Islands specimens (figure 47) from great-

er depths (37-146 m) and all Campbell Island shells (fig-

ure 46) are exclusively narrow-spired (sa 52-60°, mean

56°, SD 2.93, n = 20), though some beach shells from

the Auckland Islands may be as narrowly conical (figure

44). Most specimens from Campbell Island (figure 46)

are more weakly nodular than Auckland Islands speci-

mens, and have slower enlargement of the secondary

spirals, thougli similar forms occur in both populations.

Whilst Auckland and C^anipbell islands populations are

probably conspecific, the differences suggest that there

may be little genetic exchange between them.

Calliostoma (Maurea) foveauxanum (Dell, 1950)

(Figures 48, 50, 114, 1.35)

Venustas (Mucrinops) spectalnllt — Kiiilay, 1926:362. pi. 18,

fig. 26 (not A. Adams, 1855).

Venustas foveauxana Dell, 1950:45, fig. 13-15.

Maurea /oucaiixana— Powell, 1957:88; Powell, 1979:63, pi. 10,

fig 7,

Calliostoma spectabile foveauxanum — Beu, 1976:78.

Maurea WacAi— Abbott & Dance, 198240 (not Powell, 1950)

NOT Calliostoma (sensu lalo) foveauxanum — Beu & Maxwell,

1990:404 (C. blacki Powell, 1950).

Type Data: Hoiotype MNZ M.4727, Foveaux Strait, 53

m. Paratype (1 MNZ), Stewart Island.

Other Material Examined: 68 specimens in 8 lots MNZ,

5 specimens in .3 lots NZOI.

Distribution (figure 50): South-eastern South Island,

Stewart Island, and The Snares, 73-549 m, taken alive

at 91-220 m from bryozoan /shell substrata.

Diet: Intestinal tracts examined contained mostly thecate

hydroids (Cnidaria) with some indeterminate organic

matter.

Remarks: From Calliostoma spectabile, to which it is

most closely related, C. foveauxantim differs principally

in being more finely sculptured and in that the contour

of the early teleoconch is more weakly cyrtoconoid. Al-

though they are allopatric, there is no clinal intergra-

dation within their respective geographic ranges, so it

seems likely that they are distinct, closely related species

rather than disjunct populations of a single species. An-

other closely related species from the Antipodes Islands

is described below.

Calliostoma (Maurea) eminens Marshall, new species

(Figures 49, 50)

Description: Shell up to 51 mm high, higher than broad,

spire up to 1.68 x higher than aperture, stout, weakly

cyrtoconoid, spire angle 62-66°, suture deep, anomphal-

ous. Colour yellowish brown, spiral cords reddish brown,

nodules white, basal spirals alternately spotted reddish

brown and white. Parietal and inner lips porcellaneous

white. Protoconch unknown (etched). Teleoconch of up

to at least 8.5 convex whorls, earliest whorls unknown

(eroded — whorl numbers estimated). First 5 whorls rath-

er evenly convex, subsecjuent whorls flattened adapically,

periphery rounded, base weakly convex. After 5th whorl,

zone between suture and PI steepens until almost ver-

tical, channeling suture. Sculpture consisting of strong,

rounded spiral cords and weaker axial costae, axials

weakening and vanishing on 4th whorl. Spiral cords num-

bering 6 or (usually) 7 on adult penultimate whorl, com-

prising P1-P4 plus intercalating secondaries that com-

mence after 1st 4-4.5 whorls, additional tertiaries inter-

calate on last ailult whorl, 7-10 cords on base. First 3

whorls inserted against P4, subsecjuent whorls descending

and inserted between P4 and outermost basal spiral. Spire

spirals \\ ith rounded nodules, those on PI strongest, nod-

ules finer and more numerous on abapical half of each

whorl, basal spirals undulant. F"ine collabral growth lines

throughout, some specimens with fine spiral lirae in some

spiral interspaces. Aperture ovate. Outer lip thin at rim.

Bruce A, Marshall, 1995

Page 101

Figures 44-4-7. Calliostoma (Maurea) spectalnlc (A. Adams, 1855). 44. Auckland Island, beach, MNZ MJ02600 {48.0 x 43.0

mm). 45. West coast, Auckland Island, 9 ni, MNZ M. 36711 (51.0 x 49.3 mm). 46. Boyack Point, Campbell Island, 6 m, MNZ

Ml 17480 (47.0 x 39 4 mm) 47. North of Auckland Islands, 113m, NZOl D200 (56.0 x 46.0 mm) Figure 48. Calliostoma

(Maurea) foveauxanum (Dell, 1950). Off Otago Penmsula, 130-150 m, MNZ Ml 17268 (53.5 x 46.8 mm) Figure 49. Calliostoma

(Maurea) eminens Marshall, new species. Holotype, off Archway Island, Antipodes Islands, 13-15 m, MNZ M. 841 12 (50.8 x 45.0

mm).

thickened within, parietal and inner lips a continuous

spreading glaze.

Type Data: Holotype M. 84112 (height 50.8 mm, width

45.0 mm, 8 teleoconch whorls) and paratype MNZ, N

side of Archway I., Antipodes Is., alive on sloping rock,

13-15 m, 29 November 1978, scuba, D.S. Horning. Para-

types (18): A72S, 49°38.4'S, 178°48.7'E, off Antipodes Is.,

dead, 95 m, 7 November 1962, m.v. Taranui (1 NZOI);

49°40'S, nS'Sa'E, off Antipodes Is., dead, 103 m, USNS

Ehanin stn 1850 (9 USNM, 3 MNZ): A739, 49°40.19'S,

178°44.3'E, off Antipodes Is., dead, 113 m, 9 November

1962, m.v. Taranui (1 NZOI); A 723, 49°42'S, 178°50.3'E,

off Antipodes Is., dead, 123 m, 7 November 1962, m.v.

Taranui (4 NZOI); E side Perpendicular Head, Antip-

odes Is., under large boulder, alive, 19 m, 27 November

1978, scuba, D.S. Horning (1 MNZ); S side Ringdove

Bay, Antipodes Is , on rocks, alive, 18-20 m, 21 November

1978, scuba, D.S. Horning (2 MNZ),

Distribution (figure 50): Off Antipodes Islands, 13-123

m, taken alive at 13-20 m from hard substrata. Probably

endemic.

Diet: Guts contain mostly thecate hydroids (Cnidaria)

and some indeterminate organic matter.

Remarks: Calliostoma eminens differs from its allopat-

ric relatives C. spectabile and C. foveauxanum in having

more strongly convex whorls and in that the suture is set

lower on each whorl and is thus correspondingly deeper

and the whorls correspondingly more strongly convex.

It differs further from C. specialnle in being more lightly

built and from C foveauxanum in its stronger basal

sculpture.

Page 102

THE NAUTILUS, Vol. 108, No, 4

Figure 50. Map of New Zealand region showing distribution

of Calliostoma (Maurea) attpourianum (hollow star), C. (M.)

spectabile (circle), C. (M.) foveauxanum (square), and C.(M.)

eminens (solid star). 200 and 1000 meter contours indicated.

Etymology: High (Latin)

Calliostoma (Maurea) blacki (Powell, 1950)

(Figures .51-63, 11.5, 1:36)

Venustas blacki Powell, 1950:80, pi. 7, fig. .3, 4.

Vcnusliif: roupcri Velia, 195454.3, pi- 25, fig. 6, New synonym.

ThorUlclla chathamcnsis profunda Dell, 195644, pi. 6, fig.

57, New synonym

Maurea blacki— PoweW, 1957:88; Powell, 1979:6.3, pi. 19, fig.

10, 11.

Maurea (Mauriella) couperi — Fleming, 1966:38; Beu, 1978:

724, figs. 11, 13/13.

Calliostoma (Maurea) spectalrile Jovcauxana — Heu, 1979:88

(in part not Dell, 1950)

Calliostoma spectalrile couperi — Heu, 1981:71, pi. 32, fig. 13.

Calliostoma (sensu lato) couperi — Beu & Ma.vwell, 1990:404.

Calliostoma (sensu lato) joveauxanum — Beu & Maxwell, 1990:

404 (not Dell, 1950).

NOT Maurea Wacfci— Abbott & Dance, 1986:40 (C. joveaux-

anum)

NOT Maurea blackii [.sir]— llnrikoshi, 1989, pi 4, fig. 12 (C.

granti).

Type Data: Venustas blacki: Holotype AIM 71168, off

eastern Otago, c.128 m; Venustas couperi: Holotype

NZGS TM 4995, east side of Makara Stream, north of

Te Awaite cutting, Wairarapa (early Nukumaruan, Late

Pliocene).

Other Material Examined: 128 Recent specimens in 30

lots MNZ, 34 specimens in 13 lots NZOI.

Distribution (figure 63): Late Pliocene (early Nukuma-

ruan) to Recent, Chatham Rise, South Island east coast

from Kaikoura southwards; Stewart, Snares, Auckland,

Campbell, and Bounty islands, 73-549 m, taken alive at

95-549 m from bryozoan/shell substrata.

Diet: Gut contents examined comprised thecate hydroids

(Cnidaria) and indeterminate organic matter.

Remarks: This species has great variation in shell mor-

phology and is frequently confused with C. joveaux-

anum (Beu, 1976:79; Powell, 1979:497) and C. siinulans

in collections. The spire angle ranges from 60° to 91°

(mean 75°, SD 6.64, n = 59), and the whorls may be

more or less flat-sided throughout with an angulate pe-

riphery, or they may become convex w ith increasing shell

size so that the last whorl is evenly rounded. Shell thick-

ness may vary by up to a factor of three, and there is

considerable variation in the strength of the spiral cords

and nodules. There is cotnplete integration between all

extremes in shell morphology, and it is clear that all of

the specimens examined represent a single, highly poly-

morphic species. Specimens from the Chatham Rise (fig-

ures 51, 53) and some from off Kaikoura, at the north-

ernmost limit of its range, tend to be the most lightly

built, more finely sculptured, and more broadly conical

(sa 78-91°, mean 82°, n = 17) than the more southern

ones. The whorls tend to become markedly convex-sided

towards maturity (figure 51), although they may remain

weakly convex in some specimens from the Chatham

Rise east of the Mernoo Bank (figure 53). Specimens from

the Pegasus Canyon off Banks Peninsula (figure 52) and

from southward to off Timaru (figures 55, 56) are slightly

heavier-shelletl than specimens from the north, and most

are more narrowly conical (sa 71-79°, mean 73°, ;i = 6)

and more strongly nodular, with more weakly convex

whorls. In some specimens from off East Otago (sa 71-

83°, mean 76°, SD 3.13, n = 28) (figure 57), including

the holotype, the whorls become as strongly convex as

in most specimens from localities to the north, but most

of them are as narrowly conical as shells from Pegasus

Canyon southward to off Timaru. Specimens from off

Stewart Island and The Snares (figure 58) are indistin-

guishable from most East Otago specimens. Specimens

from off the Auckland Islands (figure 62) and Campbell

Island are the thickest and most strongly sculptured and

arc consistently narrowly conical (sa 62-70°, mean 67°,

n = 4) with almost llat-sidcd whorls. Specimens from off

the Bounty Islands (figure 60) have the most strongK

flattened whorls and the narrowest spires (msa 60-63°,

mean 61°, n = 5) and arc thiinicr and more fineK sculp-

Bruce A. Marshall, 1995

Page 103

tured than shells from off Auckland and Campbell is-

lands. Thus the shell in C. blacki tends to become more

narrowly conical towards the south and shows a roughly

oscillating dine in whorl convexity between Kaikoura

and East Otago.

Evidence that spire angle is at least partly a response

to sea temperature is suggested by the occurrence of long-

dead (probably Pleistocene), narrowly conical, thick,

strongK nodular shells in dredge samples containing liv-

ing specimens with broadK conical, thin, finely nodular

shells from the Mernoo Bank (BS 655, 43°07.2'S,

175°22.9'E, 148-150 m, MNZ M.61 184, M. 61 185) (6gure

61) and off the Chatham Islands (NZOI D876, 43°20'S,

176°50"W, 148 m) A long-dead (probabb Pleistocene)

specimen has been obtained from off Palliser Bay (41°33'S,

174°50'E, C.274 m, MNZ Ml 1534), which is north of the

northernmost known living specimens (NZOI C703, off

Kaikoura, 42°42'S, 173°37.8'E, 180-140 m), and resem-

bles living specimens taken from Pegasus Canyon and

southward to off Timaru. This specimen occurred with

similarly preserved valves of the stenothermic, cool-wa-

ter pectinid Chlamys delicatitla (Hutton, 1873) (MNZ

Ml 1521), which is currently unknov\n living north of

Cook Strait (Pantin, 1963). Past northward range expan-

sions of C. delicatula have been interpreted as evidence

of lowered sea temperature by Fleming (1944, 1951),

Borehamand Fleming ((>i Pantin 1957), Beu(1969, 1974,

1977), and Beu ct al. (1977). Past northward range ex-

tensions of C. blacki also appear to be associated with

low temperatures, and the narrowly conical, flat-sided,

strongly sculptured, subrecent northern shells are con-

sidered to have lived when sea temperatures were cooler

than at present. This explanation could account for the

occurrence of old-looking shells from submarine canyons

off East Otago, notably the Papanui Canyon (NZGS

RM4721), which have flatter-sided whorls than any known

Recent specimens from the vicinity. The holotype of the

Late Pliocene (Nukumaruan) Venustas couperi Vella,

1954 (figure 59) is essentially similar to Recent specimens

of C. blacki from off Timaru and East Otago, and it is

considered to be conspecific. Thoristclla chathamensifi

profunda Dell, 1956 is based on juveniles from the vi-

cinity of the type locality. The Late Pliocene (Nuku-

maruan) specimen from Oaro recorded by Beu (1979)

resembles Recent specimens from the vicinity (off Kai-

koura). Calliostonm blacki may be distinguished from

C. joveauxanum by the following characteristics: (1) spi-

ral interspaces translucent white or pinkish white instead

of yellowish brown; (2) spiral cords reddish brown with

white nodules after first 1.5 whorls instead of uniform

reddish brown or at least unicoloured (including nodules)

on first 4 teleoconch whorls; (3) P4 on first 5 teleoconch

whorls almost entirely covered by succeeding whorls and

alternately spotted reddish brown and white instead of

almost entirely exposed and uniform reddish brown; (4)

5th and 6th teleoconch whorls more strongly convex in

C. foveauxanuni; and (5) C. blacki attains smaller shell

size than C. joveauxanum w itiiin the geographic range

of the latter (height up to 50 mm as against 62 mm or

more). Unlike C. blacki. shell morphology in C. fov-

eauxanuin is extremelv stable.

Calliostoma (Maurea) megaloprepes (Tomlin, 1948)

(Figures 63, 64)

Maurea (Mucrinops) megaloprepes Tomlin, 1948:225, pi. 2,

fig. 1.

Venustas megaloprepes — Powell, 1955:55.

Maurea megaloprepes — Powell, 1957:88; Powell, 1979:63, pi.

19:2,

Type Data: Holotype BMNH 1951.6.13.38, BANZARE

stn 80, off north-eastern corner of Macquarie Island, 120-

80 m.

Other IMalerial Examined: 15 live-taken specimens:

54°24'S, 159°01'E, 79-93 m, 10 February 1965, USNS

Eltanin stn 1417 (6 USNM, 3 MNZ); C733, 54°25'S,

159°02'E, 104 m, 25 November 1961, m.v. Viti (2 NZOI);

54°32'S, 159°02'E, 86-101 m, 10 February 1966, USNS

Eltanin stn 1418 (1 USNM); D9, 54°52'S, 158°50'E, 113

m, 20 April 1963, m.v. Viti (2 NZOI); C730, 54°55'S.

158°47'E, 110 m, 24 November 1961, m.v. Viti {I NZOI).

Distribution (figure 63): Off Macquarie Island, living at

79-113 m on bryozoan/shell substrata. Endemic.

Diet: Gut contents examined comprised mostly forami-

niferal sand with fragments of thecate hydroids (Cni-

daria) and indeterminate organic matter.

Remarks: This species is rendered highly distinctive by

its rich chestnut-brown shell coloration and by the virtual

obsolescence of all of the spiral cords other than (rarely

including) PI on the sides and base of the last two adult

whorls. Calliostoma megaloprepes is otherwise similar

to C. blacki and the two species are evidently closely

related.

Calliostoma (Maurea) simulans Marshall, 1994

(Figures 63, 65-68, 116, 137)

Calliostoma (Maurea) simulans Marshall, 1994: 68, pl.l, figs.

1-3.

Type Data: Holotype MNZ M.87450 (height 30.5 mm,

width 29.0 mm, 8.3 teleoconch whorls) and 33 paratvpes

MNZ: BS 558, 43°30'S 173°31.3'E, head of Pegasus Can-

yon, NE of Banks Peninsula, alive, 446 m, 27 September

1976, r.v, Acheron. Paratypes (158 MNZ): BS 786, 43°25'S,

173°26'E, wall of Pegasus Canyon, alive, 329-183 m, 21

February 1979, r.v. Acheron (35); BS 785, 43°25'S,

173°26'E, Pegasus Canvon, alive, 485- 476 m, 21 Feb-

ruary 1979, r.v. Acheron (59); BS 784, 43°29.5'S,

173°30.5'E, Pegasus Canyon, alive, 402-338 m, 21 Feb-

ruary 1979, r.v. Acheron (12); BS 783, 43°31'S, 173°30.5'E,

Pegasus Canyon, alive, 256-293 m, 21 February 1979,

r.v. Acheron (52).

Other Material Examined: 3 specimens in 3 lots MNZ,

28 specimens in 13 lots NZOI, 3 specimens in 1 lot USNM.

Page 104

THE NAUTILUS, Vol. 108, No. 4

Bruce A. Marshall, 1995

Page 105

Distribution (figure 63): Recent, Challenger Plateau, off

New Plymouth, Cxxik Strait to SE of Banks Peninsula,

Chatham Rise, and off Bounty and Campbell islands,

183-1006 m, taken alive at 256-410 m from soft substrata

with shells.

Diet: Intestinal tracts of all specimens examined con-

tained fragmented chitinous polychaete tubes (Seden-

taria, probably Chaetopteridae) and indeterminate or-

ganic matter.

Remarks: Calliostoma simulans is characterised by a

rather large shell with strong spiral cords on the spire

and convex, evenly expanding whorls. Shell characters

are rather stable throughout the geographic and bathy-

metric range, though specimens from the west coast (fig-

ure 67) and from Cook Strait southward to Banks Pen-

insula, including the type material (figure 65), are more

lightly built and attain smaller size than specimens from

elsewhere (figures 66, 68). Occurring throughout most

of the range of the highly polymorphic species C. hiacki,

and locally in sympatry, most specimens of C. simulans

are strongly differentiated from C. blacki. but some forms

of C. blacki from the Chatham Rise and off the south-

eastern South Island (figures 54, 57) approach C. simu-

lans in convexity of the late teleoconch whorls. Callios-

toma simulans differs from all forms of C. blacki in

having more strongly convex early teleoconch whorls. It

differs further from C. blacki from localities other than

the eastern Chatham Rise in that SI becomes as large as

PI or (in most specimens) larger, and the nodules on SI

become correspondingly as large or larger. By contrast,

in C. blacki from other than the eastern Chatham Rise,

SI only occasionally becomes as large as PI and the

nodules on PI are consistently larger than those on SI

before the last adult whorl. Although specimens of C.

blacki from the Chatham Rise east of Veryan Bank (fig-

ure 53) have PI as finely nodular as in C. simulans, all

pass through a stage on the early teleoconch when PI is

by far the most strongly nodular spiral cord as in other

forms of the species. Despite the similarity of some Chat-

ham Rise specimens to C. blacki. the two species are

strongly differentiated where they occur in strict sym-

patry, such as off Banks Peninsula (figures 52, 65) and

particularly off the Bounty Islands and off the Auckland

Islands and Campbell Island (figures 62, 68).

Calliostoma (Maurea) antipodense Marshall, new spe-

cies

(Figures 63, 69, 117, 138)

Figure 63. Map of New Zealand region showing distribution

of Calliostoma (Maurea) blacki (solid circle), C. (M.) maiii

(open circle), C. (M.) simulans (star, solid star = sympatric

with C. (M.) blacki), C. (M.) antipodense (inverted triangle),

and C. (M:) megaloprepes (upright triangle) 200 and 1000

meter contours indicated.

Description: Shell up to 37 mm high, slightly higher

than broad, rather thin, anomphalous, glossy; spire up to

1.77 X as high as aperture, rather evenly conical, spire

angle 71-74° Colour orange buff, spiral cords reddish

brown between paler nodules. Protoconch 400 ^m wide,

sculptured with network of fine threads that enclose hex-

agonal spaces, terminal varix strong. Teleoconch of up

to 8.5 strongly convex whorls, suture becoming deeply

channeled after 4th whorl, periphery rounded, base

Figures 51-62. Calliostoma (Maurea) blacki (Powell, 1950). 51. Mernoo Bank, Chatham Rise. 95 m, MNZ M.11837S (37,5 x 40.0

mm). 52. Pegasus Canyon, off Banks Peninsula. 256-293 m, MNZ M. 64651 (41.7 x 40.4 mm), 53. Off Chatham Islands, 315-279

m, MNZ M.90037 (40.3 x 43.4 mm) 54. Off Oamaru, c 90 m, MNZ M. 102603 (44.3 x 42.8 mm) 55, 56. Off Timaru, 293 m,

NZOI E424 (55 0 x .55,3 mm, and 51 0 x 49.4 mm). 57. Continental Shelf off Dunedin, MNZ M 7209 (51.0 x .500 mm) 58.

Off The Snares, 154-168 m, MNZ M, 92443 (42,3 x 41.0 mm). 59. Holotype of Venustas couperi Velia, 1954, Late Pliocene-Early

Pleistocene (Nukumaruan), Makara Stream, Wairarapa, NZGS TM4995 (34 2 x ,35.8 mm) 60. Off Bounty Islands, 155 m, NZOI

1708 (42.8 X ,37.5 mm), 61. Subrecent (probably Late Pleistocene), Mernoo Bank, Chatham Rise, 148-150 m, MNZ M. 61185 (41.0

X 39 0 mm) 62. Off Auckland Islands, 113 m, NZOI D200 (45.4 x 40.0 mm).

Page 106

THE NAUTILUS, Vol. 108, No. 4

weakly convex. First 0.2 whorls delineated by a growth

scar, with 2 axial costae and 4 spiral threads. Subsequent

spire whorls sculptured with spiral cords that multipK

by intercalation from 4 (P1-P4) to 9 (P1-P4, S1-S3 + 2

suprasutural spirals); summit of P4 partly covered by

succeeding whorls on 1st 4 whorls, after which insertion

point progressively descends to fully expose P4 and then

2 additional spirals, the lower of which is either partly

covered by succeeding whorls or fully exposed; nodules

rounded on PI, P2, P3, SI, and S2, other spirals smooth,

spiral interspaces considerably wider than each spiral.

PI at first much weaker than P2 and P3, which are strong

and similar throughout, gradually enlarging over 1st 5

whorls then weakening and becoming obsolete. Second-

ary spirals enlarging to resemble adjacent primaries. SI

and S2 commencing on mid 4th to early 5th whorl, S3

commencing mid 4th to mid 5th whorl. Base with 6 spiral

cords of similar size, weaker than spire spirals, inner 3

smooth or with rounded nodules, others smooth. Axial

costae strong on 1st 3 whorls, weakening and vanishing

on 4th whorl. Fine spiral lirae on 1st 3 or 4 whorls, fine

collabral growth lines and obscure spiral lines through-

out; fine spiral threads in basal interspaces and interca-

lating in spiral interspaces on spire on last 1 or 2 adult

whorls. .Aperture ovate, outer lip simple, inner lip thick-

ened, parietal glaze extremely thin and transparent.

Type Data: Holotype MNZ M.80434 (height 31.8 mm,

width 29.2 mm, 8.3 teleoconch whorls) and paratype, off

Leeward L, Antipodes Is., alive, 18-73 m, 21 November

1972, r.v. Acheron. Paratypes (31): Eltanin stn 2141,

49°40'S, 178°52'E, off Antipodes Is., alive, 86-95 m (2

MNZ, 4 USNM); Eltanin stn 27/1850, 49°40'S, 178°53'E,

off Antipodes Is., alive, 103 m, 3 January 1967 (4 MNZ,

20 USNM).

Distribution (figure 63): Off Antipodes Islands, taken

alive at 18-103 m (minimum limit uncertain) from hard

substrata. Probably endemic.

Diet: The intestinal tract of the specimen examined con-

tained many calcareous octocoral scales (Cnidaria, Prim-

noidae), a few thecate hydroid fragments, and sand.

Remarks: Compared with C. simulans, to which it is

most closely related, C. antipodcnse differs in its deeper

shell pigmentation, more strongly convex whorls, deepK'

channeled suture, and obsolescence of PI after the fifth

teleoconch whorl. Interestingly, C. eminens, the other

Antipodes Islands endemic, also has an exceptionally deep

suture.

Calliostoma (Maurea) maui Marshall, new species

(figures 63, 70-72)

Description: Shell up to 44 mm high, broader than high,

thin, anomphalous or with a shallow umbilical depres-

sion, glossy; spire 1.24-1.59 X higher than aperture, even-

ly conical, spire angle 73-91° Protocoiuh and earliest

teleoconcli vvliorls translucent white. Suhseciuent whorls

either pale pink through unilorin translucent while outer

shell layer or pale buff or pinkish buff with yellowish or

reddish brown spiral cords and predominantly white

nodules. Protoconch ca. 400 ^m wide, surface sculpture

worn away in all available specimens. Teleoconch of up

to 9 strongly and rather evenly convex whorls, suture

well impressed, periphery rounded, base weakly convex.

Primary sculpture on spire consisting of spiral cords that

multiply by intercalation from 4 (P1-P4) to 7 (P1-P4, Sl-

S3) or occasionally 6 (S3 absent), up to 3 tertiary spirals

occasionally arise on penultimate and last adult whorl,

summit of P4 partly covered by succeeding whorls and

locally fully exposed. Nodules roundly conical; spiral in-

terspaces considerably broader than each spiral, becom-

ing finely spirally lirate. P1-P4 commencing immediate-

ly. PI at first much weaker than P2 and P3, which are

similar throughout, gradually enlarging to resemble them,

occasionally weakening and becoming obsolete on last

adult whorl. Secondary spirals gradually enlarging to

resemble primaries, SI commencing early to late on 4th

whorl, S2 on late 3rd to mid 4th whorl, S3 on mid 4th

to early 5th whorl. Base covered with spiral lirae, 2 or 3

nodular spiral cords on innermost part, other basal spiral

cords almost or entirely obsolete. Axial costae strong on

1st 3 whorls, obsolete thereafter. Aperture ovate, inner

lip thickened, parietal lip ver\ thin, outer lip thin and

simple.

Type Data: Holotype MNZ M. 87449 (height 27.3 mm,

width 31.3 mm, 7.1 teleoconch whorls); BS 561, 41°24'S,

174°33'E, Cook Strait, alive, 256-274 m, 29 September

1976, r.v. Acheron. Paratypes (6) : 41°.35'S, 175°00'E, off

Palliser Bay, alive, 256-490 m, 19-20 December 1966,

USNS Eltanin stn 1848 (1 USNM); C 703, 42°42'S,

173''37.8'E, off Kaikoura, alive, 180-140 m, 19 June 1961,

m.v. Viti (1 NZOI); E 759. 42°45'S, 17.3°40'E, off Kai-

koura, alive, 195-213 m, 31 March 1967, m.v. Viti (1

NZOI); BS 783, 43°31'S, 173°30.5'E, Pegasus Canyon,

NE of Banks Peninsula, alive, 256-293 m, 21 February

1979, r.v. Acheron (1 MNZ); 43°56.4'S, 179°25.1'W, W

of Chatham Is., alive, 303-296 m, 15 September 1987,

f.v. Chitjo Mart! 5(1 MNZ); off North Canterbury, alive,

ca 100 m, J. Sutherland (1 MNZ).

Other Material Examined: 10 specimens MNZ: BS 542,

41°08'S, 174°35.5'E, Cook Strait, subfossil, 282-293 m, 12

March 1976, r.v. Acheron (5); BS 561, 4r24'S, 174°33'E,

Cook Strait, subfossil, 256-274 m, 29 September 1976,

r.v. Acheron (4); VUZ 99, 41°34.3'S, 174°43.3'E, old shell,

274 m, 29 August 1957.

Distribution (figure 63): Cook Strait. Chatham Rise, and

off north-eastern South Island. li\ing at 140-490 m on

rugged substrata.

Diet: The intestinal tract of the holotype contained most-

ly fragments of thecate hydroids (Cnidaria), with some

indeterminate organic matter.

Remarks: C.allioslaina ntatii clo.seK resembles lightly

built forms of C. simulans and depressed forms of C.

blacki from ihe eastern C;hathain Rise in general facies.

Bruce A. Marshall, 1995

Page 107

Figure 64. Calliostoma (Maurca) mcgaloprepci: (Tomlin, 1948)^ Off Macquarie Island, 79-93 m, MNZ M. 23623 {32.9 x 34,2 mm).

Figures 65-68. Calliostoma (Maurea) simularis Marshall, 1994 65. Holotype, Pegasus Canyon, off Banks Peninsula, 446 m, MNZ

M, 87450 (30,5 x 29.0 mm). 66. Central Chatham Rise, 410 m, NZOI G259a (51,0 x 48,5 mm), 67. Challenger Plateau, 337 m,

NZOI D242 (27.0 x 27,8 mm). 68. Off Campbell Island, 188 m, NZOI D35 (52 0 x 48,0 mm). Figure 69. Calliostoma (Maurea)

antipodense Marshall, new species. Holotype, off Leeward Island, Antipodes Islands, 18-73 m, MNZ M 80434 (31.8 x 29 2 mm).

Figures 70-72. Calliostoma (Maurca) maui Marshall, new species. 70. Holotvpe, Cook Strait, 256-274 m, MNZ M. 87449 (27.3 x

31.3 mm), 71. Off North Canterbury, ca.lOO m, MNZ M. 74647 (42.4 x 45.4 mm). 72. Off Kaikoura, 139 m, MNZ M.102606 (45.0

X 48,7 mm).

Page 108

THE NAUTILUS, Vol. 108, No. 4

all three having similar development of teleoconch sculp-

ture, numerous fine spiral lirae on adult teleoconch whorls,

and a tendency toward obsolescence of all but the in-

nermost few spiral cords on the base. Calliostoma maui

closely resembles C. simulans in having all teleoconch

whorls markedly convex and in that PI at no stage be-

comes larger or more heavily nodular than the other

primary spire spirals as it does in C. blacki. It differs

from C. simulans in having considerably finer spiral

cords on the spire with finer, mostly more sharply pointed

nodules, and in being usually more broadly conical (sa

73°-91°, mean 81°, n = 7; instead of 66°-8r, mean 72°,

n = 54). During early stages of the present study I sus-

pected that C. maui might be an extreme phenotypic

variant of either C. blacki or C. simularis. Subsequent

recognition of a specimen living together with C. blacki

and C. simulans in a single sample from off Banks Pen-

insula (MNZ M.90065, 64651, 64650), however, suggests

that another closely related species is involved. Further

evidence is suggested by the occurrence in Cook Strait

of populations of C. maui that are geographically inter-

mediate between populations of C. simulans (figure 63).

Living specimens from Cook Strait, including the holo-

type (figure 72), are distinctive in having a shallow um-

bilical depression. Broken, worn, rust-stained specimens

from the floor of the Cook Strait Canyon (MNZ M. 52531,

54912) are evidently Pleistocene fossils that have been

reworked by current scour of soft sediment following

breaching of a Cook Strait land bridge, probably after

the last glaciation (Fleming, 1951, 1963; Pantin, 1957;

Marshall, 1978). They have considerably thicker shells

than Recent specimens from Cook Strait but are other-

wise identical.

Etymology: After Maui, the legendary Maori fisherman.

Calliostoma (Maurea) selectum (Dillwyn, 1817)

(Figures 73-77, 85, 118)

Chemnitz, 1795:168, pi. 196, figs 1896-97.

Trochus selectus Dillwyn, 1817:801, refers to Chemnitz, 1795:

168, pi. 196, figs. 1896-97; Wood, 1825:140, pi. 29, fig.

101a.

Trochus cunninghami Gray, 1834:600, pi. 1. fig 7; Fischer,

1876:119, pi. 39, fig. 1.

Ziziphinus cunninghami — Gray, 1843:237;

Trochus cuninghami [sic] — Philippi, 1855:281, pi. 41, fig 7

Zizyphinus cunningharnii — Reeve, 1863:pi. I, fig. 6

Zizyphinus cunninghami— HuHon. 1873:38; Huttoii 188098

Zizijphinus hodgei Hutton, 1875:458, pi. 21. New synonym

Zizyphinus decarinatus—HuHun. 1884:359 (not Ferry, 1811).

Zizyphintis ponderosus Hiilloii, 1885:322. New synonym.

Calliostoma selectum— Vihhry, 1889:335, pi. 65, figs. 73, 74

(not pi. 65, fig. 78 = C. pellucidum); Suter, 1897:281 ; Suter,

1913:146, pi. 40, fig. 4.

Calliostoma ponderosa — Hutton. 1893:69, pi. 8, fig. 75; Suter,

191.5:2.

Calliostoma /lOfigfi— Hutton, 1893:70; Suter, 1915:2.

Calliostoma carnicolor Preston, 1907: 140, pi. 8, figs. 6, 7 New

synonym

Calliostoma (Catliotro}}is) pagoda Oliver, 1926:1 12, pi 10, fig. 4.

Calliostoma (Calliotropis) cunningharnii — Oliver, 1926:112.

Calliostoma (Calliotropis) hodgei — Oliver, 1926:114.

Venustas (Venustas) cunninghami — Finlay, 1926:360, 371.

Venustas (Ver^uslas) ponderosa — Finlay, 1926:360, 371.

Venustas (Venustas) hodgei — Finlay, 1926:.360, 371.

Venustas cunninghami regifica Finla>, 1927:485, pi. 24, figs.

9, 10; Cernohorsky, 1972244.

Calliostoma (Calotropis) cunninghamii — Thiele, 1929:49.

Maurea (Maurea) cunninghami cunninghami — Powell, 1937:

64, pi. 13, fig. 1.

Maurea (Maurea) cunninghami pagoda — Powell, 1937:64.

Calliostoma (Calotropis) cuminghami [sic] — Wenz, 1938:282.

Maurea (Calotropis) cunninghami cunninghami — Powell,

1946:66, pi. 13, fig. 1.

Maurea (Calotropis) cunninghami pagoda — Powell, 1946:66.

Venustas cunninghami cunninghami — Dell, 1950:53.

Venustas cunninghami pagoda — Dell, 1950:53.

Maurea cunninghami cunninghami — Powell, 1957:88, pi. 13,

fig. 1.

Maurea cunninghami pagoda — Powell, 1957:88.

Calliostoma (Maurea) cunninghami — Sliikama & Horikoshi,

1963, pi 8, fig. 15.

Maurea (Calotropis) cttnninghami — Fleming, 1966:38.

Maurea hodgei — Fleming, 1966:38.

Maurea ponderosa — Fleming, 1966:38.

Calliostoma (Maurea) selectum — Cernohorsky, 1974:149, fig. 7.

Maurea selecta—PoweW, 1979:61, pi. 10, fig. 2; Abbott & Dance,

1982:40; Matsukuma, Okutani & Habe, 1991, pi 17, fig.

10.

Maurea pellucida — Abbott & Dance, 198240 (not Valenci-

ennes, 1846).

Calliostoma (sensu lata) selectum — Beu & Maxwell, 1990:404.

NOT Trochus selectus—Phi\ipp\, 1855 (C. pellucidum)

NOT Zizyphinus selectus— Reeve, 1863; Hutton, 1873, 1880.

1884 (C. pellucidum).

NOT Calliostoma selectum— Pi\shr\ , 1889: pi 65, fig. 78 (C.

pellucidum).

NOT Calliostoma (Calotropis) selectum— Wenz, 1938 (C. pel-

lucidum).

Type Data: Trochu-f sclectus: Lectotype (here selected)

University Zoological Museum, Copenhagen (Cernohor-

sky, 1974, pi. 149, fig. 7), "coasts of New Zealand"; Tro-

chus citnninghami: Holotype BMNH 1987047, ex J.E.

Gray collection, no locality data, = New Zealand (prob-

ably a Wellington west coast beach); Zizyphinus hodgei:

Holotype Otago Museum, Dunedin C, 54.50, " Wanganui,

in blue clay", probably Landguard Bluff (Late Pleisto-

cene, early Haweran); Zizyphinus ponderosa: Holotype

Canterbury Museum, Christchurch M.2766, "Wanga-

nui", horizon unspecified but probably Butlers Shell Con-

glomerate (Middle Pleistocene, early Castlecliffian); Cal-

liostoma carnicolor: Repository unknown, "C>lebes(.'*)

= New Zealand; Calliostoma (Calliotropis) pagoda: Ho-

lotype MNZ M.1602, oft Cape Campbell, c.37 m; Ven-

ustas cunninghami regifica: Holotype AIM 70823, off

Otago Heads, 55m.

Other Material Examined: Fossil — 20 Late Pliocene —

Early Pleistocene (Nukuniaruan) specimens in 8 lots AUG,

NZC;S; 21 Middle Pleistocene (Castlecliffian) specimens

in 18 lots AUG, MNZ, NZGS; Recent— 213 specimens in

86 lots MNZ.

Distribution (figure 85): Late Pliocene (Nukumaruan) to

Bruce A. Marshall, 1995

Page 109

Recent; North, South, Stewart and Chatham islands, 0-293

m, living at 27-274 m on sandy or muddy substrata with

shell or stones.

Diet: Gut contents of all specimens examined contained

mostly sand with fragments of either agglutinated poK -

chaete tubes or thecate hydroids (C^nidaria), together

with indeterminate organic matter. One specimen con-

tained some crustacean fragments.

Remarks: Recent specimens of this v\ell-known species

are characterised by large size (diameter up to 70 mm);

low, broadly conical spire (sa 73-94°); weakly and evenly

convex whorls, angulate or narrowly rounded periphery,

weakly to rather strongly convex base, weak axial costae

on early teleoconch whorls, similarity of P1-P3 through-

out, and the late appearance of the secondary spirals,

especially S3. Recent specimens from the South Island

east coast from Banks Peninsula southwards tend to have

more narrowly conical spires (sa 73-80°, mean 76.8°, SD

1.76, n = 20) than specimens from the North Island (sa

74-94°, mean 84.2°, SD 4.49, n = 40), but there is com-

plete intergradation between high- and low-spired forms

off the northern South Island (sa 77-90°, mean 82°, SD

4.22, n = 20) and the southern North Island. Two spec-

imens from off Jackson Bay, Westland (MNZ M, 17669)

are as broadly conical as material from the northern

South Island (sa 81° and 85°). North-south clinal grada-

tion in spire height is independent of depth. There is

considerable variation in the strength of nodules on the

spiral cords, and many specimens from Cook Strait tend

to be particularly weakly nodular (figure 74). Specimens

from the Chatham Islands (figure 77) are indistinguish-

able from mainland material (sa 76-81°, mean 79°, SD

1.87, n = 5).

The fragmentary holotype of Zizyphinus hodgei re-

sembles specimens from Landguard Bluff (early Haw-

eran. Middle Pleistocene) in shell morphology and pres-

ervation, suggesting that it may have come from that

horizon. Moreover, it is indistinguishable from Recent

specimens from off the east coast of the South Island.

The holotype of Zizyphinus ponderosa is a heavily

abraded adult shell from an unspecified horizon in the

Wanganui sequence. The shell morphology and state of

preservation closely match specimens from the Butlers

Shell Conglomerate (early Castlecliffian, Early Pleisto-

cene) (figure 76), suggesting that it originated from this

horizon. All but one of these specimens (see below) and

another from the Nukumaru Brown Sand (GS41 15, NZGS;

Nukumaruan, Late Pliocene) differ from all specimens

from later horizons in having distinctly cyrtoconoid in-

stead of shallovvly coeloconoid spires. They also have

considerably thicker shells than most Recent specimens,

as do most adult Middle Pleistocene specimens from the

overlying Pinnacle Sand (GS4022, NZGS) and Shake-

speare Cliff Sand formations (AUG 989), although the

few Recent specimens known from Stewart Island (MNZ)

are as heavily built. However, Early Pleistocene speci-

mens from the Lower Kai Iwi Siltstone (AUG 1028, 1034),

which underlies the Pinnacle Sand, are as thin as the

most lightly built Recent specimens. Specimens from the

Shakespeare Cliff Sand and earlier formations (notably

GS4022, NZGS) commonly show much later appearance

of some or all of the secondary spirals (especially S3)

than any known Recent specimens, often as late as the

last or second- to-last adult whorl. Fossils from these early

horizons are unlike Recent shells in that the spiral cords

frequently become smooth and sometimes very weak

after an initial nodular stage, becoming stronger and

nodular again on the last or second-to-last whorl or re-

maining weak and smooth throughout. A single specimen

(GS10964, NZGS) collected by A.G. Beu from the Butlers

Shell Conglomerate (Early Pleistocene) differs from oth-

ers from this horizon in being considerably thinner and

having a more broadly conical spire that is distinctly

coeloconoid rather than cyrtoconoid in outline. As in

some other specimens from this horizon, the shell is en-

tirely devoid of nodules on the remaining three whorls,

and the spiral cords are almost obsolete on the last adult

whorl. It is thus similar to some Middle Pleistocene shells

from the Kupe Formation (GS4041, 4121, NZGS) and

the thin, weakly sculptured specimens from the Lower

Kai Iwi Siltstone (AU1028, 1034, AUG). It is possible that

the more lightly built form lived in situ in a low-energy

environment, whereas the heavier shells may have washed

in from a shallower, high-energy environment or were

perhaps reworked from a slightly older horizon (presum-

ably late Nukumaruan, Early Pleistocene). This conten-

tion is supported by the fact that the heavier shells are

all worn to some extent and many are bored, whereas

the lightly built (fragile) form was in fresh condition. As

discussed by Fleming (1953:175, 176), many shells from

the Butlers Shell Conglomerate have undoubtedly been

transported, whereas others are clearly reworked from

older beds. All of the fossils otherwise resemble Recent

specimens in early teleoconch morphology. There is in-

tergradation between specimens with early- and late-

developing secondary spirals and between specimens in

which the spiral cords are weak or strong and nodular

or smooth, both within and between samples from the

Shakespeare Cliff Sand and older formations. Accord-

ingly, Z. ponderosa and C. selectum are interpreted as

forms of a single polymorphic species. Great intraspecific

variation during the Pleistocene was probably at least

partly a response to oscillating glacial-interglacial tem-

peratures and sea levels, with associated variations in

sediment type and habitat stability. Interpretation of the

fossils is potentially complicated by the likelihood of

reworking from older horizons during interglacial ma-

rine transgressions.

Calliostoma carnicolor Preston, 1907 was based on a

specimen reputedly from the Celebes, though from the

description and illustration (Preston, 1907, p. 140, pi. 8,

figs. 6, 7), it appears to be a perfectly typical, northern

specimen of C. selectum. Nothing similar has been re-

corded subsequently from the Celebes (or from else-

where for that matter), and it almost certainly represents

a mislocalised specimen of C. selectum. The original

specimen could not be traced despite extensive enquiries.

Page 110

THE NAUTILUS, Vol. 108, No. 4

Bruce A. Marshall. 1995

Page 1 1 1

Calliostoma (Maurea) pellucidum (N'aleiK'ieniies, 1846)

(Figures 78-84, 86, 119)

Trochus pellucidtis Valenciennes, 1846;pl 4, fig 2; Fischer,

1875:70, pi. 15, fig. 2.

Trochus torquatus Anton ;>i Pliilippi, 1848:126; Pliilippi, 1855:

261, pi. 38, fig. 13 (not H.C. Lea, 1846).

Trochus punciulatus — Kiener, 1847:pl. 15, fig. 2 (not Martvn,

1784).

Trochus selcclus — Philippi, 1855:261, pi 1, figs. 6, 7, pi. 38,

fig. 12 (not Dillwyn, 1817).

Zizyphinus selectiis — Reeve, 1863:pl 1, fig. 1; Hiitton, 1873:

38; Hutton, 1880:98; Huttt)n. 1884:359 (not Dillwyn, 1817).

Zizyphinus granatus — Reeve, 1863:pl. 1, fig. 2 (not Gnielin,

1791).

Calliostoma selectum — Pilsbry, 1889:pl. 65, fig. 78 only (not

Dillwyn, 1817).

Calliostoma pellucidum— Suler. 1897:281; Suter, 1913:145, pi

40, fig 3.

Calliostoma undulalum Finlay, 1923: 104, pi 10. figs 5a, b.

New synonym

Calliostoma (Calliotropis) pellucidum pellucidum — Oliver,

1926:111.

Calliostoma (Calliotropis) pellucidum spiratum Oliver, 1926:

111. New synonym

Calliostoma (Calliotropis) undulatum — Oliver, 1926:111.

Venustas (Venustas) pellucida — Finlay, 1926:360, 371.

Venustas (Venustas) undulala — Finlay, 1926:360, 371.

Maurea (Maurea) pellucida pellucida — Powell, 193764. pi 13,

fig. 2.

Maurea (Maurea) pellucida spirata — Powell, 1937:64.

Calliostoma (Calotropis) selectum — Wenz, 1938, fig. 603 (not

Dillwyn, 1817).

Venustas pellucida pellucida — Dell, 19.50:49, figs. 7, 8. 12.

Venustas pellucida spirata — Dell, 1950.50, fig 3

Maurea pellucida pellucida — Powell, 195788, pi. 13, fig. 2;

Powell, 1979:61, pi. 19, fig 4

Maurea pellucida spirata — Powell, 1957:88; Powell, 1979:61,

pi. 19, fig. 6.

Maurea (Calotropis) pellucida — Fleming, 1966:38.

Maurea (Calotropis) undulata — P'leining, 1966:38.

Calliostoma (sensu lato) pellucidum — Ben & Maxwell, 1990:

404.

Calliostoma (sensu lato) undulalum — Beu & Ma.xwell, 1990:

404.

NOT Maurea pellucida— AhhnU & Dance, 1982:40 (C. selec-

tum Dillwyn, 1817).

Type Data: Trochus pellucidufi: Lectotype (here select-

ed) and 6 paralectotypes MNHN; Trochus torquatus:

Repository of type material unknown, "locality un-

Figure 85. Map of New Zealand region showing distribution

of Calliostoma (Maurea) selectum. 200 and 1000 meter con-

tours indicated

known '. Calliostoma undulalum: Holotype NZGS TM

5001, Castlecliff, Wanganui, formation unknown (prob-

ably mid-late Castlecliffian or earK Haweran. Mid-Late

Pleistocene); Calliostoma pellucidum spiratum : Holo-

type MNZ Ml 1865, Spirits Bay, beach drift.

Other Material E.xamined: 6 Pleistocene specimens (from

unspecified horizons in the Wanganui coastal sequence)

in 5 lots MNZ, NZGS, and 235 Recent specimens in 94

lots MNZ.

Figures 73-77. Calliostoma (Maurea) selectum (Dillwyn, 1817), 73. Off Ahipara. 90 m, MNZ M. 72032 (40.0 x 47.8 mm). 74.

Off Palliser Bay. 274 m. MNZ M 12969 (42.7 x 51.0 mm). 75. Off Timaru. 110-128 m, MNZ M.7162 (47 0 x 50.0 mm). 76. Early

Pleistocene (Castlecliffian). Butler's Shell Conglomerate. Wanganui, NZGS (38.5 x 46 0 mm). 77. Off Chatham Islands, 248-236

m, MNZ M. 90061 (44 7 x .50 2 mm) Figures 78-84. Calliostoma (Maurea) pellucidum (Valenciennes, 1846). 78. Takapuna.

Auckland, low tide, MNZ M. 83646 (38.0 x 38.0 mm). 79. Paua, Parengarenga Harbour, low tide, MNZ M. 80.536 (37.7 x 34.0

mm) 80. Between Pandora Bank and Cape Maria Van Diemen, ,33 m. MNZ M 74676 (35.3 x .34 5 mm) 81. Off Rarawa Beach,

Northland, 40 m, MNZ M. 86657 (35.0 x 29.5 mm). 82. Reotahi. Whangarei Heads, low tide, MNZ M. 83643 (39.5 x 38,0 mm),

83. Akaroa Harbour entrance. 20 m, MNZ M.751.53 (43.1 x ,39.9 mm). 84. Foveaux Strait oyster beds, 31 m, MNZ M.21719 (48.0

X 47.8 mm).

Page 112

THE NAUTILUS, Vol. 108, No. 4

Figure 86. Map of New Zealand region showing distribution

of CaUiostonia (Maurea) pellucidum. 200 and 1000 meter con-

tours indicated.

Distribution (figure 86): Middle Pleistocene to Recent,

North, South and Stewart islands, living at 0-187 m on

hard substrata.

Diet: Intestinal tracts of specimens examined contained

thecate hydroids (Cnidaria) together with some indeter-

minate organic matter.

Remarks: Calliostoma pellucUliim has long been con-

fused with the superficially similar species C. waikanae,

with which it is locally sympatric throughout its range.

Highly distinctive characters of C. peUncidum include

reddish brown subsutural and peripheral maculations,

strong teleoconch sculpture, consistent appearance of So

before SI and S2, and strong P3 on early teleoconch

whorls. The spire is shallowly coeloconoid and 1-1.5 times

higher than the aperture, and the spire angle ranges from

56° to 86°. The most narrowly conical specimens occur

off eastern Northland north of Whangaroa (sa 57-69°,

mean 64°, SD 3.35, n = 22), especially in Parengarenga

Harbour (figures 79, 81). In specimens from the north-

western extremity of Northland (ligure 80), however,

including the holotype of C pellucidum spinituni Oli-

ver, 1926, the mean spire angle is rather constantly about

75°, and these broadly conical shells are closely similar

to material from south of Whangaroa (sa 68-81°, mean

75°, SD 3.45, n = 43) (figures 78, 82-84). The periphery

tends to be broadly rounded in southern specimens (fig-

ures 83, 84) and more sharply angulate in northern spec-

imens (figures 79, 81), but there is complete intergra-

dation from north to south. The Pleistocene type material

of C. undulatum Finlay, 1923 is indistinguishable from

Recent specimens of C pellucidum.

Calliostoma (Maurea) waikanae Oliver, 1926

(Figures 87-95, 99, 120, 139)

Calliostoma (Calltotropis) waikanae Oliver, 1926:11, pi. 10,

fig. 3,

Maurea (Maurea) waikanae — Powell, 1937:64.

Maurea (Calotropis) waikanae — Powell, 194666

Maurea pellucida morioria Powell, 1946:140, pi. 11, fig. 3;

Powell, 1979:61, pi. 19,fig. 5. New synonym.

Vcnustas pellucida waikanae~T)e\\, 1950:50, figs. 4,5,9,11.

Venufitas pellucida fomteriana Dell, 1950:51, figs. 1,2,10. New

synonym

Venustas pellucida morioria — Dell, 1950:53.

Venustas pellucida haurakiensis Dell, 1950:53, fig. 6. New syn-

onym

Maurea pellucida forsteriana — Powell, 1957:88; Powell, 1979:

61, pi 10, fig. 3.

Maurea pellucida haurakiensis — Powell, 1957:88; Powell, 1979:

61.

Maurea pellucida morioria— Powell, 1957;88; Powell, 197961,

pi 19, fig. 5

Maurea waikanae— PoweW, 1957:88; Powell, 1979:62, pi. 19,

fig. 7.

Type Data: Calliostoma (Calltotropis) waikanae: Holo-

type MNZ M.1603, Waikanae Beach; Maurea pellucida

morioria Holotype AIM 71037, Owenga Beach, Chat-

ham Islands; Venustas pellucida forsteriana: Holotype

MNZ M.2120, Preservation Inlet, Stewart Island. Ven-

ustas pellucida haurakiensis: Holotype NZGS TM 456,

Hauraki Gull, 46 m.

Other Material Kxamined: Fossil — AU1023, Kaikokopu

Shell Grit, coastal section, Wanganui (early Castlecliffian,

Middle Pleistocene) (1 AUG); GS4045, 4047, 4049, Kupe

Formation, coastal section, Wanganui (late Castlecliffian,

Middle Pleistocene) (3 NZGS); GS4 1 87, Shakespeare Cliff

Siltstone, coastal section, Wanganui (late (Castlecliffian)

(2 NZGS). Recent— 410 specimens in 148 lots MNZ.

Distribution (figure 99): Middle Pleistocene (early Cas-

tlecliffian) and possibly Middle Pliocene (Waipipian).

Recent off North, South, Stewart, Snares and Chatham

islands, and Mernoo Bank, 0-549 in, taken alive at 18-

549 m on bryozoan/shell sulistrata.

Diet: Intestinal tracts of specimens examined contained

fragments of thecate hydroids (Cnidaria) together with

indeterminate organic matter

Remarks: Calliostoma waikanae occurs throughout the

geographic range of C. pellucidum, and the two species

Bruce A. Marshall, 1995

Page 113

have been thoroughly confused taxonomically because

of their variability and superficial similarity. Calliostoma

waikanae was treated as a subspecies of C. pellncidum

by Dell (1950), whereas forms of C. waikanae have been

consistently treated as "subspecies of C, pellucidum,

namely Maurea pcllucida rnorioria Powell, Venustas

pellucida forsteriana Dell, and V. pcllucida haurakiensis

Dell, all of which are here interpreted as local populations

of C waikanae. Although they are similar in shape, size,

colour, and colour pattern, C. waikanae is easily distin-

guishable from C. pcllucidum by the constant appear-

ance of SI and S2 well before S3 instead of vice versa

(figures 119, 120). C. waikanae differs further in having

finer pustules on the side of the foot and tends to have

a more lightU pigmented shell with finer, more finely

beaded spiral cords, and the summit of P4 is usually

partly or entirely exposed on the spire instead of being

almost entirely covered by succeeding whorls. In other

words the suture is bordered by P4 in C. waikanae and

by So in C. pellucidum. S3 is seldom nodular in C.

waikanae, but constantly becomes strongl)' nodular in

C. pellucidum from an early stage of growth.

Most specimens from off the Three Kings Islands and

North Cape (figure 88), the north-western North Island

southward to off Kawhia (figure 89), and the north-east-

ern North Island as far south as East Cape (figure 87)

have narrower and correspondingly higher spires than

specimens from further south (sa 68-76°, mean 72°, n =

27, as against 73-92°, mean 82°, n = 76)(figures 90-95).

They differ further in that P3 frequently remains con-

siderably stronger than PI and P2 onto later whorls, often

until the second to last adult whorl, and typically sur-

mounts a prominent, narrow suprasutural bulge together

with S3 and P4. Such specimens intergrade completeK'

with specimens in which the primaries become similar

through progressive enlargement of PI and P2 and weak-

ening of P3 at an earlier stage of growth and in which

the suprasutural bulge is correspondingly weak or absent.

Weakness or absence of a suprasutural bulge is a char-

acteristic of type and topotypic material of V. p. hau-

rakiensis and most material from southern localities,

though some specimens from Fiordland (figure 94) have

a well-developed and persistent peripheral bulge. Spec-

imens from north of Cape Reinga attain larger size (di-

ameter up to 52.5 mm) and are more darkly and exten-

sively pigmented than specimens from elsewhere off the

northern North Island but are otherwise identical.

Calliostoma waikanae is exceptionally variable in the

area off the west coast of the southern North Island and

in Cook Strait. In many specimens from this area, in-

cluding the holotype of C. waikanae, the spiral cords

and nodules weaken with increasing shell size, and the

nodules may become almost obsolete (figure 91). There

is, however, complete gradation between weakly and

strongly sculptured forms both w ithin and between pop-

ulations. Moreover, these specimens show exceptionally

wide variation in the development of S3, which may fuse

with P4, become as large as P3, remain weak throughout,

or entirely fail to appear. Again there is complete mosaic

intergradation between the extremes. Pronounced vari-

ation in shell morphology in this area is probably at least

partly the result of repeated divergence and remixing of

populations isolated to the east and west of Cook Strait

land bridges that formed during Pleistocene glaciations

(Fleming, 1962). This explanation probably also accounts

for extreme local variation in direct developers such as

buccinids of the genera Penion Fischer and Cominella

Gray.

The only significant differences that I am able to detect

between specimens from the Cook Strait area and the

southern South Island (Jorsteriana) is the more general

tendency for the southern form to have a rounded pe-

riphery at maturity and a slightly narrower spire angle

(73-86°, mean 79°, n = 23 as against 75-92°, mean 82°,

n = 53). Again, however, there is complete intergrada-

tion both within samples and clinally between the north-

ern and southern forms.

Specimens from the Chatham Islands {rnorioria} (fig-

ure 95) are indistinguishable from mainland specimens.

Existing collections suggest that C. waikanae has a

rather patchy distribution (figure 99) and the differences

between specimens from off north-eastern North Island/

Kawhia and the Wanganui/Cook Strait area are certainly

greater than between specimens from Cook Strait/Kai-

koura area and East Otago, suggesting that the northern

population may be more strongly isolated than the other

mainland populations are from each other. In view of

the tendency for this and other New Zealand calliosto-

matids to form morphologically discrete local popula-

tions and to show pronounced clinal geographic and

bathymetric variation, it seems clear that all of the nom-

inate forms should be interpreted as local populations of

a single polymorphic species.

Middle Pleistocene specimens from the Kaikokopu Shell

Grit (AU1023, AUG) and the Kupe Formation, Wang-

anui (GS4045, 4047, 4049, NZGS), differ from Recent

specimens in being thicker at maturity and in the early

appearance and more rapid enlargement of the spiral

cord between the suture and PI, which develops to re-

semble PI. The two subadult specimens from the Shake-

speare Cliff Siltstone (GS4187, NZGS) are very lightly

built and have sculpture that becomes obsolete on the

sixth w horl and gradually reappears on the eighth w horl.

All of these fossils are indistinguishable from Recent spec-

imens in early teleoconch morphology, and they are in-

terpreted as C. waikanae that is both undergoing genetic

drift through time and varying in response to changing

sea temperature and bathymetry. A single fragmentary

specimen from below the Te Aute Limestone, 2.5km

south-east of Takapau, Takapau Surve) District, Hawke

Bay (GS2320, NZGS) — age either Nukumaruan or (more

likely) Waipipian (Middle Pliocene) (A.G. Beu, pers.

comm.) — is even more similar to Recent C. waikanae

than the Pleistocene material, but better material is re-

quired to ascertain its status.

Calliostoma (Maurea) turnerarum (Powell, 1964)

(Figures 96, 100, 121)

Page 114

THE NAUTILUS, Vol. 108, No. 4

Bruce A. Marshall, 1995

Page 115

Maurea turnerarum Powell, 196411. pi 3, fig. 1-3; Powell,

1979:62, pi. 19, fig. 9; Horikoshi, 1989:pl. 4, figs. 8,9.

Type Data: Holotype AIM 71239, off Mayor Island, Bay

of Plenty, 366 m.

Other IVIaterial Examined: 37 specimens in 19 lots MNZ.

Distribution (Bgure 100): Three Kings Islands, off Ninety

Mile Beach, and north-eastern North Island as far south

as Cape Runaway, 186-805 m, taken alive at 312-529 m

on muddy substrata with shells.

Diet: The intestinal tracts of specimens examined con-

tained numerous fragments of thecate hydroids (Cni-

daria) together with some indeterminate organic matter.

Remarks: Calliostoma turnerarum resembles C. wai-

kanae in gross fades, and the two species are undoubt-

edly closely related. Calliostoma turnerarum differs in

having a more lightly built shell with finer sculpture on

adult whorls and in having pale, wavy axial bands, within

which the strongest spiral cords are streaked deep reddish

or vellowish brown. Compared with specimens of C.

waikanae from within its geographic range, C. turner-

arum is more broadly conical, the spire angle ranging

from 83° to 95° (mean 87°, n = 5) instead of 68° to 75°

(mean 72°, n = 27). Although the early teleoconch sculp-

ture is similar in both species, P3 remains similar to P2

in C. turnerarum, whereas P3 becomes markedly stron-

ger and more strongly nodular in C. waikanae.

Calliostoma (Maurea) penniketi Marshall, new species

(Figures 97, 98, 100, 123, 140)

Description: Shell up to 57 mm high, about as high as

broad at maturity, spire 1.19-1.78 x as high as aperture,

spire outline evenly conical or shallowly coeloconoid, last

whorl slightly contracting at maturity, spire angle 65-

78°, rather thin but strong, anomphalous throughout. Pro-

toconch and 1st teleoconch whorl white. Ground colour

on subsequent whorls pale yellowish brown or buff white,

rich yellowish brown on last adult whorl of a few spec-

imens. Spiral cords spotted and spire whorls typically

maculated with yellowish or reddish brown, maculations

absent from a few specimens. Spiral cords becoming

spotted after 2nd-4th w horl, typically on each alternate

spiral on early whorls and on each spiral on later whorls,

nodules predominantly white on last 2 adult whorls, most

Figure 99. Map of New Zealand region showing distribution

of Calliostoma (Maurea) waikanae. 200 and 1000 meter con-

tours indicated.

deeply pigmented on maculations. Maculations irregu-

lar, subquadrate, occupying rather discrete subsutural

and suprasutural zones, becoming obsolete on last whorl.

Subsutural maculations typically more darkly pigmented

than suprasutural ones, almost entirely traversing early

whorls, narrowing to occupy adapical half of later whorls,

numbering 7-9 on adult penultimate whorl. Suprasutural

maculations small, occupying narrow band throughout,

numbering 13 or 14 on adult penultimate whorl. Inner

lip white, aperture nacreous. Protoconch 370 nm wide.

Figures 87-95. Calliostoma (Maurea) waikanae Oliver, 1926. 87. Paratype of Venustas pellucida haurakiensis Dell, 1950, Hauraki

Gulf, 46 m, MNZ M.15910 (36.9 x 38.3 mm). 88. Off North Cape, 178-167 m, MNZ M.74670 (51.0 x 51.3 mm). 89. Off Kawhia,

83 m, MNZ M.74656 (36.5 x 40.0 mm). 90. Erie Bay, Tory Channel, Wellington, c. 20 m, MNZ M. 17931 (41.0 x 42.0 mm). 91.

Between Mana Island and Cape Jackson, Cook Strait, 256-186 m, MNZ M. 49913 (32.0 x 36.0 mm). 92. Off Oamaru, c. 73 m,

MNZ M. 102589 (50.5 x 510 mm). 93. Foveaux Strait oyster beds. 37-46 m, MNZ M. 80567 (48.0 x ,52.5 mm). 94. Bligh Sound,

Fiordland, c, 20 m, D. Gibbs collection (42.5 x 41.2 mm). 95. Off Kaingaroa, Chatham Islands, craypots, MNZ M.92395 (36.2 x

39,0 mm). Figure 96. Calliostoma (Maurea) turnerarum (Powell, 1964), North of Three Kings Islands, 348-312 m, MNZ M. 71984

(47.5 X 56.7 mm). Figures 97, 98. Calliostoma (Maurea) penniketi Marshall, new species, 97. Holotype, off Three Kings Islands,

c. 55 m, MNZ M.74817 (57.0 x 53.0 mm). 98. Ranfurly Bank, East Cape, 76-71 m, MNZ M. 64782 (27.2 x 27.0 mm).

Page 116

THE NAUTILUS, Vol. 108, No. 4

Figure 100. Map of New Zealand region showing distribution

of Calliostoma (Maurea) penniheti (circle) and C. (M.) tur-

nerarum (star). 200 and 1000 meter contours indicated.

sculptured with a network of fine threads that enclose

roughly hexagonal spaces, terminal varix strong. Teleo-

conch of up to 10 whorls, 1st 2.5-3 whorls convex, sub-

sequent whorls more or less flat, last 2 whorls weakly

convex. Periphery tightly rounded, becoming rather

broadly rounded on last whorl; base almost flat, becoming

weakly rounded at maturity First 3 whorls sculptured

with spiral cords, axial riblets, and a few crisp spiral

threads, axials and spiral threads weakening and vanish-

ing over next whorl. P1-P4 commencing immediately,

PI thread-like on 1st whorl, gradually enlarging to re-

semble P2 and P3, which are similar throughout; P4

peripheral, covered by succeeding whorls, becoming ex-

posed by descent of last whorl. SI- S3 commencing on

3rd whorl, enlarging to resemble P1-P3; tertiary spirals

commencing on 5th whorl or late on 4th whorl, 1 in each

interspace and between IM and suture, enlarging to re-

semble secondaries and primaries. Additional finer spi-

rals in each interspace on last few whorls. Spirals about

as high as broad with considerably wider interspaces and

prominent rounded nodules on ist 5 whorls, then flat-

tening, becoming smooth, and broadening until sepa-

rated by shallow grooves; finally becoming strong, crisply

nodular and widely spaced on penultimate or last adult

whorl. P3 more strongly nodular than PI and P2 on 3rd

and 4th w horls. Adult base with about 18-20 major, spot-

ted, nodular spiral cords, additional finer threads in most

interspaces. Aperture subquadrate to subcircular. Outer

lip thin at rim, thickened within, strongly so at base.

Inner lip a spreading glaze. Parietal glaze thin, restricted

to a narrow outer spiral band.

Type Data: Holotype M.74817 (height 57.0 mm, width

53.0 mm, c. 10 teleoconch whorls) and 2 paratypes MNZ,

off Three Kings Is., c.55 m, alive in craypots, coll. A.D.

Howell, pres. M. Sanson. Paratypes (9) ; C763, 33°58'S,

172°17.6'E, off Three Kings Is., dead, 73-99 m, 18 Feb-

ruary 1962, m.v. Viti (1 NZOI); B93, 34°00'S, 172°30'E,

alive, 55-91 m, 22 September 1958, m.v. Viti (1 NZOI);

BS392, 34°08.5'S, 172°11'E, dead, 102 m, 18 February

1974, r.v. Acheron (1 MNZ); BS902 (0648), 34°10.5'S,

I72°11.4'E, dead, 153 m, 1 February 1981, r.v. Tangaroa

(1 MNZ); BS901 (0647), 34°14.1'S, 'l72°09'E, dead, 192-

202 m, 1 February 1981, r.v. Tangaroa (1 MNZ); BS910

(0656), 34°18.8'S, 'l72°18.5'E, dead, 93-88 m, 2 February

1981, r.v. Tangaroa (2 MNZ); BS911 (0657), 34°20.2'S,

172°21.8'E, dead, 121 m, 2 Februarv 1981, r.v. Tangaroa

(1 MNZ); BS769 (R127), 37°33.2'S, 'l78°50.3'E, Ranfurly

Bank, East Cape, alive, 76-71 m, 25 January 1979, r.v.

Tangaroa; BS678 (R36), 37°36.3'S, 178°53.1'E, dead, 74

m, 17 January 1979, r.v. Tangaroa (1 MNZ).

Other Material Examined (mostly juveniles): Off Three

Kings Islands — 50 specimens in 1 1 lots MNZ; Ranfurly

Bank, East Cape — 8 specimens in 4 lots MNZ.

Distribution (figure 100): Three King Islands and Ran-

furly Bank, East Cape, 55-622 m; taken alive at 55-76

m on rugged, bryozoan/shell substratum with sponges,

corals, gorgonians, etc.

Diet: The alimentary tract of a paratype from off the

Three Kings Islands at 55 m contained fragments of

thecate hydroids and indeterminate organic material.

Remarks: Calliostoma penniketi is closely related to C.

waikanae and C. pelhicidum, which it resembles in size,

colour, and colour pattern. It is readily separable from

them by the finer, more numerous nodules and weaker

P3 on the early spire whorls, and in passing through an

intermediate growth stage in which the spiral cords be-

come low, broad, and smooth. Specimens from off the

Three Kings Islands and East Cape are indistinguishable.

Calliostoma penniketi may eventually be discovered at

intermediate localities, which are as yet inadequately

sampled at appropriate depths on hard substrata.

Etymology: Named in honour of the late J. R. (Bob) Pen-

niket, a connoisseur of New Zealand Calliostomatidae,

and in recognition tor his fostering ot New Zealand nial-

act)logy, both amateur and professional.

Subgenus Olitkaia Ikcbe, 1942

Otukaia Ikebc, 1942:277. Type species (by original designa-

Bruce A. Marshall, 1995

Page 117

Figures 101, 102. Calliostoma (Otukaia) alertae Marshall, new name. 101. Off C^ape Karikari, Northland, 743 m, NZOI F913

(27,0 X 25 7 mm). 102. Off Bounl> Islands, 47.5 ni. NZOI 1704 (31.5 x 25.5 mm). Figure 103. Calliostoma (sensu lato) limatulum

Marshall, new species. Holotype, King Bank, north-east of Three Kings Islands, 128 m, MNZ M. 90129 ( 8.05 x 5 50 mm) Figure

104. Calliostoma (sensu lato) onustiim Odhner, 1924 King Bank, north-east of Three Kings Islands, 128 m, MNZ M. 50551 (9.85

X 5 50 mm) Figure 105. Calliostoma (sensu lato) kopua Marshall, new species Holotype, off Cape Campbell, 454-424 m, MNZ

M. 59698 (5.35 x 4.95 mm).

tion): Calliostoma kiheizicbisu Otuka. 1939; Recent, Ja-

pan.

Alertalex Dell, 1956:46. Type species (by original designation):

Alertalex blacki Dell, 1956 {Calliostoma alertae Marshall,

new name); Recent, New Zealand

Remarks: Most members of this group are characterised

by a moderate to large-sized shell (height up to 36 mm);

strong, smooth or weakly beaded P2 and P3; and the

extreme weakness of PI. All have a highly nacreous shell

due to transiucency of the colourless outer shell layers,

and the group (as currently interpreted) occurs world-

wide at bathyal depths The radula, jaw, external anat-

omy, and development of sculpture on the early teleo-

conch are essentially similar to those in Calliostoma (sen-

su strict a), and I am unable to justify segregation of the

group at genus level. Since I am not able to justify place-

ment of Otukaia as a synonym of Calliostoma either, I

follow McLean and Andrade (1982) in interpreting it as

a subgenus.

Calliostoma (Otukaia) alertae Marshall, new name

(Figures lUl, 102, 108, 122, 141)

Alertalex blacki Dell, 1956:46, figs 61, 120, 260; Dell 196275

(secondary homonym of Venustas blacki Powell, 1950)

Otukaia hlack'i—DeW. 1963208; Powell, 197963, pi. 19, fig.

13.

Page 118

THE NAUTILUS, Vol. 108, No. 4

Figures 106, 107. Radula of holotype of Calliostoma (sensu lata) kopua Marsfiall, new species. 106. Central and lateral teetfi.

107. Marginal teeth, innermost tooth indicated (x). Scale bars = 10 fjm.

Type Data: Holotype MNZ M.9767 and 1 paratype MNZ:

44°04'S, 178°04'W, Chatham Rise, alive, 476 m, 10 Feb-

ruary 1954, m.v. Alert. Paratypes (4 MNZ): 43°42'S,

179°55'E, Chatham Rise, alive, 512 m, 24 January 1954,

m.v. Alert.

Other Material Examined: 17 specimens in 12 lots MNZ,

30 specimens in 20 lots NZOI.

165

I : ^

175

I

Distribution (figure 108): Off Cape Brett southward to

off The Snares, Challenger Plateau, Chatham Rise, Boun-

ty Plateau, and Pukaki Rise (34°43.5'S-49''02'S), living at

280-861 m on muddy substrata with shells.

Diet: Gut contents of the few specimens e.xamined com-

prise much indeterminate organic matter together with

fragments of thecate hydroids (Cnidaria).

Remarks: Since Otukaia is here interpreted as a subgenus

of Calliostoma, C. {Otukaia) hlacki (Dell, 1956) becomes

a homonym of the prior C. (Matirea) hlacki (Powell,

1950). In the absence of junior synonyms it is here ren-

amed C. (O.)alertae. Should Otukaia prove to be worthy

of genus-group status, the specific name will revert to

the original (ICZN Art 59d).

This species is strongly characterised by its iridescent,

weakly nodular shell, very strong P2 and P3, the late

appearance and extreme weakness of PI, and the strong

development of S2, which enlarges to resemble P2.

Calliostoma alertae shows slight southward narrowing

of the spire angle, which ranges from 61° to 70° (mean

65.7°, u = 7) in material from north of East Cape (figure

101) and 55° to 60° (mean 57.8°, n = 7) in specimens

from the Bounty Platform (figure 102). Specimens from

these extreme northern and southern populations also

have respectively the most lightly and heaviK built shells.

Material from geographically intermediate localities are

intermediate in spire angle and shell thickness.

Calliostoma (sensu lata) onusluni Odhner, 1924

(Figures 104, 108, 124, 142)

Figure 108. Map of New Zealand region showing distributions

ol (.'(illiiistonia (Otukaia) alcrlac (dot), both C (slat.) lima-

litlutu and ('. (.s. lat .) (inttsliuii (star), and C. (.s. lat.) kopiia

(open circle). 200 and KHK) meter contours indicated.

Bruce A. Marshall, 1995

Page 119

Figures 109-1 17. Earl) whorls of Calliostoma species. Figure 109. Calliostoma (Maurea) gibbsorum Marshall, new species. Off

Three Kings Islands, craypot, MNZ M. 74663. Figure 1 10. Calliostoma (Maurea) tigris (Gmelin, 1791 ). Whangaroa Harbour entrance,

20 ni, MNZ M. 41060. Figure 111. Calliustonia (Maurea) regale Marshall, new species. Holotype, off Three Kings Islands, 153 m,

MNZ M. 86730, Figure 112. Calliostoma (Maurea) aupouriarmm Marshall, new species. Holotype, off Three Kings Islands, 2.52 m,

MNZ M. 86731. Figure 1 13. Calliostoma (Maurea) spectabile (A. Adams, 18.55). Off Auckland islands, 42-44 m, NZOI D71. Figure

114. Calliostoma (Maurea) joveauxarnuii (Dell, 1950), Off Otago Peninsula, 220 m, MNZ M.8939. Figure 115. Calliostoma

(Maurea) blacki (Powell, 19.50). Off Otago Peninsula, 476-549 m, MNZ M.8891. Figure 116. Calliostoma (Maurea) simulans

Marshall, 1994. Paratype, wall of Pegasus Canyon, off Banks Peninsula, .329-183 m, MNZ M. 64654 Figure 117. Calliostoma

(Maurea) antipodense Marshall, new species. Paratype, off Antipodes Islands, 103 m, MNZ M. 80469. Scale bars = 1mm

Calliostoma onustum Odhner, 192416, pi 1, fig 4.

Calliostoma (Maurea) onustum — Oliver, 1926:108.

Fautor onustus—Fin\si\. 1926:360; Powell, 1979:63, pi. 19, fig.

12.

Description: Shell up to 6.6.'3 mm high, of moderate

thickness, considerably higher than broad; spire narrowly

and evenly conical, up to .3.4 x higher than aperture;

juveniles with minute umbilical chink, larger specimens

anomphalous. Iridescent nacreous through translucent

outer shell layer, which is colourless when fresh, dead

specimens white, base and P4 pale buff in some speci-

mens, protoconch and 1st teleoconch whorl pink in some

specimens. Protoconch .320-.'3.30 /um wide, sculptured with

fine threads that enclose hexagonal spaces, terminal vari.x

strong, rounded. Teleoconch of up to 6.50 whorls, 1st

whorl rounded, next 2 angulated at P.3; subsequent whorls

rounded over abapical half, weakly concave adapically;

periphery narrowly rounded, base more or less flat. First

Page 120

THE NAUTILUS, Vol. 108, No. 4

Figures 118-126. Karly whorls of Calliusluniu species. Figure 118. Calliosluma (Maurea) aclccttim (Dillw\n, 1817). Tasman

Bank, Golden Bay, 26-24 m, MNZ M. 50.508 Figure 1 19. Calliostoma (S4aureaj pellucidum (Valenciennes, 1846). Off Stephens

Island, (^ook Strait, 183-187 m, MNZ M 50269. Figure 120. Calliostoma (Mattrea) waikanae Oliver, 1926 Pelorus Sound mouth,

29 rn, MNZ M. 51450. Figure 121. Calliostoma (Maurca) turncrarum (Powell, 1964) Off Great Island, Three Kings Islands, 440

m, MNZ Ml 18.380 Figure 122. Calliostoma (Ctiikaia) alertae Marshall, new name Wall of Pegasus Canyon, off Banks Peninsula,

1(K)6-512 m, MN'Z M 52781 Figure 123. Calliostoma (Mattrea) pennikvti Marshall, new species. Off Three Kings Islands, 187 m,

MNZ M. 34246 Figure 124. Calliostoma (scnsu lato) onustnm Odhner. 1924 Off Three Kings Islands, King Bank, north-east of

Three Kings Islands, 128 rn, MNZ M 50.551. Figure I2.'>. Calliostoma (scnsu lato) kopua Marshall, new species. Holot\pe, off Cape

Campbell, 454-424 m, MNZ M. 59698 Figure 126. Calliostoma (sensii lato) limalitlum Marshall, new species Holotype, King

Bank, north-east of Three Kings Islands, 128 m, MNZ M. 90129. Scale bars = Inim.

c. 0.3 whorl demarcated by growth scar, not .sculpturally

differentiated from .succeeding half whorl. First 1.5 whorls

with fine, crisply defined spiral threads. Spiral cords

rounded, sharply shelved adapically, nodular where tra-

versing axial costae, multiplying by intercalation from 3

{P2-P4) to 7 (P1-P4, S1-S3); interspaces concave, about

as wide as each spiral. Spiral cords enlarging Irom tlie

spiral threads. PI commencing later than P2-P4, which

are similar throughout, PI as large as them after 2nd

v\ liorl. Summit of P4 exposed on spire throughout. Sec-

ontlary spirals ra[)itlly enlarging to resemble primaries;

SI commencing on late 2rid to late 3rd whorl, S2 on mid

Bruce A. Marshall, 1995

Page 121

Figures 127-135. Protoconchs of Calliostoina species Figure 127. Calliostoma (Maurea)tigris (,(.inrliii, 1791). Whangaroa Harbour

entrance, 20 m, MNZ M. 41060 Figure 128. Calliostoma (Maurea) punctulalum (Martyn, 1784) Cook Strait, 256-254 m, MNZ

M. 54911 Figure 129. Calliostoma (Maurea) granti (Powell, 1931). Off Stephens Island,' Cook Strait, 183-187 m, MNZ M.50266.

Figure 130. Calliostoma (Maurea) gibbsonim Marshall, new species. Off Three Kings Islands, 102 m, MNZ M. 34249. Figure 131.

Calliostoma (Maurea) osbornei Powell, 1924 Off Wanganui, 33-35 m, MNZ M 118381 Figure 132. Calliostoma (Maurea) regale

Marshall, new species. Off Three Kings Islands, 102 ni, MNZ M. 80699 Figure 133. Calliostoma (Maurea) aupourianum Marshall,

new species. Middlesex Bank, north-west of Three Kings Islands, 246-291 m, MNZ M 80675 Figure 134. Calliostoma (Maurea)

spectabile (A Adams, 1855) Off Auckland Islands, 42 m, NZOI D71 Figure 135. Calliostoma (Maurea) foveauxanum (Dell,

19.50) Off Otago Peninsula, 220 m, MNZ M.8939. Scale bars = 100 ^m.

2nd to early 3rd whorl, S3 on late 1st to mid 2nd whorl.

Axial costae commencing late on 1st whorl, gently pro-

socline, non-collabral, entirely traversing all subsequent

whorls, very strong between P2 and P4, numbering 16-

20 on .5th whorl. Base with 7-9 similar, rounded spiral

cords, their outer edges beveled, at first smooth, becom-

ing weakly nodular with increasing shell size; interspaces

wider than each spiral in most specimens but narrower

in a few specimens Collabral growth lines prosocline on

spire, opisthocyrt on base. Aperture subquadrate, inner

and outer lips thin and simple, parietal glaze very thin.

Type Data: Lectotype (Odhner s described and illus-

trated specimen here selected) and 1 paralectotype. Zoo-

logical Museum, Copenhagen : 10 miles NW of Cape

Maria van Diemen, dead, 91 m, 4 January 1915.

Other Material Examined: 48 specimens in 12 lots MNZ.

Distribution (figure 108): Off Three Kings Islands and

off Cape Reinga (33°57.0'S-34°25.0'S), 55-310 m, taken

alive at 102-202 m on rugged, bryozoan/shell substratum

with sponges, gorgonians, and hydroids.

Diet: L'nknown.

Remarks: Among previously described ta.xa, Calliosto-

ma onustum most closely resembles the southern Aus-

tralian species C. retiarium Hedley & May, 1908, from

which it differs in having a flattened or concave shoulder

on the teleoconch whorls and broader spiral cords. The

smaller specimen (paralectotype) recorded by Odhner

(1924) is an immature specimen of Thysanodonta wai-

rua Marshall, 1988 (Calliostomatidae : Thysanodontin-

ae).

Marshall (1995) will refer this species to a new genus

based on the highly distinctive shell morphology and

Page 122

THE NAUTILUS, Vol. 108, No. 4

Figures 136-144. Protoconchs of Ca//tos(oma species. Figure 136. Calliostoma (Maurea) blacki {PoweW, 1950). Off Otago Peninsula,

476-549 m, MNZ M 8891. Figure 137. Calliostoma (Maurea) simtilans Marshall, 1994. Paratype, wall of Pegasus Canyon, off

Banks Peninsula, 329-183 m, MNZ M. 64654. Figure 138. Calliostoma (Maurea) antipodense Marshall, new species Paratype, off

Antipodes Islands, 103 ni, MNZ M. 80469 Figure 139. Calliostoma (Maurea) waikanae Oliver, 1926. Pelorus Sound mouth, 29 m,

MNZ M. 51450. Figure 140. Calliostoma (Maurea) penniheti Marshall, new species. Off Three Kings Islands, 187 m, MNZ M. 34246.

Figure 141. Calliostoma (Otukaia) alertae Marshall, new name. Wall of Pegasus Canyon, off Banks Peninsula, 1006-512 m, MNZ

M. 52781. Figure 142. Calliostoma (sensu lato) onustum Odhner, 1924. Off Three Kings Islands, 102 m, MNZ M. 34251, Figure

143. Calliostoma (sensu lato) kopua Marshall, new species. Holotype, off Cape Campbell, 454-424 m, MNZ M. 59698 Figure 144.

Calliostoma (sensu lato) limatulum Marshall, new species. Off Three Kings Islands, 102 m, parat>pe, MNZ Ml 18382. Scale bars

= 100 Mm.

degenerate radula (figures 106, 107). C. retiarium and

the two new taxa described below are congeneric.

Calliostoma (sensu lato) limatulum Mar.sliall, new spe-

cies

(Figures 103, 108, 126, 144)

Description: Shell up to 6.60 niin high, of niotleratc

thickness; liigher than broad, considerably so at maturity;

some juveniles with minute umbilical chink, adults an-

omphalous; spire narrowly and evenly conical, up to 2.9

X higher than aperture. Iridescent nacreous through

translucent outer shell layer, which is colourless when

fresh, dried specimens becoming white; occasionally with

a few scattered irregular patclies of pale buff; protoconcli

and 1st 1 or 2 teieoconch whorls sometimes pink. Pro-

tocorich 330 ^m wide, sculptured with fine threads that

enclose hexagonal spaces, terminal varix strong, rounded.

Teieoconch of up to 6.10 whorls, 1st whorl rounded,

subsequent whorls flattened adapically, angulate at P3

on 2nd and 3rd whorls, rounded thereafter, periphery

narrowly rounded, base more or less flat First c 0 3 w horl

demarcated by growth scar, not sculpturally ilifterenti-

ated from succeeding half whorl. First 1.5 whorls with

fine, crisply defined spiral threads. Spiral cords enlarging

from fine threads, rounded, sharply shelved adapically,

multiplying b\ intercalation troin 3 (P2-P4) to .5 or oc-

casionally 6 (P1-P4, S2±S1), interspaces narrower than

each spiral, P3 and P4 becoming most closely spaced.

S[)iral cords enlarging from fine threads. PI commencing

later than P2 and P3, alter mid 2nd-4th whorl as large

as P2; P2 and P3 of similar size on 1st whorl, P3 stronger

thereafter; summit of P4 exposed on spire throughout.

When present, SI either developing in parallel with PI

and rescnibliiig it throughout or commencing as late as

mid 4th whorl and enlarging to resemble PI; S2 com-

Bruce A. Marshall, 1995

Page 123

mencing on late 1st or on 2nd whorl, as large as PI and

P2 after late 3rd whorl. Spirals nodular on early whorls,

strongly so on P3 and P4, on 4th or 5th whorl nodules

weakening and vanishing from all spirals, v\hith are

smooth thereafter. Early spire whorls either entirely tra-

versed by strong axial costae, or costae strong between

S2 and P4 and weaker adapicalK . A,\ials obsolete after

3rd-5th whorl. Base with 7 or 8 broad, smooth, rounded

spiral cords w ith interspaces considerably narrower than

each spiral, occasionally with up to 12 finer cords. Col-

labral growth lines prosocline on spire, opisthocyrt on

base. Aperture subquadrate, inner lip thin.

Type Data: Holotype MNZ M. 90129 (height 8.05 mm,

diameter 5 50 mm, 7.00 teleoconch whorls), BS 904 (0650),

33°57.0'S, 172°19.0'E, King Bank, NE of Three Kings Is.,

dead, 128 m, 1 February 1981, r.v. Tangaroa. Paratypes

(31 MNZ): BS 898 (0644), 34°01.2'S, 17r44.4'E, dead,

206-211 m, 31 January 1981, r.v. Tangaroa (2); BS 637

(P485), 34°05.0'S, 172°24.6'E, dead, 200 m. 24 June 1978,

r.v. Tangaroa (1); BS 392, 34°0S.5'S, 172°11.0'E, alive,

102 m, 18 February 1974, r.v.. Ac/if ron (8); BS 901 (0647),

34°14.1'S, 172°09.0'E, ahve, 192-202 m, 1 February 1980,

r.v. Tangaroa (11); BS 911 (0657), 34°20.2'S, 172°21 8'E,

alive, 121 m, 2 February 1981, r.v. Tangaroa (6); BS 912

(0658), 34°22.8'S, 172°24.6'E, dead, 121 m, 2 Februarv

1981, r.v. Tangaroa (2); BS 631 (P441), 34°24.0'S,

172°16.8'E, dead, 120 m, 20 June 1978, r.v. Tangaroa

(1).

Other IMaterial Examined: 72 specimens in 10 lots MNZ.

Distribution (figure 108): Off Three Kings Islands and

off Cape Reinga, 91-805 m, taken alive at 102-310 m on

rugged, bryozoan/shell substratum with sponges, gor-

gonians, and hydroids.

Diet: I'nknown.

Remarks: Calliostorna limatuliau is closely similar to

the sympatric species C. onustuni. differing principally

in being more broadly conical, in having axial costae

confined to the early spire whorls, in having an extra

spiral cord on the spire (S3), and in that the spiral cords

on the spire become broader and smooth with increasing

shell size. The two species have identical protoconch and

first teleoconch whorl morphologies, and they are similar

in shell structure and shell size relative to the number of

whorls. Their geographic and bathymetric distributions

are similar, and they commonly occur together in dredge

samples. I have not overlooked the possibility that they

may be forms of a single sexually dimorphic species, but

without knowledge of the reproductive anatomy it is

impossible to tell.

Calliostorna (sensu lato) kopua Marshall, new species

(Figures 105, 106, 107, 125, 143)

Description: Shell up to 5,35 mm high, slightly higher

than broad, with minute umbilical chink; spire evenly

conical, up to 1.78 x higher than aperture. Iridescent

nacreous through colourless, translucent outer shell layer.

Protoconch 320 jitm wide, sculptured with fine threads

that enclose hexagonal spaces, terminal varix strong,

rounded. Teleoconch of up to 4.70 whorls, 1st whorl

convex, subseciuent whorls flat sided, periphery angulate;

base almost flat at first, becoming weakly convex. A growth

scar almost immediately after protoconch. First 1.5 whorls

with fine, crisply defined spiral threads. Spiral cords on

spire angulate in section, sharply beveled adapically, with

rounded conical nodules where traversing axials, mul-

tiplying by intercalation from 3 (P2-P4) to 6 (P1-P4, SI,

S2), summit of P4 exposed on spire; interspaces concave,

broader than each spiral, P3 and P4 becoming closer

than others with increasing shell size. PI very weak on

1st whorl, by end of 2nd whorl as large as P2 and P3,

which are similar throughout, P3 peripheral, summit of

P4 exposed on spire throughout. Secondary spirals de-

veloping to resemble primaries, SI commencing on early

to late 3rd whorl, S2 commencing on 1st half of 2nd

whorl; axial costae commencing at end of 1st whorl,

strong, prosocline, entirely traversing all spire whorls to

P4. Base with 7 or 8 similar, strong, rounded spiral cords,

at first smooth then with small rounded nodules, inter-

spaces about as wide as each spiral or wider. Surface

minutely granulate throughout. Collabral growth lines

prosocline on spire, opisthocyrt on base. Aperture

subquadrate. Radula (figures 106-107) with formula c.6

+ 3 + 1 + 3 + c.6, highly degenerate, all teeth other

than innermost 2 pairs of marginals flexible, almost ge-

latinous. Central and lateral teeth slender, narrowly ta-

pered, extremely thin in section, tips finely serrate In-

nermost marginal shortest, stoutest, cutting area angu-

late. strong terminal cusp and 3 or 4 secondary cusps on

each side. Outer marginals slender, innermost pair lon-

gest, cutting area with long series of stout, narrowly ta-

pered, curved cusps; outer marginals weakly developed.

Type Data: Holotype MNZ M. 59698 (height 5.35 mm,

width 4.95 mm, 4.70 teleoconch whorls): BS 668 (R26),

41°55.9'S, I74°43 2'E, SE of Cape Campbell, alive, 454-

424 m, 14 January 1979, r.v. Tangaroa. Paratypes (4

MNZ): BS 969 (R27), 41°55.8'S, 174°40.7'E, SE of Cape

Campbell, alive, 434-446 m, 14 January 1979, r.v. Tan-

garoa.

Other Material Examined: 1 incomplete shell MNZ,

45°45'S, 171°02'E, off East Otago, 600-520 m, r.v. Miin-

ida.

Distribution (figure 108): Off Cape Campbell and off

East Otago, 424-600 m, taken alive at 424-454 m from

rugged substrata with shells and corals.

Diet: Unknown.

Remarks: Compared with the Australian species Cal-

liostorna retiarium, which it most closely resembles, C.

kopua differs principally in being more strongly nodular,

in being larger relative to the number of whorls, and in

having a less excert protoconch. The single incomplete

shell from off East Otago is 6.55 mm in diameter (esti-

mated height 7 mm), suggesting that the type specimens

are immature. As in other members of this group the

Page 124

THE NAUTILUS, Vol. 108, No. 4

radiila is degenerate through developmental retardation,

C. kopua having the most degenerate radula of any known

cailiostomatid.

Etymology: Deep (Maori).

ACKNOWLEDGMENTS

I am grateful to the following collectors who provided

much of the material: M. Eraser (Wellington), D. Gibbs

(Auckland), G.S. Hardy, CD. Paulin, CD. Roberts, and

A.L. Stewart (Museum of New Zealand, Wellington),

P.R. Jamieson (Wellington), S. O'Shea (Wellington), JR.

Penniket (late of Warkworth), J. Raven (Wellington),

and D. Watson (Otaki). Thanks to A.G. Beu (Institute of

Geological and Nuclear Sciences, Lower Hutt) and J. A.

Grant-Mackie and J. Sutherland (University of Auck-

land) for the loan of fossil material, to A.G. Beu for advice

on stratigraphic problems, and to P. Anderson and W.

Main (National Institute of Water and Atmospheric Re-

search, Wellington), P. Bouchet (Museum National

d'Histoire Naturelle, Paris), W.O. Cernohorsky and B.

Haywood (Auckland Institute and Museum), R.G. Mool-

enbeek (Zoological Museum, Amsterdam), T. Schiotte

(Zoological Museum, Copenhagen), K.M. Way (The Nat-

ural History Museum, London), and Sorting Center staff

of the National Museum of Natural History (Washington

D.C) for the loan of Recent material. Eor comments on

the manuscript I thank A.G. Beu and J.H. McLean (Los

Angeles County Museum of Natural History). For pho-

tography and photographic printing, I am grateful to N.

Heke, M. Hall (Museum of New Zealand, Wellington).

Thanks also to W. St. George (Institute of Geological and

Nuclear Sciences, Lower Hutt) for access to the scanning

electron microscope.

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Bruce A. Marshall, 1995 Page 127

APPENDIX

CHECKLIST OF RECENT CALLIOSTOMATIDAE FROM THE NEW ZEALAND REGION WITH NEW SYNONYMS

CALLIOSTOMATINAE

Calliostoma (Matirea) antipodense Marshall, new species.

C.(M.) aiipourianum Marshall, new species.

C.(M.)bcnlhicola (Dell. 1950).

C.(M.} blacki (Powell, 1950) {= couperi Vella, 1954 = profunda Dell, 1956).

C.(M.) cmiiiens Marshall, new species

C.(M.) fovcauxanum (Dell, 1950).

C.{M.) gibbsonim Marshall, new species.

C.{M.) granti (Powell, 1931) (= ampla Powell, 1939 = multigemmata Powell, 1952).

C.(M.) jamiesoni Marshall, new species.

C.fM.j maui Marshall, new species.

C.(M.} mcgaloprepcs (Tomliii, 1948).

C.{M.) oshornei Powell, 1926.

C.{M.) pclbicidum (Valenciennes, 1846) (= nndulatum Finlay, 1923 = spiratum Oliver, 1926).

C.(M.) pcnnikcti Marshall, new species.

C.(M.) punctulatum (Martyn, 1784).

C.(M.) regale Marshall, new species.

C.(M.) selectum (Dillwyn, 1817) (= hodgci Hutton, 1875 = ponderoms Hutton, 1885 = caniicolor Preston, 1907).

C.(M.) simulans Marshall, 1994

C.(M.) spectabile (A. Adams, 1855).

C.(A4.) tigris (Gmelin, 1791) (chathamensis Dell, 1950).

C.(M.) turjierarum (Powell, 1964).

C.(M.) waikanac Oliver, 1926 (= morioria Powell, 1946 = forsteriana Dell, 1950 = haurakiensis Dell, 1950).

C.(Olukaia) alertac Marshall, new name {blacki Dell, 1956).

C. (sensti tato) kopua Marshall, new species.

C {sensu lata) limatulum Marshall, new species.

C. (sensu lata) onustum Odhner, 1924.

New genus A, new species (Marshall, 1995).

New genus B, new species (Marshall, 1995).

THYSANODONTINAE

Thijsanudonta aucklandica Marshall, 1988.

T. wairua Marshall, 1988.

Carinastelc coronata Marshall, 1988.

C. jugosa Marshall, 1988.

C. krisiellae Marshall, 1988.

Five additional calliostomatines are known from the Kermadec Islands, to the north-east of New Zealand (Marshall, 1979):

Calliostoma new species A = C. (Trislwhotrocluis) sp, cf. simplex of Marshall, 1979, 6gs. C, D, not Schepman, 1908 (Marshall

1995).

C. new species B = C. (T.) sp cf simplex of Marshall, 1979, figs. E, F, not Schepman, 1908 (Marshall, 1995).

C. (Trislichotrochus) gendalli Marshall, 1979,

C. (T.) species cf tosaense (Kuroda & Habe, 1961).

C. (Fautor) consobrinum (Powell, 1958).

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