fo. !? ?. S S

Nil''. . J

THE NAUTILUS

Volume 112, Number 1

July 21, 1998

ISSN 0028-1344

A quarterly devoted

to malacology.

Mari

JUL 3 1 7998

EDITOR-IN-CHIEF

Dr. Jose H. Leal

The Bailev-Matthews Shell Museum

3075 Sanihel-Captiva Road

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EDITOR EMERITUS

Dr. M. G. Harasewych

Department of Invertebrate Zoology

National Museum of

Natural History

Smithsonian Institution

Washington, DC 20560

CONSULTING EDITORS

Dr. Riidiger Bieler

Department of Invertebrates

Field Museum of

Natural Histor\'

Chicago, IL 60605

Dr. Arthur E. Bogan

North Carolina State Museum of

Natural Sciences

Raleigh, NC 27626

Dr. Robert T. Dillon, jr.

Department ol Biology

Coflege of Charleston

Charleston, SC 29424

Dr. William K. Emerson

Department of Living Invertebrates

The American Museum of Natural

Histor\'

New York, NY 10024

Dr. Eileen H. Jokinen

Institute of Water Resources

University of Connecticut

Storrs, CT 06269-4018

Mr. Richard I. Johnson

Department of Mollusks

Museum of Comparative Zoology

Harvard Universit)'

Cambridge, MA 02138

Dr. James H. McLean

Department of Malacology-

Los Angeles Counts' Museum of

Natural History-

900 Exposition Boulevard

Los Angeles, CA 90007

Dr. Arthur S. Merrill

% Department of Mollusks

Museum of Comparative Zoolog)'

Harvard University-

Cambridge, MA 02138

Dr. Paula M. Mikkelsen

Department of Living Invertebrates

The American Museum of Natural

Histor)'

New York, NY 10024

Dr. Gustav Paulay

Marine Laboraton,'

Universit)' of Guam

Mangilao, Guam 96923

Mr. Richard E. Petit

PO. Box 30

North Myrtle Beach, SC 29582

Dr. Edward j. Petucli

Department of Geologv-

Florida Atlantic University'

Boca Raton, FL 33431

Dr. Gary Rosenberg

Department of Mollusks

The Academy of Natural Sciences

1900 Benjamin Franklin Parkway

Philadelphia, PA 19103

Dr. Ruth D. Turner

Department of Mollusks

Museum of Comparative Zoology

HaA'ard University

Cambridge, MA 02138

Dr. Geerat J. Vermeij

Department of Geology

University of California at Davis

Davis, C:A 95616

Dr. G. Thomas Watters

Acjuatic Ecology Laboratory

1314 Kinnear Road

Columbus, OH 43212-1194

Dr. John B. Wise

Houston Museum of Natural Science

Houston, TX 770.30-1799

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THE €7 NAUTILUS

CONTENTS

Volume 112, Number 1

July 21, 1998

ISSN 0028-1344

H. Lee Fairbanks Clarification of the taxonomic status and reproductive

anatomy of Pliiloim/nis hatchi Branson, 1968 (Gastropoda:

Pulmonata; Philonivcidae) 1

Da\id G. Reid Additions and corrections to the taxonomy of the genus

Yiu-Ming Mak PcasicUa Nevill, 1885 (Gastropoda: Littorinidae) 6

Jose H. Leal Donald Richard Moore (February- 16, 1921-October 19,

1997): biographical sketch, zoological taxa, anil bibliography 34

Notices 41

THE NAUTILUS 112{l):l-5, 1998

Page 1

Clarification of the Taxonomic Status and Reproductive

Anatomy of PJiilomycus batchi Branson, 1968

(Gastropoda: Pulmonata: Philomycidae)

H. Lee Fairbanks

Department ot Biology

PennsvKania State University

100 Universitv' Drive

Monaca, PA 15061-2799 USA

[email protected]

ABSTRACT

The mantle pattern and reproducti\'e anatom\' of specimens of

Philotni/ais batchi are compared to specimens of P cnrolhii-

anus- The results demonstrate that P batchi is distinct from P

carolinianus and therefore P. batchi is reestablished as a sep-

arate species. In the onginal description of P batchi the re-

producti\e system described is that of a designated paratvpe

from a locality different from that of the holotvpe. This study

demonstrates that the reproductive anatomy of P. batchi from

the type locality- is not the same as that in the original species

description. The correct reproducti\e anatomv is described, it

includes a penial pouch, located in the wall of the penis. Tliis

pouch is not recorded in any other United States landsnail.

Key Words: Slugs, Philomycidae, Philomtjcus, reproductive

anatomy, taxonomy.

INTRODUCTION

The Philomycidae is a family of terrestrial slugs found

primarily in the eastern half of the United States. The

species in this family are characterized by a mantle that

covers the entire body e.xcept for the tip of the tail and

the anterior portion of the head. The length of an adult

slug yaries from 25 to 100 mm, depending upon the

species. There are three North American genera in this

family, Philomijcus. PalUfcra and Mcgapallifcra. In terms

of tlie reproductiye anatomy, the species of Philoim/cus

are characterized by the presence of a dart sac and dart;

the species of Pallifcra are small {approximately .30 mm

long) and lack the dart sac and dart; the species oi Me<ia-

pallifcra are large (appro?dmately 80 mm long) and also

lack the dart sac and dart but, unhke Pallifcra, haye a

bilobed "flap" on the internal surface of the upper atri-

um. Most of the original species descriptions of these

philomycids tlid not include data describing the internal

anatomy, particularly the reproductiye anatom\-.

The description of Philomijcus batctii Branson, 1968

did include a figure of a reproductixe system and a de-

scription of a portion of the terminal genitalia. Howeyer,

the author utilized the holotvpe for descriptK)n of the

external moqihology only. A ilesignated paratvpe col-

lected at another locahtv (approximately 135 km south

of the type locality) was used to describe the reproduc-

tive anatomy. Later, Hubricht (1974) synonymized, with-

out supporting data, P. batchi with P. carolinianus (Rose,

1802).

Fairbanks (1986, 1989, 1990, 1993) examined sLx of

the eight described species of Philomi/cus and all three

species of Mcgapallifcra. In all of the species examined

there were species specific differences in penis shape

and/or internal penial anatomy. Because inter-species

differences exist, it was decided that the reproductive

anatomy of P batchi and P. carolinianus should be com-

pared. In particular, it seemed appropriate to compare

the reproducti\'e anatomy of .specimens of P batchi from

the t\pe locality with that of specimens from the para-

tvpe locality.

The goals of this study were: (1) to determine whether

Philouu/cus batchi was correctly symonvmized with P.

carolinianus, and (2) to determine whether the repro-

ductive anatomy ascribed to P. batchi is the same as that

of P. batchi from the type locality.

MATERIALS AND METHODS

Specimens ot Philomijcus batchi were collected 1.9 km

south of Highway 627, on a hillside along the road on

the east side of the Kentuck"v River, elevation 210 m,

Clark Coun^, Kentucky, 16 Mav 1990. This site (site 1

in Fig. 4) is tbrectK' across a road from the tvpe locahty

(the east flood plain of the Kentucky' River, opposite

Boonesborough State Park, Madison-Clark Countv Une,

Kentucky ). Other specimens, presumed to be P. batchi,

were collected from an area along Eagle Knob Branch

of Eagle Creek near Cumberland Falls State Park,

McCreary County, Kentucky (the paratype locality), el-

evation 300 m, 6 May 1991 (site 2 in Fig. 4). The ho-

lotype of P batchi (FMNH 155478, Field Museum of

Page 2

THE NAUTILUS. Vol. 112, No. 1

Figure 1. Philointjais batchi from tlie t^pe locality.

Figure 2. Philomi/cus batchi from the paratype locality.

Figure 3. Pluloiiii/cus ciiroliniantis from South Carolina. Scale bars = 10 mm.

Natural History, Chicago) was examined, the paratype

used lor the description of P. hatchi's reproductive anat-

omy has been lost (personal communication, Branley

Branson, 6 October 1993). Specimens of P carolinianus,

from the author's personal collection, collected from logs

alongside a road 5 km north of Huger, Francis Marion

National Forest, Berkeley County, South Carolina, 22

June 1988, were used for comparative puqioses. This

site is 15 km north of the designated (Pilsbry, 1948:754)

type locahtv oi Charleston, Charleston Counts-, South

Carohna.

All specimens were identified b\ comparing external

morphology with species descriptions. The specimens

were drowned in tlistilled water and immediateK' dis-

sected. The reproductixe s\stems were removed and

pinned out in 70% ethanol tor 24 hours prior to pre-

serving them in 70% ethanol. The image of each repro-

ductive system was projected (using an overhead projec-

Figure 4. Outline of the state of Keiituclcv showing the location ol the (\pi-

Philornijnis htttrhi^ Scale bar equals 5.3 km.

localitv (site 1) and the paratope locaht\ (site 2' of

H. L. Fairbanks, 199S

Pa^e 3

tor) onto paper and traced. The atrium and penis of each

specimen were opened and photographed through a dis-

secting microscope. The slides were then projected onto

paper and traced.

Voucher specimens are in the Nation;il Museum of

Natural History, Smithsonian Institution (USNM

860578, three specimens of PhiJomijcus hatchi from

Clark Countv, Kentuck-s\ the tspe locahtv, and USNM

860.579, one specimen of Plulonu/ciis sp. from McCrearv

County, Kentucky, the paratvpe locahty), and in the

Academy of Natural Sciences, Philadelphia, PA (ANSP

A13.329 for P carolinionus).

RESULTS

The reproductive systems of four adult specimens of

Philoim/ciis hatchi collected from the t\-pe locality

(Fig. 4, site 1) are as shown in Figure 5. Thev do not

match the reproductive svstem in the original descrip-

tion of this species. Four specimens matching the de-

scription of the external moqihologv of P. hatchi were

collected from the paratope locality (Fig. 4, site 2). The

reproducti\'e system (Fig. 6) of one of these (Fig. 2)

matched the characteristics and figure given in the orig-

inal species description, and will be referred to as P.

unknowai. The reproductive anatom\' of the other three

specimens was similar to that of the specimens from the

type locahts' and is not figured. The reproductive svstem

of P. carolinianus from South Carofina is shown in Fig.

7. The reproductix'e SN'stem of P hatchi from the t^pe

locaht)' (Fig. 5) had a penis that tapered gradually- from

its proximid (atrial) end to its junction with the vas de-

ferens. In P. imkiiown from the paratype locaht\' (Fig.

6) the penis tapered rapidly for one-fourth of its length,

remiiined relati\'elv constant in diameter for the next half

of its length, and then for the distal one-fourth it rapidlv

decreased in diameter to merge with the vas deferens (a

"two stepped" appearance). The penis of P caroliuiaiiiis

(Fig. 7) was relati\eK- constant in diameter for the first

(proximal) hall of its length, then the diameter decreased

rapidly to match that of the vas deferens (a "one step"

appearance).

The internal penial anatomv of Pliilomijai.s hatchi

from the t\pe localih' is showni in Fig. 9. Figure 8 shows

a transverse section through the peni;il pouch of the pe-

nis of a specimen from the type locality. Fig. 10 shows

the internal penial anatomy of P. unkiiown whose repro-

ductive anatom\' matched that of the original species de-

scription. Figure 11 shows the internal penial anatomy

of P. carohnianits. The internal penial wall of P. batciti

from the type locahty' was smooth for the proximal (atrial

end) one fourth of its length The second quarter was

covered with \-er\- small papillae arranged in longitudinal

rows. The third quarter had a large longitudinal ridge,

it and the remainder of the internal wall was covered

with larger papillae randomly arranged. The distal quar-

ter of the penial wall was covered with narrow folds that

led into the opening of the \as deferens. A second ridge

extends from near the atrial end of the penis two-thirds

8 PS LPP

Figure 5. Reproductixe svstem of Philoiui/cus batchi from

the t\pe locality.

Figure 6. Reproducti\e system of Philomtjcus hatchi from

the paratspe locality.

Figure 7. Reproducti\e s\stem of Philomi/cus cnrolininniis

from South Carolina.

Figure 8. Section through the penis along X — X ni Fig. 5.

Scale bars for Figs. .5, 6 = 10 mm, for Fig. 7, = 5 mm. A,

atrium; AG, albumen gland; DS, dart sac; G, ovotestis; LP,

lumen of penis; LPP, lumen of penial pouch; P, penis; PR,

penial retractor; PS, penial sheath; S, spemiatheca; VD, vas

deferens.

of the way to the vas deferens. The internal surface of

the penis of P unknown (Fig. 10) was covered with very

small randomh' placed papillae (not drawii to preserve

ridge clarity). A large longitudinal ridge begins at the

proximal end of the penis and extends cUstaUv approxi-

mately two-thirds of the penial length. The ridge has a

raised portion at the atrial end The distal one-third of

the internal surface of the penis was covered bv thin

longitudind ridges that led into the opening of the vas

deferens. The internal penial surface of P. carolinianus

(Fig. 11) had three to four large folds on the proximal

third, a second set of three to h)ur small ridges on the

middle third, and the chstal third had narrow ridges that

led into the \as deferens.

Page 4

THE NAUTILUS, Vol. 112, No. 1

Figure 9. Internal penial aiiatomv o( Philomi/rit.s hatchi from

the hpe localitw

Phih

uuijms

batchi

Figure 10. Internal penial anatomy

from the parat\pe localit\'.

Figure 11. Internal penial anatomy of Philomijcus carolini-

anits from South Carolina. Scale bars = 10 mm. A, atrium;

DS, dart sac; OPP, opening into penial pouch; P, penis; PP.

penial pouch; PS, penial sheath; VD, yas deferens.

A small circumferential chamber (tlie penial pouch)

having a sht-hke opening into the lumen ot the penis

(Figs. 9, 10) was present in the penial w;d] of all speci-

mens from Kentucky used in thi.s study. This chamber

extended for nearly tsvo-thirds of the circumference of

the penis, and its outer wall was coyered with \illum-

Uke papillae (Fig. 8). The holotype of P. batchi was im-

mature, its reproductive system was one-half to two-

thirds fully developed. It was possible to detennine the

shape of the penis, it was long, narrow with a slight taper

from the proximal to the distal end. Two characteristics

of the internal penial anatomy were discernible: a peni;il

pouch, which was located appro.ximatelv one-third of the

length of the penis from tlie proximal (atriali end, and

a single large ridge on the internal wall oi the proximal

third of the penis.

The correct description of the reproductive anatomy

of Philomyctis batchi, based on material from the t\pe

locality is as follows:

Genitalia of Philomijcus batchi Branson, 1968 (Figs.

5, 8, 9)

Atrium approximately 45% penial length, outer sur-

face glandular, light orange to light pink in color, with

finely ridged pad on the inner surface near the opening

into the vagina. Vagina approximately 10% penial length.

Origin of spermathecal duct approximately 1.7 times the

diameter ol the oviduct, slight taper to spermatheca.

Spermatheca ovoid. Dart sac approximately twice sper-

mathecal diameter Proximal penis diameter approxi-

mately e(jual to atrial diameter, penis tapers to diameter

of vas deferens; penial sheath appro.ximately 70% of pe-

nis length. Internal penial surface smooth for proximal

25% of length, tinv papillae in longitudinal lines for next

25%, large papillae for ne.xt 25% with one large ridge,

thin folds leading into vas deferens for distal 25% of

penial length. A second ridge extends from the atrial end

of penis two-thirds the distance to the vas deferens; a

circumferential pouch in the peniid wall with opening

into lumen oi penis, pouch located in second 25% of

penial length, pouch extent approximately 60% of cir-

cumference of penis, pouch with villum-Uke papillae

covering outer wall. Vas deferens length approximately

3 times penial length, enters distal end of penis, diam-

eter of vas deferens at distal end 2.5 times that of prox-

imal end. Oviduct length approximately 30% penial

length. Penial retractor muscle length approximately

35% penial length, inserted at junction of penis and vas

deferens. Accessory retractor muscle robust, located on

atrial surface near junction of penis and atrium. Uterus,

;dbunien gland, spermoviduct, and gonad tvpical of ge-

nus.

DISCUSSION

Hubricht (1974) svnonvanized Philoim/cus batchi with P.

caroUuiauiis because he saw P. batchi as a ". . . melanistic

color form of P. caralinianiis. " However, the usual cri-

teria used to identify P- carolinianus are the presence of

a double row of black spots along the dorsal surface of

the mantle with a brown stripe between them (Fig. 3).

Philoiiujciis batchi does not have these characteristics

(Fig. 1). Furthermore, the internal penial anatomy also

is different (Figs. 9, 11). Phihiim/ciis batchi has two large

ridges on the internal surface ot the penis and a penial

pouch (Figs, 8, 9), whereas P. carohiiianiis has from .six

to eight much snuJler ridges and lacks a pouch (Fig. 11).

There are differences in the atrial anatomy as well. Phi-

Joiui/cits batchi has a large pad, with several small folds

in it, in the upper atrium near the opening into the va-

gina (Fig. 9). Philoinifciis carohnianus has a few small

folds that lead into the opening of the vagina, but no

pad (Fig. 11). These data demonstrate that Pliilomt/cits

batchi is distinct from P. carohnianus and therefore must

be reestablished as a species.

The external appearance of Phihmtjcus batchi from

the tvpe locality was indistinguishable from that of all

specimens from the paratspe localitv. Howe\'er, specific

features of the reproductixe system of P. batchi from the

type locahtv (Fig. 5) were clearK' different from those

of the original description, as well as from P. unknown

from the paratype locality (Fig. 6), and did match the

original description (see Table 1 for a brief comparison

of the reproductive anatomies). Two additional tlifier-

ences betyveen the specimens oi Phihimi/ciis batchi from

the type locality and P. unkiiown from the paratvpe lo-

cality, were the position and size of the penial pouch

(Figs. 9, 10). That is, the penial pouch is larger in P.

batchi from the type locahty than m P. unknown from

the paratype localiK', and the latter specimen's peni;d

pouch is more pro>dmal than that of the specimens from

the type loc;ility'.

The data obtained from the immature holotype were

consistent with the Philomijcus batchi specimens col-

lected from the type locality. That is, the general shape

of the penis was similar, and the location of the penial

pouch and the presence of the ridge on the internal sur-

face near the pro.\im;d end of the penis were the same.

When Branson (1968) described Phihiimjcus batchi, he

apparently assumed, based upon external appearance,

that slug specimens he had collected in McCreary Coun-

ty, Kentuck-y- were of the same species as the one a col-

league had collected in Clark Ct)untv, Kentuck-y 135 km

to the north. He then described a new species using hvo

specimens. Based upon the data presented in this studv,

it appears that Branson utihzed two different species to

describe P. botchi. Therefore, a description of the cor-

rect reproductive anatomy for P. batchi is provided. The

specimen that Branson used to describe the reproduc-

tive system of Philoimictis batchi appears to be an un-

described species. Because only one specimen of this

species is presentK a\;iilable. its description will appear

in a later paper.

An unexpected result of this stud\' was the disco\ery

of a chamber in the penial wall of the specimens of Phi-

lonu/ciis from Kentucky. The gross structure and posi-

tion of the chamber is not recorded from any other

group of land snails in the United States. Studies are

underyvav to ascertain the number of different species

of Philomijcus that have this chamber, and to try to de-

termine hov\' widely spread these species are geograph-

ically The function or functions of this chamber are un-

knowii at this time.

ACKNOWLEDGMENTS

Financial support for the fiekl trips associated with this

study was provided bv grants from the Research Devel-

opment Grant Fund of Penns\lyania State University.

LITERATURE CITED

Bosc, L.A.G. IS02. Histoire naturelle des coquilles, contenant

leur description, et leurs moeurs. Vol. 1. Paris, 34.3 pp.

Branson, B. 1968. Two new slugs (Pulmonata: Philomvcidae:

Philomijcus) from KentuckT and N'irginia. The Nautilus

8I(4):I27-I.33.

Fairbanks. H. L. 1986. The taxonomic status of Phihvni/ciis

togatus (Pulmonata: Philomvcidae): a morphological and

electrophoretic comparison with Phih^mijcu.'i cnrohnianus.

Malacolopa 27(2):271-2S0.

Fairbanks, H. L. 1989. The reproducti\e anatomy and taxo-

nomic status of Philonujam ventisttis Hubricht, 1953 and

Fhilomi/ciis bisdosiis Branson, 1968 (Pulmonata: Philo-

mycidae). The Nautilus 103(l):20-23.

Fairbanks, H. L. 1990 Morphological comparisons of the spe-

cies of Mi'gapnllifcra (Gastropoda: Philomvcidae). The

Nautilus 104(2):71-75.

Fairbanks, H. L. 1993. The reproductive anatomy of Philo-

im/cus scUatus Hubricht, 1972 and Philomijcus virainiais

Hubricht. 1953 (Gastropoda: Philomycidae). The Nautilus

107(1):9-13.

Hubricht, L. 1974. A re\iew of some land snails of the eastern

United States. Malacological Re\iew 7:3.3-.34.

Pilsbry, H.A, 1948. Land Mollusca of North America (North

of Mexico). Vol. 11, Part 2:754.

THE NAUTILUS 112(l):6-33, 1998

Page 6

Additions and Corrections to the Taxonomy of the genus

Peasiella Nevill, 1885 (Gastropoda: Littorinidae)

Da\id G. Reid

Department of ZooIogN

The Natural Histon Museum

Lxmdon S\V7 5BD,' UK

D.reidCS'nhm.ac.nk

Yiu-Ming Mak'

Department of Ecology and

Biotliversity

The University of Hong Kong

Hong Kong

ABSTRACT

Peasiella is a genus of the Littonmdae with nnnute, trochoidal

shells, found onlv in the Indo-West Pacific region. A prexious

monograph recognized slx Recent species. As a result of new

anatomical information (reproductive anatomy, egg capsules,

radulae) two of these are dixided into five and two species,

respectively, to make a total of 11 species in the genus. Three

of these are descnhed as new. Additional notes on distribution,

variation and nomenclature are provided on otlier species.

Characters of the radula and pallial oxiduct are relatively uni-

form throughout the genus, hut penial shapes are often diag-

nostic. Shell shape and sculpture show extreme mtraspecific

x-ariabilitx'

INTRODUCTION

Pca.sicllo is one of the more poorly knoxvii genera of the

Littorinidae. In common xxith all members of the ,sub-

family Littorininae, its species occur t)n hard substrates

in the httoral zone. Although easily accessible and often

abundant, they are infre(juently collected oxving to their

small size and crvptic habit in rock crexaces and among

oyster and barnacle shells. All knoxvn species occur in

the Indo-West Pacific province and most are entirely

tropical, although several extend into the temperate

zones of southern Africa and the northxx'esteni Pacific.

Among Utt(jrinids, they are readily recognized by their

small (1-7 mm diameter), trochoidal or depressed shells,

lacking spines, with open umbihcus and multispiral oper-

culmn. These characters are superficially similar to those

of trochids (but lacking the nacreous interior of that

group), with which they were often contused by eark

authors (see review of taxonoinic histon- bv Reid,

1989a). Other littorinids with trochoidal sliells include

the genera Bciiihicimu and m.scllop.sis irom Australia

and New Zealand (Reid, 1988), and Tcctiiniis and C.i'u-

chritis from the Indo-West Pacific and western Atlantic

(Ro.sevvater, 1972; Reid, 1989b; Reid & Celler, 1997).

' Present address: 1.3/F, Canton Road Covemment Offices, Ag-

riculture and Fisheries I^epartinent, Hong Kong

For long these small littorinids were neglected ta.\o-

nomicallv. The most recent monograph of e.xtant and

some fossil Pca.sicllo species was that by Reid (1989a);

before this the onlv compilations were a fist by Nevill

(1885) and a monograph bv Trvon (1887). The anatomy

(male and female reproductive tracts, paraspermatozoa,

egg capsule, alimentary tract, head-foot pigmentation),

radula, protoconch and teleoconch were described in de-

tail bx' Reitl (1989a). As a result, the genus xxas defined

not onlv bv shell and opercular characters, but iilso by

the uni(jue combination of the elongate penis with single

mamilliform peni;il gland and closed vas deferens, to-

gether xvith the double-looped form of the pidfial ovi-

duct, and other anatomical features. These same char-

acters were included in a phvlogenetic an;ilysis of the

morphology of the genera and subgenera of the Litto-

rinidae (Reid, 1989b). However, since none of the coded

anatomical characters appeared to be apomoqihic xxithin

the famil), the (juK' formal svnapomorphies of the genus

in this analysis were the trochoidal shell shape and mul-

tispiral operculimi, and neither of these xx'as unitjue.

Nexertheless, the close moiphological similarities among

its members leaves little doubt that the genus is a mono-

phvletic one; the arrangement of the loops of the p;dli;d

oxiduct is one unitjue character, but oxving to difficulties

of coding such structures it did not appear as such in

the formal auiilvsis. As a result of Reid's (1989b) phylo-

genetic analysis, it was clear that Peasiella was a member

of the derived subfamilv Littorininae, and probably a

relatively basal member, although its precise relation-

ships xxith such genera as Maimvarin^ia, Ccnchhtis and

Tcctariiis were not resolved. A recent molecular study

included a species of Peasiella. vxith members of Tcctar-

iiis. Ccncliriti.\ and Sodiliftorina. but did not find any

close relationships xxith these other genera (Reiil & C;ell-

er, 1997).

In th(- previous systematic account ni Peasiella species

(Reid, 1989al, anatomical details were provided for all

the six species then recognized. Hovxexer, the amount

of material available vxas limited; for example, for P.

'isseli' onlx one preseived male was dissected, and tor /'

'iiifraciKtalii' oulx seven. As a result, it was loiuluded

D. G. Reid and Y.-M. Mak, 1998

Page 7

that the genus was anatoinicallv rather uniform, without

the striking differentiation in reproductive characters

that has proved so useful for the discrimination of spe-

cies complexes elsewhere in the faniilv (e.g. Reid, 1986a,

1988, 1996). Instead, it was necessar\' to base species

definitions largely on the shape, sculpture and color of

the shell (although these characters were evidently sari-

able), and on geographical distributions. Anatomical

characters such as head-toot pigmentation, the number

of cusps on the outer margiuiil radular tooth, and pres-

ence or absence ot a copulatory bursa provided only mi-

nor input.

Our attention was once more drawn to the svstematics

of PcasicUa as a result of a stud\- iiv one of us (Y-MM)

of the littorinids ot Hong Kong. The fauna of Hong Kong

was thought to include three TcasicUa species (Reid,

1992), the highest number then known to occur s\in-

patricallv. However, it was tound that two distinct shell

forms of P 'infracostata' scnsti Reid (1989a), both rare

in Hong Kong and not yet found svnitopically (i.e. on the

same shores), produced egg capsules of different shape.

This prompted a reexamination ot this stipposed species

throughout its wide range in the IncUan and Pacific

Oceans. Meanwhile, further collections were made by Y-

MM throughout the Far East. During the past decade

much new material has also accumulated in the major

museums. Using this new material, we found that penial

torm was, after all, a character showing useful interspe-

cific variation. As a result, we have revised the taxonomy

of two species complexes; PcasicUa 'infracostata' scnsii

Reid (1989a) is here showai to consist of five species,

and P. 'isscli' scnsti Reid (1989a) of two. These seven

species are fully described. We also record additional

information on the variability, distribution and nomen-

clature of other PcasicUa species. The number of species

recognized in the genus is therefore increased from six

to eleven.

MATERIALS AND METHODS

The descriptions and records are based on material in

the following museums: Natural History Museum, Lon-

don (BMNH); Museum National d'Histoire Naturelle,

Paris (MNHN); Instituut voor Svstematiek en Populatie-

biologie, Amsterdam (ZMA); Nationaal Natuurhistorisch

Museum, Leiden (NNML); Institut Royal des Sciences

Naturelles de Belgique (IRSNB); National Museum of

Natural History, Smithsonian Institution, Washington,

DC (USNM); Academ%- of Natural Sciences of Philadel-

phia (ANSP); Museum of Comparatixe ZoologN, Harvard

Universit}- (MCZ); Los Angeles Countx' Museum of Nat-

ural History (LACM); Natal Museum (NM); National

Science Museum, Tok-yo (NSMT): Department of Earth

and Planetary' Science. Kyushu Universit)- (KU); Austra-

lian Museum, Sydney (AMS) and the private collections

of H. Dekker, The Netherlands (HD) and J. Le Renard,

Paris (LR). In addition, t\pe specimens have been ex-

amined from Oxford Uni\ersit\ Museum (OUM); Mu-

seo Civico di Storia Naturale Giaconio Doria', Genoa

(MGD) and Zoological Survey of India. Calcutta (ZSI).

Unless otherwise indicated, all figured specimens are in

BMNH. LectoKpes ha\e been designated from svmtypic

series onK' for \-alid species names.

Shell height (H) was measured parallel to the axis of

coiling, and maximum shell diameter (D) perpendicular

to this iLxis. The height/diameter ratio was used as a sim-

ple index of shell shape. The number of whorls of the

protoconch was counted as described by Reid (1996: 9).

Where counts of the number of ribs above the periphery

are given, this includes the strong rib at the periphery;

this rib is not included in counts of the basal ribs. Li\ing

material was preserved in 80% ethanol, and dissected

under a binocular microscope. PiJUal oxiducts were ex-

amined by dissection; in \ery small animals the copula-

tory bursa can be impossible to find unless filled with

recentK deposited sperm; to confirm absence of the bur-

sa woukl re(juire histological preparation, but this was

not done in the present study. The loops of the albumen

and capsule gland follow a similar path in all species

examined, and are therefore not re-described (see Reid,

1989a, tor description and figures). Radulae were

cleaned ot tissue in a cold, ililute solution of hypochlorite

bleach, thoroughly rinsed in distilled water, and mount-

ed on a thin laver of polyvinyl acetate glue on glass cox'er

slips. Ratlulae were examined with a scanning electron

microscope. Unworn radular sections were photo-

graphed from three orientations: vertically above flat

radula (to show shape of teeth), at an angle of 45° from

front end of radula (to show shape of tooth cusps), and

at an angle of 45° from the side of the radula (to show

relief). For three species, egg capsules were obtained

from living animals. Females were placed inchvidually in

plastic vials (20 X 47 mm) half filled with filtered sea-

water Vials were covered and kept at room temperature

(24-26° C) with no aeration. The water was renewed

daily, and eggs were collected from the bottom of the

containers. Eggs were preserved in 5% formalin in sea-

water, for subsequent examination and measurement us-

ing light microscop\-.

The fists ot material examined are those records used

tor the cUstribution maps. Numbers of specimens dis-

sected are incUcated by M (male), F (female); R indicates

radula preparation, and P protoconch examination. For

the seven .species described in detiiil, the material listed

mcludes that examined during the previous study by

Reid (1989a), and the descriptions supercede the earlier

account. This earfier work should still be consulted for

detailed descriptions of the histology of the reproductive

system and of the sperm For those species for which

only additional notes antl records are given, full descrip-

tions and other records can be found in Reid (1989a).

SYSTEMATIC DESCRIPTIONS OF REDEFINED

AND NEW SPECIES OF PEASIELLA

PcasicUa infracostata (Issel, 1869)

(Figures 1-5, 35-41, 76-79, 108-109, 131)

Trochus sp, Audouin, 1826:42

Risella infracostata Issel, 1869:19.5-196, .348 [refers to Savigiiy,

Page 8

THE NAUTILUS, Vol. 112, No. 1

1817: pi. 5, fig. 40.1, 40.2; 16^01)^6 (Reid, 1989a; fig.41)

+ .3 paraleetot)pes MNHN, seen; Suez (Issel)]. Pallary,

1926:86, pi. 5, fig. 40.1, 40.2. Bouchet & Danrigal, 1982:

13, fig. 64.

Risella (Peasiella) infrocoslnta. — Ne\ill, 188.5:160. Tryon, 1887:

264, pi. 50. ficrs. 41, 42.

Peasiclhi infracostata— -Reid. 1989a:61-64, figs. 106-109,112

[in part; includes P fasciata. P fuscopiperata. P. habei, P

patula]. Bosch et al. 1995:46, fig. 118.

? Risella (Peasiella) tantillus var. subinfracostata Nevill, 1885:

160 [lectonpe (Reid, 1989a; fig. 35) + 2 syntypes ZSI;

lectotype seen; Nicobar Islands].

Nomenclature: In his description of this species, Issel

(1869) referred to Savigny's (1817) unlocalized figures of

Egyptian material, and ;ilso mentioned three specimens

of his own from Suez; the latter were not found by

Bouchet & Danrigal (1982) in Issels collection in MGD.

Reid (1989a) therefore designated one of four shells in

the Savignv Collection in MNHN as the lectotype.

The identit\- oi Risella (Peasiella) tantillus var. subin-

fracostata Nevill, 1885, is uncertain. The lectot\pe (Fig-

ure 35) measures 2. .3 mm in diameter by 2.2 mm in

height (H/D 0.96), is almost smooth above the periph-

ery, but for fine spiral microsculpture and traces of about

four spiral grooves near the periphei;)'; the base bears

four subequal ribs around a small umbihcus; the color is

cream with a single brown spiral line on the first whorl

of the teleoconch, and no other pattern. Of the known

species of the genus, this shell can be compared with P.

infracostata and P. fasciata. In outline it resembles P.

infracostata, for the whorls and aperture are less round-

ed than in P fasciata; however, it lacks the enlarged mid-

basal rib that is often present in the former. The sparse

color pattern is unlike any k-nown specimens of either of

these .species. The locafity of Nevills shells, from the

Nicobar Islands, is far from the clo.sest known occur-

rence of P. infracostata in Bombay, and from that of P.

fasciata in Borneo. The only other Peasiella species re-

corded from the Andaman Sea is P. roepstorffiana (Reid,

1989a), but that .species usually has stronger spiral sculp-

ture, flatter whorls and a strong peripheral keel. Until

more information is a\iiilable, the name is tentatively

included in the synonvmy of P. infracostata.

Reid (1989a) confu.sed five species under the name P

infracostata (here distinguished as P infracostata s.s.. P.

fasciata, P. fuscopiperata. P. Iiahei and P. patula), misled

by their similar and confusingly variable shells.

Shell (Figures 1-5, 35-41): Adult size range 1.1-2.6

mm diameter. Shape: thickness varies from dehcate and

translucent, to thick-walled and opaque; equilaterally

conical to depressed-globular (H/D 0.7-1.0); outline

domed; whorls usuallv rounded or with rounded shoul-

der, becoming almost flat-sided on last whorl of largest,

thick-walled shells; suture always distinct; periphery oc-

casionally uniformly rounded, but usually marked by

sharp angle or rib, rarelv de\eloped as a narrow flange;

base usually shghtly rounded, becoming flat or even

shghtlv concave in largest, thick-walled shells; umbilicus

small to large, occasionally closed in thick-widled shells;

columella narrow, uniformly rounded. Sculpture: proto-

conch not seen, apex usuallv eroded; teleoconch whorls

always lacking ribs above periphery, microscopically

smooth or with fine spiral microstriae (rarely, about 6

striae mav be more prominent, but not developed into

ribs. Figure 39); 1—4 ribs on base (rarely absent, or 5-

6), of which that at mid-point of radius is usually most

prominent (producing shght angulation of base). Color:

translucent yellow-browni, opaque white where shell is

thicker at suture and periphery (entirely white in thick-

walled shells); pattern of p;ile or dark brown or black,

oblique or zigzag, narrow lines from suture to periphery,

numbering .5-19, darkest at periphery and sometimes

idso at suture; pattern occasionally represented only by

peripheral stripes; spire mav be brown, or frequently

marked only by single spiral brown line; base usu;dly

pale with brown line in outermost groove, line some-

times absent, base rarely brownish with paler ribs; col-

umella and parietal area usually not colored, rarely

browni.

Animal: Head-foot: front of head blackish, with a sep-

arate black band across base of snout; tentacles unpig-

mented; foot may have black pigment around opercu-

lum. Penis (Figures 76-79): filament long (50-60% total

length), tapering; base slightly thicker, with single small

mamilliform penial gland. Pallial oviduct: bursa not seen.

Radula (Figt/rc.v 108, 109): cusps pointed, central cusp

of rachichan tooth slightly apiculate; outer marginal tooth

with 6-7 long, pointed cusps, outermost more rounded,

becoming smaller toward micUine.

Distribution: Habitat: Among barnacles in upper eu-

littoral on surf beach (Oman; J. D. Taylor); rock pool on

hmestt)ne, in upper eulittonil of sheltered shore (Oman;

E. Glover); under rocks and stones in intertickil (Oman;

R. G. Moolenbeek & H. Dekker); under coral blocks in

eulittoral (Eg\pt; D. G. Reid). Abundant at some local-

ities in Oman and at Karachi, but elsewhere appears to

be scarce.

Figures 1-5. Peasiella infracostata. 1. Has ;il-Ha<kl, Oman (BMNH 19963S1 ). 2, 4. Kaniclii, Pakistan (BMNH 1996383). 3. Tiwi

Beach, .30 km S Mombasa, Kenya (BMNH 1996.384). 5. 4 km S Ras Qudntali, M;isiiuh I., Oman (BMNH 1996385). 6-10. Peasiella

fuscopiperata. 6. LectoUpe oi Cijelostretna fuscopiperata Turton, 1932, Port AltVed, South Africa (OUM). 7. North Bay, Bengnera

I., Mozambique (NM k7412). 8-10. Tiwi Beach, .30 km S Momb;isa, Kenya (BMNH 1996386). 11-13. Peasiella patula. 11.

Holotype, Chaugi l^)inf, Singapore (BMNH 1996276). 12, 13. St. John's I., Singapore (BMNH 1996387). 14-17. Peasiella fasciata.

14. livol Mission, C;ape Dampier, New Britain. Papua New Guinea (AMS C16312S). 15. HolotApe, Turtle Bay, Cape Ferguson,

Queensland, Austraha (BMNH 1996272) 16, 17. Cape d'Aguilar, Hong Kons; (BMNH 1996.388) Scale bar = 2 nun.

D. G. Reid and Y.-M. Mak, 1998

Page 9

Page 10

THE NAUTILUS, Vol. 112, No. 1

Range (Figure 131): East Africa, Reel Sea, Oman, Pa-

kistan, possibly Nicobar Islands.

Records and material examined: Mozambique; Con-

duda Bav (NM). Tanzania: Zanzibar (BMNH). Kenya:

Tiwi Beach, 30 km S Mombasa (BMNH, 2M). Somalia:

Mogadiscio (ANSP). Egvpt: Hurghada (BMNH; HD);

Suez (Issel, 1869). Yemen; Perim Island (MNHN); Aden

(MNHN): Haui al-M;ihr;ih (HD). Oman: Ravsut, Sal-

aliih (BMNH; ZMA); Kuria Muria Islands (ZMA); Ma-

sirah Island (BMNH, 2M, IF; ZMA); Ras al-Hadd

(BMNH, IM, IF, IR). Paki.stan; 7 km W Bulegi Point,

Sind (LACM); Karachi (BMNH, 3M, 5F 2R). India:

Mada Island, Salsette Island, Bombay (ANSP); Nicobar

Islands (ZSI; uncertain identification, see Nomenclature,

above).

Remarks: The shell of this species shows considerable

\ariation in shape, thick-ness, opacit\' and coloration.

Thin-walled, translucent shells with inchstinct pattern,

rounded whorls and no peripher;il rib (Figures 2, 36, 40)

look very different from larger, thick-wiilled, opaque,

strongly domed shells with striking black and white pat-

tern, shaqDlv angled periphery and flat or concave base

(Figures 1, 3). Yet these extremes are connected by

many intermediates, and even within samples from a sin-

gle localit)' there can be great variation (Figures 36, 37,

39, 40). Whether there is a geographical or ecological

correlation to this variation is not clear Shells from Pa-

kistan are mostly of the translucent form with pale pat-

tern, those from East Africa are opaque and strongly

marked, whereas those from the Red Sea and Oman

span the entire range. A common feature of the shells

is the generallv small number of basal ribs, often 3 or

less, of which that at the mid-point of the radius is usu-

ally slightly larger than the rest. The color pattern of

narrow olilique or wavy a,\ial lines is also a useful rec-

ognition character

This species is closely similar in shell characters to P.

fiiscopiperata, and some of the translucent, rounded

shells are indistinguishable. That thev are indeed distinct

species is confirmed by the difference of the penis in

each (long filament and small mamilfiform gland in P.

infracostata, short filament and large gland in P. fiisco-

piperata), which is maintained in svntopic samples from

Tiwi Beach, Kenya (Figures 76, 77, 80, 81). At this lo-

cality (as in a dry collection from Conducia Bav, Mo(jam-

bique) both species are represented by thick-walled

forms; the shells of P. infracostata have a taller spire,

more steeply domed, with flatter final whorl and sharper

peripheral angle, the base is flatter vvqth 1-2 or rareK' 3

coarser ribs (3—4 in P. fuscopipcrata) and the umbilicus

smaller or closed; the color pattern is sometimes very

similar, but in P. infracostata the oblique lines are dark

and more regular (Figure 3), whereas in P. fuscopiperata

the coloration is much more variable in intensitv and

pattern, including irregular markings, spiral bands and

fine marbhng (Figures 8-10). Thinner-shelled forms of

these two species are distinguished by the slightly taller

spire of P. infracostata, and sometimes by the basal ribs,

of which the centr;il one is often enlarged in P. infra-

costata, whereas in P. fuscopiperata the ribs become

larger toward the periphery.

As a consequence of this similarity, the southern geo-

graphical range of P infracostata is uncertain. Available

material from southern Mozambique and South Africa

consists mainly of dry sheUs, oi the translucent tvpe. AU

anatomical material seen from this area has been of P.

fuscopiperata, but more is required to confirm that P.

infracostata does not occur No anatomical material has

been seen from the Red Sea (including the tvpe locality,

Suez), and the association of the name with the speci-

mens from Oman and Pakistan is based on apparently

identical shells and geographical proximity. The eastern

limit of P. infracostata remains imclear: adtlitional ma-

terial is necessary to confirm the synonviny of NeviU's

(1885) subinfracostata from the Nicobar Islands (see

Nomenclature, above).

Peasiclla fuscopiperata (Turton, 1932)

(Figures 6-10, 42-48, 80-83, 110, 111, 132)

Cijclostrcnm fiiscopifwrata Turton, 19.32:198-199, pi .51, fig.

1.379 [lectot\pe (here designated, 1.9 mm diani.. Figure

6) -I- paralectotype, OUM, seen; Port Alfred. South .\fri-

ca].

Peasiclla infracostata. — Held, 1989a:61-64 [in part; includes P.

infracostata. P. fasciata, P. habei, P. patula; not P. infra-

costata (Issel, 1869)].

Nomenclature: Suiprisingly, since this species is not

unconnnon in southern Africa, it does not appear to have

been mentioned in works on moUusks of the region, ex-

cept in Turton's (1932) origin;d description of shefls from

the southern extremit^• of its range.

Shell (Figures 6-10, 42-48): Adult size range 1.3-

2.7 mm diameter Sluipe: thickness \aries from delicate

and translucent, to more sofid and opacjue; depressed to

flattened (H/D 0.54-0.91); outline domed; whorls

rounded, suture impressed; in thin-walled shells periph-

Figurcs 18-24. Peasiclla halm. 18, 19. 4-5 km SW Tsutsu, Tsushnna I., Nagasaki l^rel'., Japan (BMNH 1996.389), 20. IloloUpe,

Esu Cape, Shiraliama, Wakayama Prcf , Jajian (BMNH 1996274). 21. Tai Ping Bav, Qingdao, China (BMNH 1996.390) 22. Lok

Wo Sha, Mong Kong (BMNIl 1996391). 23. Baten, Okinawa, Japan (BMNH 1996.392). 24. Ma Lin Shi, Tolo ILidioun Hong Kong

(BMNH 1996393). 2.5-29. Peasiella isseli. 25, 28. Oaliab, Sinai, Egypt (BMNH 1996394). 26, 27. Hurghada, E,g\pt (BMNH

1996.395). 29. Suez, Eg>pt (BMNH 1882.8.7.316). .3()-.34. Pca.siclla m/iiiritiana .'50. Ll■c•to^pe o( Ri.sclla isseli var maiiritiana

Viader, 1951, Grand Bav, Mamitius (BMNH 19S9()()4i 31. Arabian C;ult (BMNH 1996.396). .32. Inhaca I., Movamhiiiue (NM

L3435). 33. Sadh, Oman (BMNH 1996397) 34. •/an/il)ar, Tanzania (BMNH 1904.10.20.97). Scale bar = 2 unn.

D. G. Reid and Y,-M. Mak, 1998

Page 11

Page 12

THE NAUTILUS, Vol. 112, No. 1

ery uniformly rounded or marked by slightly thickened

rib onlw in thicker shells peripher\' angled and marked

bv sharp keel or narrow flange; base rounded; umbilicus

of moderate size; columella rounded, shghtK thickened

and angled at base. Sculpture: protoconch 240 |xm di-

ameter, 2.5 whorls, protoconch II sculptured by scat-

tered granules and narrow raised ridges (as in Figures

106, 107 of P. fasciata): teleoconch whorls usually mi-

croscopically smooth above periphery, or with fine spiral

microstriae, rarely with 2-6 slight or indistinct ribs; 2-6

(rarely 0, 1 or 8) ribs on base, outermost strongest, be-

coming weaker toward umbilicus; basal ribs fine, indis-

tinct or absent in thin-walled shells; rarely minute per-

iostracal bristles present on basal and dorsal ribs. Color:

thin-walled shells translucent pale yellow-brown with

faint pattern of 6-14 oblique or waw bro\\ni lines (some-

times present onlv as faint marbling), distinct only at

suture and periphery, base unmarked; thicker shells

opaque white or beige, with highly variable pattern: red-

brown to black coarse, irregular pattern of bands, chev-

rons and zigzags, usually leaxdng an unpigmented sutural

band; sometimes a continuous broad spiral band on

shoulder; sometimes a paler brown pattern of diffuse

marbhng, with or without 9-12 darker peripheral spots;

spire orange-brown, often with dark red-brown spiral

band; base sometimes red-brown with 6-10 dark spots

on white peripheral keel; lower columella and parietal

callus sometimes red-brown.

Animal: Head-foot: front of head blackish, with a sep-

arate black band across base of snout; tentacles unpig-

mented; foot with some black pigment on sides or un-

pigmented. Penis (Figures 80-83): filament small (10-

20% total length), tapering; base thicker, sometimes

wrinkled, with single very large mamilliform penial gland

occupying most of its length. Pallial oviduct: bursa not

seen. Radula (Figures 110. Ill): cusps pointed, major

cusp of lateral and inner marginal tooth sometimes

sUghtly rounded, central cusp of rachidian tooth sUghtly

apiculate; outer marginal tooth with 5-8 long, pointed

cusps, outermost more rounded, becoming smaller to-

ward midline; base of rachidian tooth sometimes devel-

oping a pair of small posterior denticles.

Di.stribution: Habitat: Eufittoral; low, undercut lime-

stone cliffs with fringing reef platform beyond (Kenya;

D.S. Brown): alive in coralline algal debris washed up

on strand line (South Africa; D. Herbert).

Range (Figure 132): East coast of South Africa to Ke-

nya.

Records and material examined: South Africa: Port Al-

fred (NM, OUM); East London (NM); Mbot\i. Pondo-

land (NM); Port Edward (NM); Umdhloti, Natal (NM);

Mapelane, Zululand (NM, 2M); Leven Point, Zululand

(NM, IM). Mozambique: Benguera Island, Bazaruto Ar-

cliipelago (NM); Conducia Bay (NM). Kenya: Tiwi

Beach, 30 km S Mombasa (BMNH, 7M, 4F, 4R, IP).

Remarks: Reid (1989a) identified smooth-shelled

forms of this species as P. infracostata, and those rare

examples with spiral sculpture above the periphery as P.

isseli (with which P. mauritiana was then included). The

form of the penis pro\dded the first evidence that this

species is distinct; the short filament is found elsewhere

in the genus only in P. patula, but there the mamilhform

gland is of a different shape. The present species is svm-

patric with tvvo others, P. mauritiana and P. infracostata,

in East Africa, and penial shape is diagnostic of each.

The shell is most similar to that of P. infracostata, and

separation of the two has been discussed in the Remarks

on that species. In both, the shell shows parallel variation

in thickness, opacity, spire height, angulation of the mar-

gin and intensitv^ of color pattern. Occasional shells with

spiral sculpture abo\e the periphery' could be confused

with those of P. mauritiana; P. fuscopiperata lacks the

shoulder angulation of that species, is usually a more

dehcate shell and rarely shows the same pattern of

strong oblique radial lines.

PeasicUa patula new species

(Figures 11-13, 52-56, 84-87, 112, 113, 123, 127, 128,

133)

Peasiella infracostata. — Reid, 1989: 61-64, fig. 110 [in part; in-

cludes P. infracostata, P. fuscopiperata. P hahei. P. fa.scia-

ta: not P infracostata (Issel, 1869)].

Etymology: Latin patulus, open, in reference to the

relatively large aperture.

Types: Holotype BMNH 1996276 (Figure 11); 5 para-

types BMNH 1996277, 49 parat\pes in alcohol BMNH

1996278 (Figures 87, 112, 113)'. Type locality: Changi

Point, Singapore.

Shell (Figures 11-13, 52-56): Adult size range 1.2-

2.0 mm diameter Shape: depressed to coniciil (H/D

0.75-0.85); somewhat patulous, last whorl enlarged, ap-

erture relatively large; protoconch (if present) projects

as papillose tip to spire; whorls well rounded or inflated,

often with rounded shoulder, suture impressed; angled

peripheiT marked by shaq3 but bareh' projecting keel;

base slightly rounded; umbihcus wide; columella narrow,

cuned at base; viewed from above or below, apertural

Figures 35—41. Peasiella itifracostala. .'$5. Lcctoh pi* u{ Risellu [Peasiella) lunlillus vm. subinfracostata Ne\ill, 1885, Nicobar Is

(ZSI). 36, 37, 39, 40. Karachi. Pakistan (liMNH l'996383). 38. 4 km S Ras Qudufah, Masirali I., Oman (BMNH 1996.385). 41.

Lectohpe of Risella infracostata Ls.sel, 1869, Red Sea (MNHN). 42^8. Peasiella fuscopiperata. 42, 43, 46, 48. Tiwi Beach, 30

km S Mombasa, Kenya (BMNH 1996.386). 44. Leven Point, Zuhiland Sontli Africa (NM E2745). 45. Mzaniba, Transkei, South

Africa (NM .3028). 47. Mapi'lanc, Znhiland South Africa (NM D2415). Scale bar = 1 mm.

D. G. Reid and Y.-M. Mak, 1998

Page 13

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THE NAUTILUS, Vol. 112, No. 1

edge is curved (i.e. growing edge is prosocyrt). Sculp-

ture: protoconch 260 ^I,m diameter, 2.7 whorls, sculpture

not preserved; teleoconch whorls smooth or with 6-7

indistinct to strong spiral ridges; 4— .5 fine rihs on base;

surface smooth or with fine spiral microstriae; basal ribs

occasionally bear minute periostracal bristles. Color

cream to vellow ochre, with 4—7 fine bnmTi spiral lines

above periphery' (including dark line at suture; some-

times increasing to 9 at end of last whorl); 7-12 browm

obhque short stripes at periphery and suture, often faint

or absent, occasionally continuing indistinctly across dor-

sal surface; base with 1-2 spiral brown lines at peripherv',

margin of umbificus sometimes puiplish brown; colu-

mella and parietal area purplish brown.

Animal: Head-foot: usually entirely unpigmented;

sometimes slight blackish pigmentation behind head. Pe-

nis (Figures 84-87): filament slender, tapering, 30-40%

total length; base thick, single large mamilliform penial

gland (occup)ing 60-80% total length) with narrowly

elongate reservoir; subepithelial glandular tissue at base

of filament and around base of reservoir. Pallial oviduct:

bursa not seen. Egg capsule (Figures 123. 127, 128): di-

ameter 221.7 (j.m (SD 13.1 |jLm, n=10; ovum diameter

66.0±1.6 Jim; Singapore; this study), of depressed cu-

pola type with 3 concentric rings on upper side and

pleated margin (like a thick cogwheel), containing single

ovTim. Radula (Figures 112. 113): all cusps shaiplv point-

ed, central cusp of rachidian tooth slightly apiculate;

base of rachidian tooth with two sharp posterior denti-

cles; outer marginal tooth with .3—4 pointed cusps, be-

coming smaller toward midline.

Distribution: Habitat: Among barnacles and oysters

in upper euUttoral, on rocky shores and sea walls (Sin-

gapore).

Range (Figure 133): Singapore, Gulf of Thailand,

northern Borneo.

Records and material examined: Singapore: Changi

Point (BMNH; 1M,4F, 3R); Changi South (BMNH; 4M,

2F); St. Johns Island (BMNH; 2R, lP).Thailand: Pattava

(Le Renard Colin). Sabah: Berhala Channel, Sandakan

(USNM).

Remarks: The expansion rate of the shell is larger in

this species than in other members of the genus, so that

it achieves a sUghtly, but tlistinctly, patulous shape which

is characteristic. The color pattern resembles that of F.

fasciata, ;ilthough the fines are usually less pronounced

and the sutural and peripheral spots more developed.

These two species are distinguished b\ their penial form.

the filament being slender and short in P. patula, and

the mamilfiform gland relatively larger. Their egg cap-

sules differ in shape in samples of P. patula from Sin-

gapore and P fasciata from Hong Kong The\ have not

yet been found s\ntopically, but their known ranges ap-

pear to overlap in northern Borneo; additional collecting

in the region may well rev'eal wider SMiipatn. The radula

of P. patula is peculiar, showing a p;ur of shaip posterior

denticles on the base of the rachicfian tooth (Figures

112, 113). Among other littorinids, similar denticles have

been described only in Melarhaphe neritoides (Bandel,

1974: figs. 58, 59), but the stnictures are clo.ser to the

posterior edge of the tooth in P. patula, and are probably

not homologous. Sfight development of basal denticles

has been found in one example of P. fuscopipcrata and

one of P. isseli. One specimen from the tvpe locality

showed both a fulK- formed pallial oxiduct and a well

developed penis (see remarks on pseudo-hermaphrodit-

ism under P. habei). This is the rarest of all the Pcasiella

species in museum collections, perhaps because of its

limited geographical range; in Singapore it occurs abun-

dantly.

Pcasiella fasciata new species

(Figures '14-17, 49-51, 8.8-91, 106, 107, 117, 122, 125,

126, 134)

Peasielln infracostnta—Keid. 1989a:61-64, fig. 12, fig. .30 (rad-

vila), fig. 48 (penis), fig, 112 [in part, includes P infracos-

tata, P. fuscopipcrata, P patula, P. habei; not P. infracos-

tata (Issel. 1869)]. Reid, 1992:194, fig. Ic (penis), fig. 2c

(oviduct), pi. 1, figs, i, 1, 111 [in part, includes P. habei].

Etymology: Latin fasciatus, banded, in reference to

color pattern.

Types: Holotvpe BMNH 1996272 (Figure 15); 5 para-

types BMNH 1996273; 1 parat>pe AMS C203254. Type

locality-: Turtle Bay, Cape Ferguson, Queensland, Aus-

tralia.

SheU (Figures 14-17, 49-51, 106, 107): Adult size

range 1.3-2.5 nun diameter. Shape: turbinate to de-

pressed turbinate (H/D 0.78-1.12); outhne domed;

whorls well roundeil, suture distinct; peripher\' uniform-

1\ rounded, or angled, or marked by a more or less

prominent rib; base rounded; umbificus snuill or almost

closed; columella narrow, uniformly rounded. Sctdpture:

protoconch 240 [xm cfiameter, 2.6 whorls, protoconch I

smooth, protoconch II with scattered small granules and

narrow wavy or straight ridges, of which 6 are visible

above suture (Figures 106, 107); teleoconch whorls

Figures 49-51. Pcasiella fasciata, \}\o\ Mission. Cape Danipier, New Bntam, I'apua New Guinea (AMS C163128). 52-56. Pcasiella

patula. 52. Changi Point, Singapore (BMNH 1996278). 53-55. St. Johns I.. Singapore (BMNH 1996387), 56. Pata\a, Thiulaiid

(LR 990f2), 57-«l. Pca.'iiella habei. 57. 4-5 km SW Tsutsu, Tsushima I., Nagasaki Pref , Japan (BMNH 1996389). 58, 59. Nejiko,

Hirado I., Nagasaki Pref., Japan 60. Ma Liu Shi, Tolo Harbour, Hong Kong (BMNH i996.393). 61. Zanpa Point, Okinawa, Japan

(AMS C146702). Scale bar = 1 mm

D. G. Reid and Y.-M. Mak, 1998

Pa^e 15

Page 16

THE NAUTILUS, Vol. 112, No. 1

smooth above periphery, with fine spiral microstriae,

rarelv with .3—4 faint groo\-es: 3—4 ribs on base (rarely 1-

6), becoming stronger toward periphen: Color translu-

cent cream to opatjiie gra\ish white; pattern of brown,

dark red-brown or black lines or bands, numbering 2-8

above periphery on last whorl (only 2—4 on spire whorls),

bands often altematelv thick and thin on last whorl; in

addition sometimes 6-13 f;unt brownish spots at suture

and periphery ; 2—4 dark lines in grooves at periphery of

base; base of columella and umbilical area sometimes

dark purphsh brown.

Animal: Head-foot: front of head blackish, with a sep-

arate black band across base of snout; tentacles unpig-

mented or with single longitudinal black line; toot un-

pigmented or with blackish sides. Penis (Figures 88-91):

filament 40—50% total length, cylindrical and rounded at

tip; base thicker, with single maniillitorm penial gland of

moderately large size (30—40% total length). Pallial ovi-

duct: bursa present. Egg capsule (Figures 122. 125, 126):

diameter 215.4 ^.m (SD 5.8 jim, n = 10; ovum diameter

77.8±2.9 |jim; Hong Kong; this study), of depressed cu-

pola Kpe with 4 concentric rings on upper side and

pleated margin with peripheral keel, contiiining single

ovum. Radula (Figure 117): cusps pointed, central cusp

of rachidian tooth shghtly apiculate; outer marginal tooth

with 5-6 long, pointed cusps, outermost more rounded,

becoming smaller toward midline.

Distribution: Habitat: among algae in rock pools

(type locahtv; A. Mitchell); among short calcareous

green algae and Caidctya, on wave-exposed rocky plat-

form (Cape Dampier; P. H. Colman); among euhttoral

cobbles, cre\dces in granite and sedinientarv rocks,

among Saccostrca, on fairly sheltered and wave-exposed

coasts (Hong Kong; J. D. Taylor, Y.-M. Mak). Appears

to be generally scarce.

Range (Figure 134): Queensland, New Guinea, Bor-

neo, Vietnam, Hong Kong, Taiwan.

Records and material examined: Queensland, Austra-

Ua: Facing Island, Port Curtis (AMS); Turtle Bay, Cape

Fergu.son^AMS; IM, IR); ILdfmoon Bay, Cairns (AMS);

Green Island (AMS); Port Douglas (AMS). New Britain,

Papua New Guinea: Uvol Mission, near Cape Dampier

(AMS; 3M, 2F, 2P); Duke of York Island, Ral)aul (AMS).

Sabali: Bak-Bak, 9.6 km N Kudat (USNM). Vietnam: La

Table Island. Tonkin (MNHN). Hong Kong: Wu Kwai

Sha (BMNH; 2M, 2R); Peng Chan (BMNH; IM); Hoi

Ha (BMNH; 2F); C:ape d'Aguilar (BMNH). Tmwan: Shi

Cheng, Taipei (BMNH).

Remarks: The first evidenc-e that this species was chs-

tinct from P. hahei (both confused under P. infracostata

by Reid, 1989a, 1992) was provided bv their differently

shaped egg capsules in Hong Kong (Figures 122, 124).

Differences in shell shape and penial lorni were then

recognized, as describi'il in the Remarks on P. hahei.

The shell of P. fasciata is easily recognized by its com-

bination of dark spiral lines, tinliinate shape with well-

rounded whorls, and small umbihcus. Spiral color bands

are found in some other Peasiella species, especially on

the spire whorls, but are onlv well-developed on the last

whorl in one other species, P. patula. which is distin-

guished by its patulous shape. Anatomically, P. fasciata

is characterized by its penis with long, cylindrical fila-

ment and moderatelv large mamilhform gland. As in P.

infracostata and P. fuscopiperata. the shell can be

opaque and solid, or translucent, although it is never as

dehcate as in some examples ot those species. This spe-

cies is rare in collections.

Peasiella hahei new species

(Figures 18-24, 57-61, 92-98, 114-116, 124, 129, 130,

135)

Litlorinn-capsuln hahei Tokioka, 1950:151-1.52, fig. 6.1 (egg

capsule) [Ago and Taiiabe Bavs, Japan; name una\ailable,

see below]."Habe, 19.56:117-121.

Peasiella roepstorffiana. — Habe. 1956;lliS-121, fig. A (egg cap-

sule). Oyama & Takemura, 1961: Peasiella and Littorinop-

sis pi., figs. 1-3. Yaniamoto & Habe, 1962:16, pi. .3, figs.

3, 4, figs. 34, 35 (egg capsule). Amio, 1963:.303, figs. 22a,

b (egg capsule). Habe, 1964:28, pi. 9, fig. 23. Higo, 1973:

46. Habe, 1984:11. fig. 1. Ohtsuka & Yoshioka, 1985:232,

fig. 4B. Okutaiii, 1986:71, unnumbered pi Fukuda, Ma-

shino & Sugimura, 1992:57, pi. 43, fig. 126. Higo & Goto,

1993:74. [All not P. roepstotfiana (Ne\ill, 1885)[.

Peasiella infracostata— Reid, 1989a:61-64, fig. 10, fig. 29 (rad-

ula), fig. 47 (penis), fig. 64 (egg capsule), fig. Ill [in part;

includes P infracostata, P. fuscopiperata. P patula. P. fas-

ciata]. Choe, 1992:290, .591, fig. 56. Reid, 1992:194, pi. If

[in part: includes P fasciata]. Choe & Park, 1993: 19-20,

text fig. 2. [All not P infracostata (Issel. 1869)].

Etymology: To honor Prof T Habe, who has contrib-

uted much to the studv of Japanese httorinids.

Types: Holotxpe BMNH 1996274 (Figure 20); 9 para-

types ni alcohol BMNH 1996275. Type locidity: Esu

Cape, Shirahama, Wakayama Prefecture, Japan.

Nomenclature: Uniquely for a Uttorinid, the first

name for this species was applied to its egg capsules.

Tokioka (1950) described two types of httorinid egg cap-

sules in plankton hauls from Ago and Tanabe Bays in

southeastern Honshu, to which he gave Latin names.

Since the httorinids that produced the capsules were not

kniowni, he coined a new genus Littoiina-capsula. Sub-

secjnentlv, Habe (1956) showed that one of these cap-

sides, Littorina-capsula hahei, was the egg of the com-

mon Japanese species of Peasiella (which he misidenti-

fied as P. roepstoiffiana). However, it is debatable

wlu'ther the specific name gi\en by Tokioka is nonien-

ilaturalK a\ailable. Habe (1956) apparenth' considered

it so, printing it in italics. Others have not done this, and

thus appi^ar to have judged it unaviiilable (Amio, 1963;

Reid, 1989a). In its favour it can be argued that the

name is uniuubiguous, since the egg capsule is distinctly

different from those of other known Japanese httorinids.

A name based on ouK one stage in a fife cycle, while

undesirable, is not iherebv made unaNaikdile (ICZN Art.

D. G. Reid and Y.-M. Mak. 1998

Page 17

17). The generic name Littorina-capsula is una\;dlable

(since it is not accompanied bv fixation of a t\pe species,

ICZN Art. 13b), but this does not affect the standing of

a specific name combined with it (ICZN Art. llh). The

inclusion of a hvphen in the generic name is an incorrect

spelling (ICZN Art. 32c), and does not indicate that the

name is non-binominal. On the other hand, it appears

that Tokioka did not intend to introduce new names for

httorinid species, for he stated: For con\enience of re-

cording these capsules met with during our plankton

studies, I propose to give them the following provisional

names'. This, we believe, debars the names from no-

menclatural consideration, since the ICZN Code specif-

ically excludes names proposed "as means of temporan.

reference and not for formal taxonomic use as scientific

names' (ICZN Art. lb). This case is a complex and un-

usual one, and is open to altemati\e interpretations. If

Toldoka's name were to be accepted, the egg capsules

studied by him would have the status of type material;

inquiries from the Seto Marine Laboratorv have shown

that no such material exists there (S. '^'amato, pers.

comm.). In this case, our choice is to re-describe the

species as new, based on type material of adult aniniiils

collected near the locality of Tokioka's capsules. How-

ever, in recognition of Tokioka's discoverv of the egg cap-

sules, his intention to honor Prof T Habe, and to pro-

vide continuitv, we use the same specific name.

In the Japanese literature this species has generallv

been misidentified as P. rocpstotjfiana, while Reid

(1989a, 1992) incorrectlv included it under P infracos-

tata.

Shell (Figures 18-24, 57-61): Adult size range 1.6-

3.8 mm diameter Shape: equilaterally conical or sfightK-

more depressed (H/D 0.60-1.03); outhne domed; whorls

almost flat-sided or rounded or shghtlv shouldered, su-

ture inconspicuous or impressed; peripheral keel prom-

inent, often a projecting flange, rarelv slightlv undulat-

ing; base flat to slightlv rounded; umbilicus usualK' nar-

row; columella narrmv, cun'ed at base. Sculpture: pro-

toconch 230 |xm thameter, 2.5 whorls, sculpture not

preserved, apex usually eroded; teleoconch whorls usu-

ally smooth, with spiral microstriae, sometimes with 6-

11 equidistant spiral grooves above peripherv in largest

specimens; .3-.5 (rarelv 6) shaip ribs on base, equidistant

or becoming closer around umbilicus; basal ribs (and oc-

casionaUy dorsal ribs) may bear periostracal bristles. Col-

or: variable; fawn, cream or ochre, paler at peripherv,

with obhque (sometimes axial or zigzag) anastomosing

browm stripes (sometimes faint, especiaUv in Hong

Kong), always darkening to form single row of 7-12

more or less conspicuous large brown or black spots near

periphery (occasionallv fusing to give a sofid spiral band)

which become narrower as they extend over pale pe-

ripheral keel; pattern on spire whorls darker, often

blackish where eroded (especiallv in Japan); occasionallv

2^ spiral Isrown lines on spire whorls (common in Oki-

nawa), usuallv disappearing on last whorl, but rarelv per-

sisting as 5-9 spiral lines; base ochre to brown, darker

near umbilicus and in peripheral groove adjacent to keel;

sometimes shell more darklv patterned, spire and base

black, last whorl with obhque anastomosing black stripes

on whitish ground (some from Japan and northern Chi-

na).

Animal: Head-foot: front of head blackish, with sepa-

rate black band across base of snout, occasionallv almost

unpigmented; tentacles unpigmented; sides of foot

sometimes slightly pigmented. Penis (Figures 92-98): fil-

ament long (50-60% total length), vemiiforni; base

shghtly thicker, 1 (rarely 0 or 2) small mamilhform penial

gland. Pallia] oviduct: bursa present. Egg capsule (Fig-

ures 124. 129, 130): diameter 261.3 |xm (SD 0.5 |xm,

n=10; ovum diameter 68.3±2.3 |xm; Hong Kong; this

study), 150-250 jjim (Japan; Tokioka, 1950; Amio, 1963;

Ohtsuka & Yoshioka, 1985), of cupola Upe with 4 con-

centric rings on upperside, containing single ovum. Rad-

ula (Figures 114-116): cusps pointed, central cusp of

rachidian tooth shghtly apiculate, major cusp of lateral

and inner marginal teeth sometimes shghtlv rounded;

outer marginal tooth with 4-6 (rarelv 3) long, pointed

cusps, outermost more rounded, becoming smaller to-

ward midline.

Distribution: Habitat: In Japan this species is abun-

dant in crevices and among barnacles in the middle and

upper eulittor;il zone, on sheltered and moderatelv ex-

posed rock"\ shores; on exposed shores it shows a pref-

erence for surfaces protected from wave action (Mori et

al. 1985a, b; Tanaka et ai, 1985). In Hong Kong and

Taiwan it appears to be rare, and is found among bar-

nacles and oysters in the uppermost euhttoral zone on

sheltered shores.

Range (Figure 135): China, Korea, Taiwan, Rvukxii Is-

lands, Japan.

Records and material examined: China: Ma Liu Shi,

Tolo Harbour Hong Kong (RMNH; IM, IF, 2R); Lok

Wo Sha Bay, Tolo Channel Hong Kong (BMNH); Pao-

tai, Xiamen (BMNH; IM, IF); Gu Leng Yu, Xiamen

(BMNH; IM, 3F,1R, IP); Tai Ping Bay, Qingdao

(BMNH; 2M, 2F, IR); Hui Quan Bav, Qingdao

(BMNH). Korea: Taesori, Sangch'uja I. (Choe, 1992;

Choe & Park, 1993). Taiwan: Yehhu, Taipei (BMNH).

Japan: 1.5 km WNW Onna, Okinawa (AMS); Bolo Point,

Okinawa (AMS, IF, IR; USNM, NSMT); Baten, Oki-

nawa (BMNH; IM, 2F, 3R); Tomioka. Kumamoto Pref

(NSMT, 2R); Nagasaki (USNM); Omura Bav, Nagasaki

Pref (NSMT); Goto, Nagasaki Pref (AMS); Nejiko, Hir-

ado I., Nagasaki Pref (BMNH; IM, IF); Tsut.su, Tsush-

ima I., Nagasaki Pref (BMNH); Fukoivoshi, Fuk-iK)ka

Pref (KU); Kure, Hiroshima Pref (KU); Hanazura, Ko-

chi Pref (MNHN); Tatsukoislri, Kochi Pref (USNM,

NSMT); Shirahama, Tanabe Bay, Wakavama Pref.

(BMNH; 5M, 2F, IR; USNM, NSMT); Shionomisaki,

Wakavama Pref. (NSMT); Kuchino, Shiznoka Pref.

(BMNH, 4M, 2R); Arasaki, Kanagavva Pref (USNM,

NSMT); Oga Peninsula, Akita Pref (KU); Asamushi, Ao-

Page 18

THE NAUTILUS, Vol. 112, No. 1

mori Pref. (USNM, NSMT); 2.5 km N Tomari, Aomori

Pref. (AMS); Kominato, Aomori Pref. (LACM).

Remarks: This .species is variable in conspicuous fea-

tures of the shell including color, spire profile and sculp-

ture, but consistent characteristics are the row ol dark

spots above the periphery', which e.\tend onto the pale

peripheral keel, the darker and often black spire whorls,

and the prominent keel at the periphery. There are some

recognizable geographical trends in shell characters. In

Japan and northern China the shape is taller, with flatter

whorls and a sharper peripheral keel; periostracal bris-

tles are present at least on the base of most well-pre-

served specimens; die spire, usually eroded, is black or

darklv patterned (Figures 18-21, 57-59). Shells from the

southern parts of the range are distinctly different.

Those from Okinawa have a pale pattern; the dark pe-

ripheral spots are most conspicuous on the spire whorls,

vv'hich are also often marked bv 2-4 spiral brown lines;

the whorls are sometimes rounded at the shoulder, and

usually bear strong spiral grooves above the periphery,

and the peripher;il keel is less pronounced; periostracal

bristles have not been seen (Figures 23, 61). Shells from

Hong Kong and southern China are similar in shape to

those from Okinawa, but are usually smooth above the

periphery, sometimes bear periostracal bristles on the

base, and the color is ochre with strong peripheral spots

and darkly patterned spire (Figures 24, 60). However,

these southern populations are also variable, and inter-

mediates apparentlv connect them with the typical

northern forms (Figure 22). Additional material, partic-

ularlv from the R\-uk-\ii Islands, would be desirable to

investigate this \ariation in more detml, and to confirm

our interpretation. The penial shape is similar through-

out the range. The egg capsules from Hong Kong (fig-

ures 124, 129, 130) are almost identical in shape to those

figured from Japan by Toldoka (1950; reproduced b\ Ya-

mamoto & Habe, 1962; Reid, 1989a), Habe (1956) and

Ohtsuka & Yoshioka (1985). However, at a diameter of

261 fjim they are slightlv larger in size than Japanese

capsules, measured as 150-2()0(xm (Tokioka. 1950), 150

|xm (Amio, 1963) and 220-250 (xm (Ohtsuka & Yoshi-

oka, 1985).

The rounded shells with lined pattern on the spire

that can be found in Okinawa bear some similaritii- to

those of P. fcisciata from Hong Kong. This led Reid

(1989a) to cemsider them conspecific (as P. infracostata) .

However, new material from Hong Kong has showni that

both species occur there, although so tar the\ have not

been collected syntopicallv (i.e. on the same shores). In

Hong Kong the shells of P. luihci only occasionally bear

1-2 indistinct spiral brown lines above the periphery on

the last whorl, and their conspicuous peripheral spots

and darkl\' patterned spire differ from the 4—8 strong

spiral lines of local examples of P fa.sciata. Anatomically,

the mamilliform penial gland of P. Iiahei is smaller and

the penial filament relatively narrower and shghtly more

elongate than in P. fasciata. The egg capsules of these

two species (sampled in Hong Kong) also differ (Figures

122, 124).

Two other species are sometimes svnipatric with P.

luibci in the southern part of its range and might be

confused with it, P. hitulcnta in Hong Kong and P. roep-

stoijftana in the southern Rviik-yu Islands, Taiwan and

Hong Kong. PcasicUa hitulcnta has 5-9 strong spiral ribs

above the peripherv, usuallv a gap between inner and

outer series of ribs on the base (ribs all eijuidistant in P.

liabei), the columella has a thickened angulation at the

base, and the color pattern is of 7-13 broad and irreg-

ular, dark axial stripes (see Reid, 1989a, 1992). Anatom-

icallv the two are doselv similar; the penial filament of

P. Iiahci is shghtly narrower and more elongate, and

there are 4-6 cusps on the outer marginal tooth, but

only 3 in P lutulcnta. Peasiella rocpstotjfiana (Figures

142-145) has 4—13 strong spind grooves above the pe-

ripherv, frefjuently shows radial plications near the su-

ture and periphery, which give rise to strong crenulations

of the peripheral keel; the innermost one third of the

basal rathus lacks ribs; the color is often bright chrome

vellow or orange, with peripheral brown spots or band.

Anatomically, P. rocpstoijfiana from Hong Kong lacks

the mamilliform peniid gland, and the copulators- bursa

is absent.

Some abnormalities of the reproductive tract of P. lui-

bci have been found. A single specimen from Shiznoka

Prefecture had no maminiform penial gland (Figure 98),

and one (parasitized h\ trematodes) from Tanabe Bay

had two glands of normal size (Figure 97); in each case

normal specimens occurred in the same lots. One other

specimen from the latter sample had both a small, fully

formed penis and a small oxaduct. Genuine hermaph-

roditism has been described in onlv one littorinid genus,

Mainwariufiiii (Reid, 1986b), but pseudo-hermaphrodit-

ism (involving development of a penis and other male

characters in females) is known in several littorinid gen-

era (Reid, 19S6b, 1996), and is sometimes a response to

pollution bv organotin compounds (Bauer ct at.. 1995).

Peasiella species are not known to be tnily hermaphro-

ilite, antl tiiis example is probably a case of pseudo-her-

maphrocbtism.

Peasiella i.s.seli (Semper in Issel, 1869)

(Figures 2.5-29, 62-67, 99-101, 118, 119, 136)

Trochiis sp. Audonin, 1826:42

Riselhi isseli Semper in Issel, 1869:194, .347 [refers to Savigiiy,

Figures 62-67. Pctisicllti isseli. 62. Lectotype of Risella isseli Semper in Issel, 1869. Red Sea (MNHN). 63, 65, 66. Sue/., Egvpt

(BMNII ISSS.IO 14) 64, 67. llmijliada, Eirvpt (BMNH 1996.39,5). 68-75. Peasiella mauritiana 68. Tiwi Beach, 30 km S Mombasa,

Kenya. 69, 70, 73. Sedli. A! Kliassali, Oman. 71. Kuwait. 72, 74. Bandar Kliasran, Oman. 75. Bale Temay, Mahe, Seychelles.

Scale bar = 1 mm.

D. G. Reid and Y.-M. Mak, 1998

Page 19

Page 20

THE NAUTILUS, Vol. 112. No. 1

1817: pi 5. figs. .35.1, .35.2; lectohpe (Reid, 1989a: fig. 62)

+ 2 paralectohpes MNHN, seen; Suez (Reid, 1989a); 2

paralectohpes in MGD ex-Seniper Colin, seen, from Zan-

zibar are P mauritiana]. Pallarv, 1926:84, pi. 5, fig. 35.1.

35.2. Laniv, 19.38:71. Moiizzo, 1939:183. Bouchet & Dan-

rigal, 1982:13. fig. 65.

Risella {PeasieUa) isseli. — Nevill, 1885:160-161 [in part: in-

cludes P. mauritiana]. Trson, 1887:263, pi. 50. figs. .39, 40.

PeasieUa ).s.se/i.— Franc, 1956:25. Mastaller. 1979:40. Reid,

1989a:60-61, fig. 13, fig. 50 (penis), fig. 56 (sperm), figs.

97, 98, 101, 103 [in part, includes P. mauritiana].

Trochus sismondae Issel, 1869:225-226, pi. 2, fig. 13 [holotype

MGD, seen; Suez].

Risella isseli var. carinata Pallarv', 1926:84-85, pi. 5. fig. 36 [re-

produced from Savngnv, 1817: s\mt\pe MNHN, seen;

Suez].

Risella i.sseli van undata Pallarv', 1926:84, pi. 5, fig. 35.3 [re-

produced from SaMgnv, 1817; t\pes not in MNHN; Suez].

Nomenclature: Throughout its taxonomic history this

species, endemic to the Red Sea, has been confused with

P. mauritiana, and the two taxa have not previously been

distinguished at the specific level. In the original de-

scription, Issel referred to the figures of Sa\dgnv (1817)

based on Eg\ptian specimens. He also mentioned his

own material from Suez and Semper's from Zanzibar, of

which the latter (in MGD) is P. mauritiana. Reid (1989a)

designated one of the shells from the Sa\ignv Collection

as the lectot)'pe, and restricted the type locality to Suez,

thereby fixing the identity of this species.

Shell (Figures 25-29, 62-67): Adult size range 1.7-

4.2 mm diameter Shape: conical to depressed conical

(H/D 0. 5.5-1. 23); outhne domed; whorls usually with

rounded or angled shoulder, occasionalh' flat or only

gently rounded; suture usually distinct; periphery sharply

angled, with prominent rib or flange; dorsal sculpture

sometimes slightlv nigose, but only rarelv does periph-

eral flange show slight crenulation; base flat or slightly

rounded; umbilicus small to moderate; columella round-

ed, sometimes with a slightly thickened angle at base.

Sculpture: protoconch 240 |xm diameter, 2.3 whorls,

protoconch II v\ath scattered small granules and narrow

wavy or straight ridges, of which 5 are visible above su-

ture (as in Figures 106, 107, of P. fasciata): teleoconch

whorls with 6—15 narrow ribs, uniform or unequal in

size, above periphery; rib at shoulder sometimes slightlv

enlarged; surface above periphery' covered with fine spi-

ral microstriae; base with 4-6 (rarely 2-7) ribs, larger

toward peripherv: Color: extremely variable; uniform

cream, ochre, orange-browai, dark chestnut brown or en-

tirely black; often with darker pattern of fine dots or

lines in dorsal grooves; dorsal surface sometimes with

fine white and brown marbled pattern; rarely a dark spi-

ral band on dorsal surface; sometimes 7-15 brown spots

on white peripheral keel; rarely pale sutural spots in ad-

dition to darker peripheral spots, but only very- seldom

does an indistinct pattern of broad radial bands appear;

spire sometimes pink or crimson; apex often lilac or

blackish; base lacking color pattern or with small spots

in grooves.

Animal: Head-foot: head unpigmented or blackish in

front, separate black band across base of snout; tentacles

usually with 2 longitudinal black fines; sides of foot with

some black pigmentation or unpigmented. Penis (Fig-

ures 99-101): filament long (509f total length), tapering

at tip; base thicker, with single mamilfiform penial gland

of moderate size. Pallial oviduct: large anterior bursa

present. Radula (Figures 118, 119): cusps pointed, major

cusp of lateral and inner marginal teeth slightK rounded,

central cusp of rachidian tooth sometimes slightK- apic-

ulate; base of rachicfian tooth occasionally with slight de-

velopment of pair of posterior denticles; outer marginal

tooth with 4 pointed cusps, outermost more rounded,

becoming smaller toward midline.

Distribution: Habitat: Abundant under coral blocks

in upper eulittoral on a moderatelv sheltered shore

(Egypt; D. G. Reid).

Range (Figure 136): Red Sea and Aden.

Records and material examined: Israel: Eilat (BMNH,

IM, 2F; USNM, ANSP). Egypt: Faraun Lsland, Gulf of

Aqaba (HD); Ras el Kura, Daliab, Sinai (HD); Suez

(BMNH, IP, 2R; MNHN); Suez Canal (Tillier & Bavay,

1906; Moazzo, 19.39); Hurghada (BMNH, 3M, 2F 2R;

HD). Sudan: Dungunab Bay (BMNH); Port Sudan

(BMNH; ANSP). Saudi Arabia: Jeddah (USNM,

MNHN). Eritrea: Mas.sawa (BMNH, NM); Dahlak Ar-

chipelago (BMNH). Yemen; Perim Island (MNHN);

Aden (MNHN).

Remarks: The shells of P. i.s.seli and P. mauritiana are

very similar, and the two have not previously been dis-

Figures 76-79. Penes of PeasieUa infracostata 7fi, 77. Tivvi Beach, 30 km S Mombasa, Ketna. 78. Karachi. Pakistan. 79. Ras

al-Hadd, Oman. 80-83. Penes of PeasieUa Juscopiperata. 80, 81. Tiwi Beach. .30 km S Mombasa, Kenva. 82, 83. Mapelane,

Zululand, South Africa (NM D5822). 84-87. Penes of PeasieUa patula 84-86. Cliangi South, Smgapore' 87. Paratvpe, Changi

Point, Singapore (BMNH 1996278). 88-91. Penes of PeasieUa faseiata 88, 89. U\ol Mission. Cape bampier. New Britain, Papua

New Guinea (AMS C163128), 90. Wu Kwai Sha, Hong Kong. 91. Peng Cliau, Hong K.ong. 92-98. Penes of Pea.siella habei. 92.

Tai Ping Bay, Qiugdao, China. 93. Sliirahama, VV'akayama Pref , Japan. 94. Batcn, Okinawa, Ja]ian 95-97. Fujisliima, Tanabe Ba\.

VVakayama Pref., Japan. 98. Kuchino, S\miga Bay, Sliiznoka Pref., Japan. 99-101. Penes of PeasieUa isseli. Hurghada, Egvpt. 102-

105. Penes of PeasieUa mauritiana 102. Khor Hajar, Ras al-Hadd. Oman. 103. Anse La Mouche, Mahe, Seychelles. 104. Mera-

wwah I., Abu Dhabi. 105. Tiwi Beach, .30 km S Mombasa, Kenva. Note that stnictnre of mamilliform penial glands is visible by

transparency; mucous reservoir is shown by solid line, and extent of subepithelial glandular tissue by dotted line. The penial base

is .sometimes wrinkled.

D. G. Reid and Y.-M. Mak, 1998

Page 21

76

77

78

79 80

81 82 83

92

93 94

95

96

97

98

99 100

101

102

103

104 105

Page 22

THE NAUTILUS, Vol. 112, No. 1

tinguished as separate species. From the new anatomical

material av;dlable for each species, it is clear that they

can be diagnosed by the form of tiie penis. This is of

the common type for the genus in P. isseli, with long

filament and small mamilliform gland, whereas in P

mauiitiana from throughout its large range it has a short

filament with mucronate and hooked tip, and a large

gland. Having separated the two using this character,

correlated differences in shell shape and coloration can

be recognized. In both, the development of the shoulder

angulation is variable; in P. isscli it is often absent or

represented only by an angulation of the profile, more

rarely bv a prominent rib; in P iiiaiiritiana the shoulder

is usuidly marked by a shaip angulation, often with an

enlarged or even carinate rib. The coloration is also dif-

ferent; in P. mauiitiana there is usually a pattern of

obliquely ratlial lines or stripes, often darker at the su-

ture and periphery; in P. isseli the color is highly variable,

but radial stripes rarely appear, and the apex is often

pink or blackish. Using these characters, together with

knowledge of the regional variation in the shell of P.

mauritiana, it has been possible to identify shells with

confidence. So far, it appears that P. isseli is restricted

to the Red Sea and Aden, whereas P. mauiitiana is wide-

spread in the Indian Ocean and Persian Gulf The two

apparently occur sympatrically just at the mouth of the

Red Sea. Two dead shells of P. isscli have been recorded

from Aden (MNHN), from which locality five other sam-

ples contained only P. mauritiana. Both have been found

together in a single lot of dead shells from Perim Island,

South Yemen (Jousseaume Colin, MNHN), which con-

tained 50 P. isseli and 8 P. mauritiana, readily separated

by their .shell characters.

The only other species to occur sympatrically with P.

isseli is P. infracostata. These two are readily distin-

guished by the presence of spiral ribs above the periph-

ery in P isscli (absent in P. infracostata}, anil by the color

pattern of obh(jue or zigzag lines in P. infracostata. The

shape of the penis is, however, similar in both.

Most Peasiella species show variation in color pattern,

sometimes on a regional or inteqiopuhitional scale. This

species is unique in the genus m its extreme intrapop-

ulational variation; shells from beneath the same rock

mav range from cream to p;itterned to black. Further-

more, occasional specimens can be found in which the

color changes from cream on the spire to entirely black

on the last whorl (Figure 26).

Peasiella mauritiana (Viader, 1951)

(Figures .3()-.34, 68-75, 102-105, 120, 121, 1.37)

Ri.sella mr/i — Lssel, 1869:194 [in part; includes P. mr/i].

Riwlta {Peasivlla) me/i— Ne\-ill, 188.5:160-161 [in part; in-

cludes P issi'ti]. MeK-ill & Standen, 1901:.364 [not Semper

in lssel. 1869[.

Peasiella me/i.— Reid, 1989a:60-61, figs. 14, 1.5, fig. 28 (rad-

ula), fig. 49 (fiead), fig. 97, 99, 100, 102, 104, 105 [in part;

includes P isseli]. Bosch et ai, 1995:46, fig. 119 [not P.

isseli (Semper in lssel, 1869)].

Ri.selln infracostata. --Ddutzenherg, 1929:496 [not lssel, 1869].

Risella is.seli van mauritiana Viader, 1951:149, pi. 3, figs 10, 11

[lectotype, here designated. Figure .30, -I- 2 paralecto-

types, 'seen, BMNH 1989004; Grand Bay, Mauritius; 2

probable paralectotvpes Mauntius Institute, not seen].

Nomenclature: Hitherto, this species has always been

confused with P isseli. It was first named, as a variety

of that species, bv Viader (1951). Twt) specimens of Ri-

sclla isseli var. mauritiana were exhibited in the natural

history museum of the Mauritius Institute (R. Gajeelee,

pers. comm. 1985); these have not been examined, but

are likely to have been syntypes. In addition, there are

3 specimens in the Winck-worth Collection in BMNH,

labeled 'Peasiella isseli v. mauritiana Viader MS' in

Winckworth's hand; these are from Grand Bay, one of

the two lociilities mentioned in Viader's (1951) descrip-

tion. It is knowai that Winck-worth received specimens

from Viader (see also acknowledgement of Winck-worth's

assistance in determinations, Viader, 1951:1.36), and it is

likely that this material was sent by Viader before Winck-

worth's death in 1950. Since Viader (1951) chd not des-

ignate tvpe specimens, a lectotype is here designated

from the BMNH materiid (ICZN art. 74; recommen-

dation 72B).

Shell (Figures 30-34, 68-75): Adult size range 1.6-

5.6 nun diameter Shape: highly variable; depressed con-

ical to Hattened (H/D 0.42-1.03); outline ilomed,

straight or spire sfightly concave; shoulder usually sharp-

Iv angled, or with projecting rounded carina (some Per-

sian Gulf and northern Oman specimens), or sometimes

becoming rounded or even flattened on last whorl; su-

ture usuallv distinct; periphery sharply keeled or flanged,

or with flaring rounded carina (some Persian Gulf and

northern Oman specimens); shoulder occasionally reg-

idarlv nigose and peripheral flange with 7-8 slight cren-

ulations (some East African specimens); base flat to

rounded; umbilicus small to large; columella rounded,

sometimes indistinctlv thickened and angled at base.

Figures !()(>, 107. Frotoconch of Peasiella fasciatn, Uvol Mission. Cape Danipier, New Britain, Papua New Guinea (AMS

C163128). Figures 108, 109. Radulae oi Peasiella infracostata. 108. Karachi, Pakistan (flat view, shell D = 1.5 nun). 109. Ras al-

Hadd, Oman (flat view, shell D=1.9 mm). 110, 111. Two \iews (flat and at 45°) of radula of Peasiella fiiscopiperata, Tiwi Beach,

30 km S Mombasa, Kenya (slieii D=1.7 mm) 112, 113. Two views (flat and at 45° from side) of radula of paratype o( Peasiella

patula. Changi Point, .Singapore (BMNH 199627S, shell D=1.7 mm); note posterior denticles on base of rachidian tooth (arrow).

Scale Inirs: 106, 107 = 100 |jlui; 108-113 = 20 jxm.

D. G. Reid and Y.-M. Mak, 1998

Page 23

Page 24

THE NAUTILUS. Vol. 112. No. 1

Sculpture: protoconch 220 [im diameter, 2.2 whorls,

protoconch II with .scattered small granules and narrow

wa\y or straight ridges, of which 6 are visible above su-

ture (as in Figures 106. 107. of P. fasciata); teleoconch

whorls usually with 6-12 fine spiral ribs above periphery;

sometimes onlv single rib at shoulder, or absent; surface

above peripher\' covered with fine spiral microstriae;

base with 4-7 (rarely 1-3) ribs, becoming weaker and

more closely spaced around umbiHcus; periostracum

thicker than in other species of the genus, and may flake

off from dn- shells; ribs on base and rarely on dorsal

surface mav bear periostracal bristles (some Persian Gulf

and northern Oman specimens). Color: cream to ochre;

usually with dark brown oblique (opisthocline) hnes or

narrow stripes, numbering 6-11. often darkest or broad-

est at suture and periphery; in northern Oman and Per-

sian Gulf oblique lines may number up to 23 on last

whorl, anastomosing to produce 8-15 spots at suture and

periphen-; diagon;il stripes occasionally present only at

suture and periphery (some East African specimens); di-

agonal pattern only rarely entirely absent or developed

only on spire (most specimens from Seychelles); often a

single brown spiral line on early spire whorls; base un-

pattemed or with small brown spots on ribs, occasionally

forming radial fines; parietal callus, columella base and

umbifical margin sometimes browaiish.

Animal: Head-foot: front of head blackish or with nar-

row black band only, separate black band across base of

snout; tentacles with 2 longitudinal black lines; sides of

foot with some black pigmentation or unpigmented. Pe-

nis (Figures 102-105): filament short (20-30% total

length), with elongate, mucronate tip, distinctly hooked

or minutely hammer-shaped at very tip; base thicker,

finelv wrinkled, with single large mamilliform penial

gland (rarely absent) with elongate reservoir. Pallial ovi-

duct: bursa present, apparently at posterior end of

straight section. Radula {Figures 121, 122): cusps point-

ed, major cusp of lateral and inner marginal teeth slight-

ly rounded, central cusp ol rachidian tooth slightly apic-

ulate; outer marginal tooth with 4 pointed cusps, out-

ermost more rounded, becoming smaller toward mid-

fine.

DLstribution: H(i]>itat: intertidal rocks, pools and Tlwl-

as.sodr lid roil Hats in a sheltered bay (Inhaca Island; R.

N. Kilburn); barnacle zone on beach rock (Aldabra; J.

D. Taylor); among barnacles in high eufittor;il, on surf

beach (Masirah Island; J. D. Taylor); crevices in fime-

stone, in upper eulittoral, on sheltered coast and in tidal

lagoon (Ras al-IIadd; E. Glover); among baniacles on

mangrove pneumatophores (Abu Dhabi; D. George);

among Crassostrea (Kuwait; D. Jones).

Ran^e (Fiffire 137): Zululand to Persian Gulf, Mauri-

tius, Madagascar, Seychelles; excluding Red Sea.

Records and material examined: South Africa: Mission

Rocks. Zululand (NM). Mofambique: Inhaca Island

(NM, IM); Benguera Island (NM); Mozambique Island

(NM). Tanzania: Zanzibar (BMNH). Kenya: Tiwi Beach,

30km S Mombasa (BMNH, 2M. 2F). Madagascar: Tu-

lear (MNHN); Maliajanga (IRSNB); Nossi-Be (USNM.

ANSP MNHN. ZMA); Diego-Suarez (IRSNB). lie Ma-

yotte (MNHN, IM). Mauritius: Grand Bav (BMNH,

NM); Pointe Radeau, S of Roches Noires (NM); Pointe

d'Azur (NM). Seychelles: Aldabra (BMNH); Baie Ter-

nay, Miilie (BMNH, 2R); Anse la Mouche, Mahe

(BMNH, IM, IF; USNM); Northwest Bay, Mahe

(ANSP). Djibouti (MNHN). Yemen: Perim Island

(MNHN); Little Aden (BMNH, 2F); Aden (BMNH,

MNHN). Oman: Hamran. Zufar (BMNH); Sudh

(BMNH); Sadh (BMNH); Kuria Muria Islands (ZMA);

Masirah Island (BMNH. IF; ZMA. 2F); Sedli. Al Khav-

sah (BMNH); Khor Hajar, Ras al-Hadd (BMNH. IM,

2F, IR); Bandar Kliayran (BMNH); Al Bustan, 3.2 km

E Muscat (ZMA); Muscat (NM); N of Sharm (BMNH).

Abu Dhabi: Merawwah Island (BMNH. 3M, IF). Qatar:

Khor. Doha (BMNH). Kuwait (BMNH).

Remarks: Although shape and coloration are highly

variable in this species as a whole, some geographical

patterns can be tfistinguished. In southern Arabia the

spire is a little taller, the shoulder more rounded, the

peripheral keel less pronounced, and spiral sculpture on

the dorsal surface is weaker or even absent (Figures 33.

69. 73). In the Persian Gulf and northern Oman there

is a tendency to flattening of the spire, and dexelopment

of heaw carinae at the shoulder and peripher\-. while

the dark obli(jue fines become more numerous (Figures

31, 71, 74). Specimens from Kenya and Zanzibar are

more nigose. sometimes with regular racfial folds and

slightlv crenulate peripheral flange (Figures 34. 68). In

the flattened and weakly shouldered specimens from the

Sevchefles (Figure 75) the fined color pattern is absent

or occasionallv present on the spire only. In typical spec-

imens from the Mascarene Islands. Madagascar and Mo-

9ambi(jue, the shoulder is scarcely marked and the color

pattern is pronounced (Figures 30, 32).

The shell of this species is most likely to be confused

with tliat of/' (.s,st7( (see Remarks on that species), which

Figures 114-116. Radulae of Peasiella halm. 114. Baten, Okinawa, Japan (flat view, shell D=3.2 mm). 115. Ma Liu Shi. Hong

Kong (view at 45°. shell D = 2.2 mm). 116. Gu Leng Yu, Xiamen. China (view at 4.5° from side, shell D=1.8 mm). 117. Radula of

paratvpe of Pcasidla fasciat a. Turtle Bay. Cape Ferguson, Queensland. Australia (BMNH 1996273; view at 45°, shell D = 2.1 mm).

118,'ll9. Radulae of PrasirUa mc/i. 118. Hunj;liada, Kgvpt (flat view, slid! D = 2.4 mm). 119. Hvirghada. Egvpt (view at 45°, shell

D= :?,() luuii. 120, 121. Two views (flat and at 4.5°) of radula of AY;.vic//« inmiritiana. Khor Hajar. Ras al-Hadd. Oman (shell D=3.6

mm). Scale hars = 20 (Jini.

D. G. Reid and Y.-M. Mak, 1998

Page 25

Page 26

THE NAUTILUS, Vol. 112, No. 1

122

123

124

-T^^X

100 ;jm

Figures 122-124. Egg capsules of Pcnsiella species. 122. Pcasiclla fnsciata. Cape d'Aguilar, Hong Kong. 123. PeasieUa patida.

Changi Point, Singapore. 124. PensU-Un Itahri. Ma Liu Shi, Tolo Harbour, Hong Kong.

is endemic to tlie Red Sea; the two are only known to

be sympatric close to the mouth of the Red Sea. Two

other species, P. infracostata and P fuscopipcrota. can

be found s\nitopically with P nmuiitiana, and all three

have diagnostic penial shapes. The shell of P. infracos-

tata is alwavs smooth abo\'e the periphery, and is thus

easily distinguished from all ribbed and shouldered

forms off matiritiana. However, examples of the latter

from southern Oman and Yemen may lack ribs above

the peripherv, and if the shoulder angulation is poorly

developed mav be confused with P. infracostata. The ba-

sal ril)s may then be useful; in P. mmiritiana the outer-

most rib is the largest, but in P. infracostata the mid-

basal rib is most prominent. Rare examples of P. fiisco-

piperata which ha\'e spiral sculpture above the periphery-

could lie confused with this species, lint lack the shoul-

der angulation.

ADDITIONAL NOTES ON OTHER SPECIES OF

PEASIELLA

PeasieUa roepstorffiana (Nevill, 1SS5)

(Figures 142-145)

Di.stribution: Additional records: Indonesia: Ambon

(NN.ML). T;iiwan: Kcnding, Kaohsiang (BMNH); Yehhu,

Taipei (BMNH). Japan: Kauuon Saki, Ishigaki, Rvuk-yu

Islands (BMNH).

Remarks: The range of this species is now extended

from Hong Kong (Reid, 1989a, 1992) to T;iiwan and the

.southernmost RvTikyu Islands (Figures 142, 144, 145),

and also includes Ambon in Indonesia. A curious feature

of this species is the apparent dimoiphism ol penial

form; a mamilliform gland is present in material from

Queensland, but not in specimens from Asia (Reid,

1989a). The absence of a penial gland has been con-

firmed in m;it('rial from Ishigaki and .Xmbon. .Vdditional

anatomical material has also been seen from Lautoka,

Viti Levu, Fiji (BMNH); penial glands were found to be

absent in two specimens, but in a third a nidimentary

gland was present, with a cluster of subepithehal tissue

but no reservoir or papilla. This supports the earlier sug-

gestion that the peni;d dimorphism is indeed intraspe-

cific. A similar case of geographic;d \ariation in presence

or absence of a single penial gland has since been re-

corded in Littoiina kasatka (Reid, 1996). The new Fijian

material also supports the identification of the distincti\'e

darklv tessellated shell from the eastern part of the range

as merelv a color variant of this species. Sb; additional

radulae examined (Hong Kong, Singapore, Tiiiwan, Ja-

pan) each have 3 cusps on the outer marginal tooth, as

recorded by Reid (1989a). The egg capsule of this spe-

cies was described bv Mak (1995) from Hong Kong; it

is nuich larger than those capsules described here, 360-

390 |xm in diameter, and similar in shape to that of P.

hahei, although without the h\'o inner concentric rings

on the upper surface.

Pcasiclla conoidalis (Pease, 1868)

(Figures 138-141)

Remarks: AdtlitionJ anatomical materi;il of this species

has been seen from Taiwan and Ishigaki, Ryuk)ii Islands

(BMNH). The penial shape is the same as that illustrated

b\- Reid (1989a), with a small and slender penial gland.

Shells from these loc;Jities are often darker in cok)r than

those found elsewhere, with brown dots (or occasionally

fine lines) in the spiral grooves abcne the peripher)- and

strong brown marks in the folds behveen the peripheral

creiHilations (Figin-e 138). In some shells the pattern con-

.sists of 1(>-17 dark browni axial stripes (Figiue 140). Peas-

ieUa rocpstoiffjana ;ilso occurs at these localities, and

shells of the t\\t> are ea.silv confused since both are veUovv,

D. G. Reid and Y.-M. Mak, 1998

Page 27

' i.

125

126

im

]

I'

127

129

130

Figures 125-130. Egg capsules oi' Ptnsirlla species. 125, 126. Peasiella fasciata. Cape d'Aguilar, Hong Kong. 127, 128. Peasiella

pattiln. Chaiigi Point, Singapore. 129, 130. Pensiclln hahci. Ma Liu Shi, Tolo Harhour. Hong Kong Scale bar = 200 \xm.

have radially plicate sculpture and peripheral crenulation.s

(Figures 138, 141, 142, 145). They are separated by the

following characters: in P. conoidalis the spiral grooves

above the periphery usually contain sni;ill lirovMi dots

rather than continuous lines and the periphenil lirovvni

marks are weaker; die middle of the base of P. conoklalis

shows 1-2 rows of small brown spots, whereas that of P.

roepstoijfiana is unmarked but for dark blotches on the

underside of die peripheral keel and puiple-tinged colu-

mella; the wide unsculptured area surrounding the mn-

bilicus is characteristic of P. roepstoijfiana: anatomically,

the single mamilliform penial gland of P. conoklalis is

lacking in P. roepstorffiana (note that these characters do

not necessarily apply to the two species in other parts of

their ranges, see Reid, 1989a). These two species are only

occasionally found SMiipatriciillv elsewhere (e.g. Lizard Is-

land, Australia; soutlieasteni Papua New Guinea; Ambon,

Indonesia; some islands in Fiji and the Philippines), since

P. conoidali.s has an oceanic distribution while that of P.

roepstoijfiana is more continental in character. An un-

identified Peasiella species from Chichijima, in the Oga-

sawara (Bonin) Islands, illustrated bv Fukoida (1993), is a

Page 28

THE NAUTILUS, Vol. 112, No. 1

small, worn example of this species (specimen now in

BMNH). PeasieUa corwidalis has also now been recorded

on die African mainland for die first time, from Tiwi

Beach, Kenva (BMNH), where it occnrred v\ith P. infra-

costata, P. fuscopipcrata and P. mauiitiona. Another new

record is from Ambon, Indonesia (NNML).

Pca.siclla lufiilcuta Reid, 19S9

Remarks: This species is now recorded from Singa-

pore (BMNH), where it appears to be rare. One of the

five specimens found was a female with a mature paUial

oviduct as well as a small penis (see Remarks on P. habci

and P. pattda). Two additional radulae examined each

have 4 cusps on the outer marginal teeth, whereas Reid

(1989a) found 3 cusps.

PeasieUa petiti Le Renard, 1994

Remarks: This species from the Eocene of France

was described and figured as P. minitta (Deshayes, 1824)

bv Reid (1989a); this name is preoccupied, and the spe-

cies was renamed by Le Renard (1994).

DISCUSSION

Snf:i,l, V.\RIATION

Our reassessment of the taxonomy of PcasicUa has not

altered the earUer conclusion (Reid, 1989a) that the shell

characters of these littorinids are unusualK- variable.

Elsewhere in the family, species with likewise plankto-

trophic development do not normally show such con-

spicuous variation (e.g. Reid, 1986a, 1996). The varia-

tions in whorl outline and sculpture are most striking,

since these characters are traditionallv used to diagnose

httorinid species. For example, in P. injracostata and P.

fuscopiperata the periphery may be strongly keeled or

smoothlv rounded. In P. isscli and P. maitritiana the

whorl profile may be almost flat or may develop a prom-

inent carina at the shoulder In P. fuscopiperata, P. pa-

tula, P. habei and P. mauritiami the surface above the

periphery may be smooth or bear spiral ril)S. Shell thick-

ness is also \ariable, ranging from solid and opaque to

thin and translucent in both P. injracostata and P fus-

copiperata. One sculptural character that is more useful

for identification is the number and size of the ribs on

the base. Manv littorinids arc conspicuously variable in

shell color and pattern, and this is also the case in some

PeasieUa species, notably P isseli. Nevertheless, we have

found the number and arrangement of the obUque or

spiral color bands t(j be one of the more useiul charac-

ters for their identification. The cause and adapti\e sig-

nificance of shell variation in non-planktotrophic littorin-

ids has been the subject of much researcli, but that oi

the plank-totrophic species is less well understood (re-

views by McQuaid, 1996; Reid, 1996).

Anatomy

Throughout the family Littorinidae the shape of the pe-

nis is known to be one of the most useful taxonomic

characters, and probably plays a role in species recog-

nition in nature (e.g. taxonomic accounts and reviews by

Reid, 1986a, 1989b, 1996). It was therefore surprising

that the prexaous account oi PeasieUa (Reid, 1989a) con-

cluded that penial shape was similar throughout the ge-

nus. All the six species for which anatomical materiid was

then available showed a penis with a long filament and

a single mamiHiform penial gland (although in P rocp-

storffana the gland was found to be absent over much

of the geographical range). The new material available

to us in the present study has re\'ealed that s\nipatric

species do in fact frequenth' show diagnostic differences

in penial form, particularly in the relative sizes oi fila-

ment and penial gland.

The gross moipholog\' of the female reproductive sys-

tem does not, however, show useful diagnostic charac-

ters. The arrangement of the loops of the egg groove

through the glands of the palfial oviduct is the same in

all 11 species oi PeasieUa (as illustrated bv Reid, 1989a:

figs. 57, 58, 65, 66). The copulaton bursa is apparently

absent in six species, but this character is not useful for

identification since it can be difficult to trace without

histological preparation. Furthermore, in at least one ht-

torinid, Littorina saxatilis. there is intraspecific variation

in this feature, for the bursa is sometimes vestigial or

absent in small, but mature, adults (Reid, 1996: 312).

The egg capsules of four PeasieUa species have now

been described (P habei, P. patula, P. fasciata, herein;

P rocpstotffana in Mak, 1995). At least in P habei the

form of the capsule is constant over a wide geographical

range, although there is some variation in size (see de-

scription of this species). These four species each show

a unique capsule shape, and there is an almost twofold

range of diameters among them. Similar interspecific

variation is present in the genera Wxhlittoiina and Lit-

toraria (reviews bv Bandel 6c Kadolsky, 1982; Reid,

1986a, 1989b; Mak', 1995), although in most cases intra-

specific variation is poorly kiiovvni. The radial plication of

the capsules of P patula and P. fasciata has hitherto been

described onlv in Sodilittorina species

In general, the radula of fittorinids is taxononiicallv

useful for the diagnosis of generic groups (Reid, 1989b),

lint not at the species level (e.g. Reid, 1986a, 1996). As

in tlu' earlii'r studv, we have found the radula of PeasieUa

species to be relatively uniform in the genus, showing

interspecific variation onlv in the mmiber of cusps on

Figures 131-135. Distribution maps tA l','/isirlln species, compiled from records listed in text. 131. PeasieUa iufnuosltitn. 132.

PeasieUa fiisiopipmild 1.33. Pi-asiillii patula 1.34. Priisii-lhi fiisciata 1.3.5. Pitisiclla hahei.

D. G. Reid and Y.-M. Mak, 1998

Page 29

Page 30

THE NAUTILUS, Vol. 112, No. 1

Figures 136, 137. Distrilnition nmp.s of Piasiella species, compiled t'roni records listed in te.\t. 136. Peasiella isseli. 137. Pensiella

mnuritiana.

Figures 138-141. I'casirUa ronoidalis. Kcndint;, Kaolisiaiii;, Taiwan (BMNH 199(i-59.S). 142-145. I'laswUa nvpstorffiana. 142.

Kannon Saki. Isliiijaki. Japan (HMNd 1996:5991. 143. Ma Lin Slii, Honj; Kong (BMNH 1996400). 144. Yelilin, Taipei, Taiwan

(BMNH 1996401). 14.5. Kciidini;, Kaolisianj;, Taiwan (BMNH 1996402). Scale bar = 2 mm.

D. G. Reid and Y.-M. Mak, 1998

Page 31

tile outer marginal tooth, ami m the development oi ha-

s;il denticles on the raehiilian tooth.

Phvi,(ic;env and Bi()(:i'.()(;hai'ii\

The phvlogenetic relationships of PcasicUa within the

Littorinidae remain uncertain. In the nioqihological

phvlogenetic analvsis of Reid ( iy89h), the gcTius ap-

peared as the sister-taxoii ot Maimcahngia in the con-

sensus of all equallv parsimonious trees; however, no

unique s\iiapomoq)hv tor these two taxa was found. An-

atomicalK, thev are similar in their penial form (single

mamilliform gland and closed sperm duct), pallial om-

duct loops (although these are shghtlv more complex in

Mainwaringia. Reid, 1986b) and egg capsule shape

(compare Reid, 199():fig. 3j with Mak, 1995: fig. 2d), hut

some of these resemhlances mav he plesiomoqihic. Thev

are so different in their shell and opercular shape, rad-

ular teeth and in the unique hermaphroditism oi Main-

waringia, that a close relationship might seem unlikely.

Nevertheless, prehminar\' molecular data offers some

support for their sister-group relationship, and is cur-

rentlv under investigation (R. H. Thomas, pers. comm.).

Morphological features do not provide sufficient in-

formation for a cladistic analysis at the species level, and

in the absence of an une(juivocal sister-group the polar-

ities of the few aviulable characters are doubtful. The

five species confused by Reid (1989a) under the name

P. infracostata (P. infracostata. P. fiiscopipcrata. P. pa-

tula. P. hahci, P fasviata) are a possible clade, showing

some similariU in shell form (tendencv to reduction of

peripheral keel and spiral ribs, presence of spiral cokjr

bands), and in radular cusps (4^8 cusps on outer mar-

ginal tooth, except 3—4 in P. pattila: cf 3-4 in all re-

maining species).

Four probable pairs of sister-species can be suggested.

PeasicUa infracostata and P. fuscopipcrata are closeK

similar in shell characters; both lack a copulators' bursa,

and radular cusps are similar. Their geographical distri-

butions are knowm to overlap only between Mo^ambi(jue

and Kenva, which might suggest an allopatric mode of

speciation followed b\ limited secondarv contact. These

two have strikingly dissimilar penial shapes, as would be

predicted if this were either an isolating or species-rec-

ognition mechanism (Reid, 1996). A second likely sister-

species pair is P. isscli and P. mauritiana. Again these

are similar in shell characters, but differ in penial shape.

These are aliuost completel)' allopatnc; P isscli is re-

stricted to the Red Sea, while P. mauritiana occurs wide-

ly in the Indian Ocean and Persian Gulf, and thev over-

lap only in the vicinih' of Aden and at the mouth of the

Red Sea. While endemism in the Red Sea fauna is high

in some groups, among mollusks it is generally low

(Sheppard et al, 1992). During the substantial falls in

sea level produced bv Pleistocene glaciation, the Red

Sea became hvpersaline, and recolonization from the

Arabian Sea has occurred only since the Holocene trans-

gression. Speciation in the Red Sea mav therefore be a

recent phenomenon. At present, the currents at the

mouth of the Red Sea do not appear to prevent passage

of plankionic lanae, but the cold upweUing on the

southern Arabian coast during the summer months may

cause a partial isolation of the Red Sea from the Arabian

Sea and Indian Ocean (Sheppard rf al., 1992). The third

likelv pair is P hahci and P. fasciata. vs'hich both belong

m the infracostata group'. These are known to overlap

only between Hong Kong and Taiwan, and show a shght

but consistent difference in penial shape. Finally, P. co-

noidalis mvX P tantilla mav be sister-species. Their rad-

ula, o\iduct and penis are identical, shells are similar in

sculpture, and both occur only on oceanic islands (Reid,

1989a). The former is widespread throughout the Indo-

West Pacific, whereas the latter is restricted to the Ha-

w;dian Islands, suggesting a case of speciation by pe-

ripheral isolation. The species status of P. tantilla is

based only on its distinct shell. The two are not biown

to occur svmpatricallv and, assuming that penial shape

is an isolating mechanism, there has presumablv been

no selection for its divergence.

ACKNOWLEDGMENTS

This work was carried out while Y-MM was in receipt

of a post-doctoral fellowship from the Croucher Foun-

dation, Hong Kong. We thank E. Glover and J. D. Tavlor

(BMNH), S-Y. Chan (Singapore), H.L. Strack (Dor-

drecht, The Netherlands), Y. Shikancj, (Tokx'o) and E.

Platts (Winchester, U.K.) for providing specimens. For

assisting Y-MM with field collecting we thank K. S. Tan

(National University of Singapore), Fu-xue Li (Xiamen

University), C. K. Tseng (Institute of Oceanography,

Qingdao), Huang Shong (National Taiwan Normal Uni-

versity), Michael Ilin-kiu Mok (National Sun Yat-sen

University, Taiwan), S. Yamato and S. Ohgald (Kyoto

University, Shinihama), Y. Takada (Ishigaki Tropical Sta-

tion) and all their colleagues. For providing loans of ma-

terial we are grateful to P. Bouchet (MNHN), H. Dekker

(The Netherlands), H. Fukiida (Tok-vo Metropofitan

University), R. N. Kilburn (NM), I. Loch (AMS), R.

Moolenbeek (ZMA) and ). Pickering (OUM). For their

expert photography of shells we thank H. Tavlor and N.

Hayes (BMNH). P. York (BMNH) kindly assisted with

the photomicroscopy of egg capsules. We appreciate the

helpful comments of two anon)mious referees.

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THE NAUTILUS 112(l):34-4(), 1998

Page 34

Donald Richard Moore (February 16, 1921-October 19, 1997):

Biographical Sketch, Zoological Taxa, and Bibliography

Jose H. Leal

The Bailev-Matthews Sliell Museiini

3075 Sanibel-Capti\u Road

Sanibel, FL 33957 USA

[email protected]

Donald R. Moore was born in West Palm Beach, Flor-

ida, son of Walter R. and Estelle K. Moore. He attended

elenientar\- school in Venice, C^alifornia and Hobe

Sound, Florida, and high school in Stuart, Florida.

Growing up in Florida provided Don with, in his words,

"plentv of sea, swamp, lagoon, and woods to roam in".

He joined the U.S. Marine Coq^s in September 1941.

Don's love for shells and marine life consoUdated after

his stay in the Central and Southwest Pacific, something

that happened with so many young naturahsts who

fought the Second World War in that part of the world.

Don took his dive mask witli him to the Pacific, and

dived as much as events would permit, at Midway, New

Hebrides, and the Solomon Islands.

After receiving an honorabk- discharge in October

1945, Don entered the undergraduate program at the

Universitv' of Miami in June 194S, and graduated in Feb-

marv 1954 with a Bachelor of Science degree in Zoology.

He joined the Graduate School of the Universit\' of Mi-

ami shortly thereafter, aiming for a M.S. degree in Ma-

rine Science. In the summer of 1954 Don traveled to

Houston, Texas, taking on a temporaiy assignment with

the Exploration and Product Research Dixdsion of Shell

Oil Company as field biologist. A job offer as Research

Scientist at the Institute of Marine Science, University

of Texas, caused Don to put his academic efforts tem-

porarily on hold. From Texas he moved to Ocean

Springs, Mississippi, to join the staff at the Gulf Coast

Research Laboratory, working on oyster culture with

Gordon Gimter as an Assistant Marine Biologist It was

there that he published his first scientific article (Moore,

1956). Taking advantage of his proximity to the Univer-

sitv of Southern Mississippi camp\is in Hattiesburg, he

enrolled in the graduate program at that institution with

Gordon Counter as his advisor, earning the Master of Sci-

ence degree in June 19fi().

In September of 1960, he again entered the Graduate

School at the Institute of Marine Science (now the Ro-

.seustiel School of Marine and Atmospheric Science,

RSMAS), University of Miami, graduating under the

guidance of Clilbi'rt L. Voss on June 7, 1964, with a

Ph.D. degree in Marine Science. His Ph.D. dissertation

revised the systematics of western Atlantic \itrincllid

gastropods. Don once told me of his first meeting with

the great Henry Pilsbry. In the early stages of his dis-

.sertation work, Don visited Pilsbrs at the Acadeins of

Natural Sciences in Philadelphia. In Pilsbry's office, he

brieflv introduced himself and his research goals. After

hearing Don's desire to revise the western Atlantic Vi-

trinellidae, Pilsbrv' offered his hand, saving, "Here, I sa-

lute a hero! ". His unpubli.shed dissertation remains the

best source of information about this difficult faniilv of

gastropods.

From the historic perspective of the entl of the twen-

tieth centur)', the timing of Don's Ph.D. defense could

not have been better: the "golden age" of tropical west-

ern Atlantic marine science, centered at the lab in Mi-

ami was under full sail in the mid-196()s. Don was hired

in 1964 as an Assistant Professor of Marine Science at

that institution. There he shared an extremely produc-

tive academic environment with marine biologists such

as (Gilbert Voss, Ravmond Manning, Frederick Bayer, C.

Richard Robins, and Hillary B. Moore, as weD as with

geologists of the caliber of Robert Ginsburg and Cesare

Emiliani. During the sixties, Don cniised extensively on

the Institute's research vessels in the Caribbean Basin,

northern South America, Brazil, Straits of Florida and

the B;iliamas, and the eastern seaboard of the U.S. In

1971 Don was promoted to Associate Professor of Ma-

rine Geology, remiiining in that position until he retired

to become Professor Emeritus in 1984. Don continued

working and participating in students' committees until

the dav of his death.

Don showed great appreciation of other cultures anil

was proud of his travels and abilitv to read and ,speak

several foreign languages. Among his many travels and

professional visiting appointments, he v\'as a visiting sci-

entist at the Institnto de Investigaciones de Punta Betin

in Santa Marta and taught for Sea Grant in Cartagena,

both in Colombia. In 1980, Don worked at the Natural

Histon Museum in Basel, Switzerland, by invitation of

the Swiss National Science Foundation. An active par-

ticipant in the American Miilacological Union, he sen'ed

as Conniillor-at-Large of that organization in 1968-69,

Vice-President in 1973-74, and Presulent in 1974-75.

Following an invitation by then-editor R. Tucker Abbott,

l^on joined the first group of consulting editors of The

Naiifilit.s in julv 1972, a position he retained for the rest

of his life (l')(in's first three peer-reviewed articles were

piiblislicd m rlir Saiitilus). He was also a member and

strong supporter ol the Miami Geological Society, serv-

in<' as tluir Vice-President in 1967, President in 1968-

J. H. Leal, 1998

Page 35

Young Donald R. Moore in 1954, suneving Gull ot Mexico

niolliisks on the coast ol Texas for Shell Oil Co., as featined

on the cover of Shell News Magiizine {Anonviiious, 1955).

69, and Secretai-v in 1977-78. He wa,s a .strong .snpporter

of Unita.s Malacologica a.s a global organization, and

played a key role, as a Scientific Committee member, in

the formation of the Committee for Latin American

Congresse.s in 1990.

Don Moore sened on more than one hundred Uni-

versit)' of Miami graduate .student committees, niiiinly at

RSMAS, but also at the University of Miami's Depart-

ment of Biolog) and Department of Geological Sciences,

on the Cora! Gables Campus. He was committee chair

or co-chair for the students: Richard S. Houbrick (M.S.),

Jose H. Leal (Ph.D.), John Meeder (Ph.D.), James

Quinn, Jr. (Ph.D.), Nidia Homer (M.S.). Don iilso was a

committee member for the following malacologists:

George Darcv (M.S.), Thomas Brackoniecld (M.S. and

Ph.D.), Charles DAsaro (Ph.D.), Steven He.ss (Ph.D.),

Edward J. Petuch (M.S. and Ph.D.), James Quinn, Jr.

(M.S.), Ronald Toll (Ph.D.).

Both as mv personal friend and academic advisor, Don

was always present and supportive, in good and bad

times. A large portion of my personal librarv' comprises

used books and reprints that were gifts from Don (and

I know the same to be tnie for many of his students)

An a\id reader and staunch book"\vorm, Don used to

spend a great deal ol his time in bookstores and book

k ■ ■'• ' -J

Dr Donald R. Moore m 1989, Coral Cables, Flonda Photo

courtesN RSMAS Archi\'es.

lairs. He also used to gauge the merit ot small towns by

the presence or absence of good second-hand book-

stores. Although most of his scientific producti\itv took

place in the 1960s and 1970s, Don was allowed to retain

for hfe his office in the North Gro.sxenor Building at

RSMAS. His office doors were always open, a s\nnbolic

depiction of his view of the world and of the special

affection and warmth he had for students, no matter

how inexperienced, or how distant their homelands.

Don's immediate family survive him: his wife C\iithia

is Researcli Associate with the Division of Marine and

Atmospheric Chemistry- at RSMAS, and son Walter is

Systems Administrator with Eckerd College at St. Pe-

tersburg, Florida

ZOOLO(;iCAL TAXA

Abbreviations im repositories of Wpe material:

AMNH American Museum of Natural Historv, New

York

ANSP The .Academv of Natural Sciences, Philadel-

phia

BMNH The Natural History Museum, former British

Musemn (Natural Histoi-y), London, En-

gland

CNMS C^olombian National Museum of Science, Bo-

gota, Cokjmbia

DMNH Delaware Museum of Natural Histon, Wil-

mington

FMNH Field Museum of Natural Historv, Chicago

FSBC Florida Marine Research Laboratorv, St. Pe-

tersburg

Page 36

THE NAUTILUS, Vol. 112, No. 1

MNHN Museum national d'Histoire naturelle, Paris,

France

MNRJ Museu Nacioniil, Rio de Janeiro, Brazil (also

MNHSL)

MCZ Museum of Comparative Zoolog\\ Harvard

University', Cambridge

MORG Museu Oceanografico, Funda^ao Universida-

de do Rio Grande, Rio Grande, Brazil

NNM Nationaal Natuurhistorisch Museum, Leiden,

The Netherlands (formerly Rijksmuseum

van Natuurlijke)

UF Florida Museum of Natural Historv, Univer-

sity of Florida, G;iines\ille

UMML Marine Invertebrate Museum, Rosenstiel

School of Marine and Atmospheric Science,

Uni\ersit\ ol Miami, Miami

USNM National Museum of Natural History, Smith-

sonian Institution, Washington, DC

Family-Level Taxon

CYCLOSTREMELLIDAE Moore, 1966, Bulletin of Marine

Science, 16(.3):4S0— 4S4 (diagnosis on page 4S1). For Cij-

clostremella Bush, 1897. Robertson (1973) indicated that

this genus belongs to the PyTamidellidae. Ponder & Waren

(198S) included the Cvclostremellinae as as a subfamily o)

P\TaniidelIidae.

Genus-Level Taxon

Dimijella Moore, 1969, Journal de Conchvlioloepe 107(4):137-

141 (diagnosis on page 137). T\pe-species: Dinu/rlhi star-

cki Moore, 1969, by nionotvpy. Dinividae.

Species-level Taxa

All type-specimens collected by D. R. Moore except

where otherwise indicated.

bulli.si, Linckia Moore, I960, Bulletin of Marine Science

of the Gulf and Caribbean I0(4):414-4I6, fig. I. Holo-

type: USNM E-8I20, off Guiana, South America,

8"15'N, 58"I7'W, 72 to 91 m. (=Lmckia nodosa Perrier,

1S7.5, fide Downey, 1968) (Echinodermata, Asteroidea,

Ophichasteridae ) .

calcicola, Ntictda Moore, 1977, The Nautilus 91 (4): 120-

121, figures 1-3. Holotype: USNM 7.585.3.5, Chancanab

[Chankanaab] Lagoon, Cozumel, Quiutana Roo, Mexico,

2 m; All paratypes from type locafity: USNM 758537, 7

paired specimens -I- 3 single valves; MCZ 288445, 3

paired specimens + 4 single valves, from tvpe localit\';

UMML 28.2812, 3 p;ured specimens + 4 single valves,

from type locality; AMNH 183857, 10 single valves;

ANSP 344387, 5 paired specimens + 4 single valves;

MNHN unnumbered, 5 paired specimens. 2 left valves,

1 right valve; DMNH 120581, 6 paired specimens + 1

single valve; BMNH 1977089, 3 paired specimens + 3

single valves; NNM 55302, 4 paired specimens + 4 sin-

gle valves, all dry. Nucididae.

colombiana, Alvania (Alvania) Romer and Moore, 1988,

The Nautilus 102(4): 131-133, figures 1-5. Holotvpe

USNM 859339, off west Florida, 29°35'N, 87°20'06'"W.

107.3 m; Paratvpes: USNM 859340, 1 parat\pe, t\pe lo-

cality; FSBC 3311.3-114, 2 paratvpes, off west Florida,

28°24'N, 8.5°15'06"W. 164.6 m; MCZ 297220, 1 para-

type, off Puerto Rico, 17°53'24"N, 66°35'10"\\' 221 m;

MCZ 297219 and UMML 30.8349, 2 parat\pes, off Mi-

ami, Florida, 25°47'N, 80°01'30"W, 137 'm; UMML

.30.8350 and CNMS ICN-MHN(MO) .523, 2 paratvpes,

off Miami, Florida, 2.5°46'.30"N, 80°00'08"W, 76.81 m;

CNMS ICN-MHN(MO) 522, 1 parat%pe, off Caribbean

coast of Colombia, 09°52'.3.5"N, 75°47'25"W, 72 m. Ris-

condijlum, Caecum Moore, 1969. Bulletin ol Marine Sci-

ence 83(l):26-28, figure 1. Holotype: USNM 679.348,

Pavardi Island, near Colon, Atlantic coast of Panama.

Parat\pes: USNM 679348, 1 paratvpe. East Flower Gar-

den Bank, Gulf of Mexico, 27°54'30"N, 93°35'45"W. cor-

al debris, 1 8.. 3-23. 8 m. Caecidae.

cynthiac, Condylonucida Moore, 1977, The Nautilus

91(4): 12.3-124, figures 6-7. Holotype: USNM 7.58.534,

about 800 m west of outer reet (lagoon), Courtown Cays

(Cayos del E.S.E.), off Nicaragua, western Caribbean

Sea, 7.5 m. Paratvpes: USNM 7.58.538, 5 paired speci-

mens + 1 broken right valve + 1 fragment, Courtown

Cavs, off Nicaragua, 1.5 m; ANSP 344388, I p;iired spec-

imen + 1 right valve, Courtown Cays, off Nicaragua, 1.5

m; DMNH 120580, 1 paired specimen + 1 left vidve;

MCZ 288444, 1 piiired specimen + 2 left valves, Cour-

town Cavs, north end, 2 m; UMML 28.2811, 1 paired

specimen + 1 left v;ilve, Courtown Cavs, outer reef, 1

m, Nuculidae.

rsperanza, Thala Leal and Moore, 1993, The Nautilus

107(2):58-62, figures 1-6. Holotype: USNM 860280,

Playa Esperanza, 40 km west of San Juan. Municip;Uity

of Manati, northern Puerto Rico .\11 parahpes from tvpe

localit%-: USNM 860281, parat^pes 1-4; UF 193.382-.i83,

paratvpes .5-6; ANSP 391938, paratype 7; AMNH

232313, paratvpe 8; MCZ .302588, paratvpe 9; MNHN

unnumbered, paratype 10; FMNH 2233S8, paratype 11;

UMML 30.8375, paratype 12; USNM 860262, paratypes

1.3-16. Costellariidae.

fioridana, Macromphalina Moore, 1965, The Nautilus

'78(3):7.5-76, plate 7, figure 1-3. Holotype: USNM

6.36610, Madeira Beach at 150"' Avenue, Saint Peters-

burg, Floritla; paratvpes: ANSP 295621, 1 shell, type lo-

cality; UMML 302773, 1 shell, east of Sokfier Key Bis-

cavne Bay, Florida, I m. Vitrinellidae.

iiidiscrcta. Rissoiua Leal and Moore, 1989, Bulletin of

Marine Science 45(I):I39-147, figures 1-12. Holotype:

MNRJ 5760. Ponta Verde Beach, Maceio, Alagoas State,

J. H. Leal, 1998

Page 37

Brazil, 09°38'S, 35°44'W, beach at low tide. Paratxpes:

MNHSL 6025, paratopes 1-4, t\pe Iocalit\-; MORG

4164, paratvpe.s 5-7, t\pe locality; MNHN unnumbered,

paratopes 8-11, t\pe localit\'; USNM 859335-338. para-

t\pes 12-15, t\pe loc;iliK; MNRJ 5761, paratxpes 16-18,

type localitv-: BMNH l'9S8.()43, paratvpes 19-22, type

localit\-; MORG 25463, parat\pe.s 23-24, Atol das Rocas,

Brazil', ()3°52'S, 33°4y'W; MORG 25464, parat\pe,s 25-

28, Abrolhos Reef Complex, B;ihia, Brazil, ■l7°58'S,

38°42'W; UMML 8347-48, parat\pes 29-30, Jaragua

Beach, Maceio, Alagoas State, Brazil, 09°38'S, 35°44'W.

Rissoidae.

insiilaniin. Caecum Moore, 1970, Bulletin of Marine

Science 20(2):368-373, figures lA-B, Holot>pe: USNM

679178, open sandy bottom at head of Lesser Lameshur

Bav, St. John, Virgin Lslands, 1.5 m. All paratypes from

type localitv: USNM 679179, 12 paratypes; ANSP

315477. 10 ■ parat\pes; MCZ 271.305. 11 paratxpes;

AMNH 147965, 10 paratypes; UMML 30.3203, 14 para-

types; "more than a hundred paratypes ... kept bv the

author". Caecidae.

mai/a. Condi/lonitcula Moore, 1977, The Nautilus 91(4):

124-126. figures 4-5. Holot\pe: USNM 758536, Chan-

canab [Chankanaab] Lagoon, Cozumel, Quintana Roo,

Mexico. 2m. All paratypes from type locaht)': USNM

758536, 7 paired specimens + 1 right valye, type local-

ity; MCZ 288443, 2 p;ured specimens -I- 1 right yalye;

ANSP 344389, 3 paired specimens + 1 left yalye;

AMNH 183858, 2 paired specimens -I- 1 left yalye;

DMNH 120579, 3 paired specimens + 2 single valves;

FMNH 198080, 1 paired specimen. Nuculidae.

semipuncttis, Solariorhis Moore, 1965, The Nautilus

7S(3):77-78, figures 1-3. Holot>pe: USNM 636309,

northwest Campeche Bank, Mexico, 18 m, muddy bot-

tom. Paratype: UMML 30.2774, 1 paratype, Baie de

Aquin, Haiti, beach drift. Vitrinellidae.

starcki, DumjicUa Moore, 1969, Journal de Conchvliol-

ogie 107(4): 138-139, one plate with two figures. Holo-

tvpe: USNM 678363. complete shell with txvo larxal

specimens in brood chamber, Cozumel Island, Mexico.

All paratypes from type locality: USNM 678364, 1 paired

specimen + 1 detached upper valve; ANSP 312218, 1

paired specimen -I- detached upper valve; BMNH

1967.6171, 1 paired specimen + another with broken

lower valve; MNHN 1967.226, 1 paired specimen -I- de-

tached upper valve; MCZ 257727-257726, 1 paired

specimen with brood chamber in upper valve + de-

tached upper \alve; UMML 28.784, 1 paired specimen

+ another with both v;ilves broken + three small de-

tached upper valves. Dimvidae.

tcxana. Vitrinclla Moore, 1965, The Nautilus 78(3);76-

77, plate 7, figures 4-6. Holotype: USNM 636311, Mus-

tang Island, near Port Aransas, Texas. Parat\pes: ANSP

295622, 1 paratope, t>pe locality; USNM 636312, 1 para-

type, type localitv'; UT 1015, t\vo shells, Cline's Point,

Port Aransas, Texas; UT 1016, 1 paratvpe, Ferr\' Land-

ing, Port Aransas, Te.xas; UMML 30.2775, 1 paratype.

Port Aransas. Texas; UMML .30.2776. 1 paratvpe. Port

Aransas, Texas. VitrinelUdae.

GRADUATE THESES

Moore, D. R 1960. Tlie marine and brackish water MoIIusca

of the Mississippi Gulf Coast. ,Master of Science. Univer-

sity of Sontheni Mississippi. 116 pages.

Moore, D. R, 1964. The faniilv N'itnnellidae in South Flonda

ant! the Gulf ot Mexico. Doctor ot Philosophy, Uni\'ersity

of Miami, xi -I- 235 pages, 35 figures, -I- vitae. [7 June]

PUBLICATIONS

This hsting includes peer-re\iewed articles as well as

Don's reports and contributions to several scientific or-

ganizations. Also included are the extended meeting ab-

stracts pubUshed up to 1980 bv the American Malaco-

logical Union (AMU) in its Annual Reports and Bulletin.

References are hsted in chronological order.

Moore. D, R 1956. Observations of predation on echinoderms

h\ three species of Cassididae. The Nautilus 69(3);7.3-76.

[il Fehniarv]

Moore, D. R. 1957. A note on Cuna daUi. The Nautilus 70(4):

12;3-125, pi. 8, figs. .5-4. [29 April]

Moore, D. R, 1958. Additions to Texas marine MoIIusca. The

Nautilus 71(4):124-129. [24 April]

Moore, D. R. 1959. Notes on Blanquilla Reef the most north-

erlv coral reef in the western Gulf of Mexico. Publications

of the Institute of Manne Science 5:151-1.55 ["Decem-

ber. 195S-]

Moore, D. R. and H. R. Bullis. 1960. A deep-water coral reef

in the Gulf of Mexico. Bulletin of Marine Science of the

Gulf and Caribbean 10(1): 125-128. figures 1-2. [March]

Moore. D. R. 1960. Linckia hnlhsi, a new asteroid from the

northeast coast of South America. Bulletin of Marine Sci-

ence of the Gulf and Caribbean 10(4):414-416, fig 1.

[December]

Moore, D R 1961a. The manne and brackish water mollusca

of the state of Mississippi. Gulf Research Reports [Ocean

Springs, Mississippi] 1(1): 1-58. [April]

Thomas, LP., D. R. Moore and R. C. Work. 1961. Effects of

hurricane Doniia on the turtle grass beds of Biscayne Bay,

Florida. Bulletin of Marine Science of the Gulf and Ca-

nbbean 11(2): 191-197, figs. 1-2 [June]

Moore, D. R. 1961b. The occurrence of Stephanoscijphuscor-

nifonnis Komai (Sc\phozoa) in the western Atlantic. Bul-

letin of Marine Science of the Gulf and Caribbean 11(2):

319-.320. [June]

Moore, D. R. 1961c. The molluscan sea grass communities in

Biscayne Bav. American Malacological Union Annual Re-

ports for 1961 [Bulletin 28], page 20, The American Mal-

acological Union Twenty-eighth Annual Meeting, Wash-

ington, DC, June 19-23, 1961. [Abstract] [1 December]

Moore, D. R. 1962a. Notes on the distribution of the spinv

lobster Fanuhnis in Florida and the Gulf of Mexico. Crus-

taceana 3(4):3L8-319. [March]

Moore, D. R and G Gunter 1962. Notes on the Pliocene

molluscan fauna from one site in the western Florida Ev-

erglades. Bulletin of Marine Science ot the Gulf and Ca-

ribbean 12(l):66-72. [March].

Olsson, A. A. and D. R. Moore. 1962. A neglected west At-

Page 38

THE NAUTILUS, Vol. 112. No. 1

lantic stronib. The Nautilus 75(4):127-128, pi. 11, figs. 1-

3. [9 April).

Moore, D. R. 1962b. Occurreuce and distribution o( Netnopsis

hachei Agassiz (Hvdrozoa) in the nortlieni Gulf of Mexico.

Bulletin of Marine Science of the Gulf and Caribbean

12(3):399-402. [September].

Moore, D. R. 1962c. The systematic position of the famiK-

Caecidae. American Malacological Union Annual Report

for 1962 [Bulletin 29], page 5, Twenty-ninth Annual Meet-

ing of the American Malacological Union, St. Petersburg,

Florida, 31 ]uly-.3-August 1962. [Abstract] [1 December"].

Moore, D. R. 1962d. The systematic position of the family

Caecidae (MoUusca: Gastropoda). Bulletin of Marine Sci-

ence of the Gulf and Caribbean 12(4):69.5-7()1, figs. 1-3.

[December]

Moore, D. R. 1962e. An ophiurian shoal on the Mississippi

coast. Quarterly Journal of the Florida Academy of Sci-

ences 25(l):70-72. [23 August]

Moore, D. R. 1963a. Turtle grass in the deep sea. Science

1.39(.3.560): 12.34-12.3.5. [22 March]

Moore, D. R, 1963b. Mecoliotia. a gastropod genus new to

Florida. Bulletin of Marine Science of the Gulf and Ca-

ribbean 1.3(1): 7:3-76, fig. 1. [March]

Moore, D. R. 1963c. Distribution of the sea grass, Thalassia,

in the United States. Bulletin of Marine Science of the

Gulf and Caribbean 13(2):.329-.342. [June]

Moore, D. R. 1964. The evolution of the Mesogastropoda,

American Malacological Union Annual Reports for 1964

[Bulletin 31], pages 17-18, Thirtieth Annual Meeting of

the American Malacological Union, New Orleans, Loui-

siana, 21-24 July 1964. [Abstract]

Hoffmeister, J. E., J, 1. Jones, J. D. Milliman, D. R. Moore

and H. G. Multer. 1964. Field guidebook to the hving

and fossil reef types of South Florida Miami Geological

Society, Coral Gables, 2S pages, [Noxember]

Moore. D, R. 1965a. New species of Vitrinellidae from Gulf

of Mexico and adjacent waters. The Nautilus 78(3):7.3-7S,

plate 7, figs. 1-6; plate 8, figs. 1-3. [25 Januar\']

Moore, D. R. 1965b. The Marquis de Folin and his work on

the Caecidae. American Malacological Union Annual Re-

ports for 196.5 [Bulletin .32], pages .3.5-36, Thiity-first

Meeting of the American Malacological Union, Staten Is-

land. New York, 20-23 JuK. 1965.'[Abstract| [1 Decem-

ber]

Moore, D. R 1966a. A note on Pseudoparastrophia. Journal

de Conciiyliologie 105(2):57-61, figs. 1-2. [1 Marclil

Moore, D. R. and K. J. Boss. 1966. Records for I'liralxu-nid

squiUimi. The Nautilus 80(1 ):.34-.35. [6 Julyj

Moore. D. R. 1966b. The Cyclostremellidae, a new lamiK ot

prosobraiicli mollusks. Bulletin of Marine Science 16(3):

480-484, figs. 1-6. [September]

Moore, D. R 1966c. Systematics and zoogeography of the

Ctiloceratidae. American Malacological Union Annual

Report for 1966 [Bulletin 33], page 40, Thirty-.second

Annual Meeting of the American Malacological Union.

Chapel Hill, North Carolina, 22-27 August 1966. [1

December]

Boss, K. J. and D. R. Moore. 1967. Notes on Malleus {Pari-

rruillexis) amdeanus (d'Orbignv) (Mollusca, Bi\'aKia). Bul-

letin of Marine Science 17(1 1:8.5-94, 2 figs. [Marcii]

Bock, W. D. and D. R. Moore. 1968. A commensal relation-

ship between a foraininifer and a mollusk. Gulf Research

Reports [Ocean Springs, Missi,ssippi] 2(3):27.3-279, plate

1. [No\ember|

Moore, D. R. 1968. Obser\'ations on the Caecidae. American

Malacological Union Annual Report for 1968 [Bulletin 35,

page 39, Thirty-fourth Annual Meeting of the American

Malacological Union, Corpus Christi, Te.xas, 15-19 July

1968. [Abstract] [27 December]

Moore, D R 1969a. A new Caecum from the tropical western

Adantic. Bulletin of Marine Science 83(1):26-2S, fig. 1.

[July]

Bock, W. D., D. R. Moore, A. C. Newmann and P. R Supko.

(organizers). 1969. Late Pleistocene geology in an urban

area. Third annual field trip of the Miami Geological So-

ciety. Miami Geological Societ\'. Miami, 26 pages. [July]

Moore, D. R. and P R. Supko. 1969. Introduction. In: Bock.

W. D., D. R. Moore, A. C. Newmann and P R. Supko.

(organizers). Late Pleistocene geolog\' in an urban area,

pages 1-2. Third annual field trip of the Miami Geological

Society. Miami Geological Society, Miami. [Jul\']

Moore, D. R. 1969b. Cijclostrema miranda Bartsch, a syn-

onym of Tormis subcarinatus (Montagu). The Vehger

12(2): 169-170. figs. 1-3. [1 October]

Moore, D R. 1969c. Systematics, distribution, and abundance

of the West Indian micromollusk Rissoina catesbijana

d'Orbigny. Nineteenth Meeting of the Gulf Coast Asso-

ciation of Geological Societies, October 29-November 1,

1969. Transactions [of the] Gulf Coast Association of Geo-

logical Societies [Tallahassee, Florida] 19: 42.5-426. [Oc-

tober]

Moore, D. R. 1969d. A new genus and species of Dinnidae

from the Caribbean coast of Mexico, Journal de Concliv-

liologie 107(4):137-141, 1 plate. [30 December]

Moore, D. R. 1970a. The subgenus Armata de Folin, of the

genus Caecum. Journal de Conchvliologie 108( 1 ):.3-.5. [31

March]

Moore, D, R. 1970b. A new Caecum from Puerto Rico and

the Virgin Islaiuls. Bulletin ot Marine Science 20(2):.36S-

.373, figs, 1-2, [June]

Moore, D. R, 1971a. Cochliolcpis parasitica, a non-parasitic

marine gastropod. American Malacological Union Annual

Reports for 1970 [Bulletin 37], page 80, Thirty-sixth An-

nual Meeting of the American Malacological Union, Key

West, Florida, 16-20 July 1970. [18 Febniarv]

Moore. D. R. 1971b. A deep water Omalofii/ra in the western

Atlantic, The Nautilus 84(4): 11.3-1 17, fig, 1. [26 .\pril]

Bock, W, D, and D R. Moore. 1971. The Foraminifera and

micromollusks of Ilogstv Reef and Serrana Bank and their

paleoecological significance. Transactions of the Fifth Ca-

ribbean Geological Conference, Geological Bulletin

[(Queens College Press, Flushing, New York] No. 5, pages

14.3-146, 2 figs., 2 tables. [May]

Meeder, J. F, and D R Moore 1971. The extension of

range of Beiilulinia carih}>ea Edmunds to Brazil and

Panama (Molliisa [sic]; Gastropoda). Caribbean Journal

of Science 11(.3-1):1.59-I61, figs. A-B. [September-De-

cember].

Moore. D R. 1971c. Disccners' of the Carolina marsh clam,

Poli/mesoda caroliniana (Rose), a supposeil Florida dis-

junct species, in Everglades National Park. Florida. Gulf

Research Reports [(.)cean Springs, Mississippi! 3(2): 26,5-

277, figs. 1-.3. table 1. [December]

Moore. D. R. 1972a. Ecological and .SN.stematic notes on Cae-

cidae collected at St. Croix, Virgin Islands. Bulletin of the

American Malacological Union for 1971, page 11, Thirt\-

se\'enth Annual Meeting of the American Malacological

Union, Cocoa Be;ich. Florida, 15-19 |ul\ 1971 [Febman]

J. H. Leal, 1998

Pase 39

Moore. D. R. 1972b. Cochliolepis parasitica, a nonparasitic

marine gastropod and its place in the V'itrinellidae. Bul-

letin of Marine Science 2iZ(l): 100-1 12. figs. 1-6. [March]

Moore. D. R. 1972c. Coral reefs, new and old. Pages 1-6,

Field trip. 2-3 December 1972, American Quateniarv' As-

sociation, [2 December]

Moore, D. R 1972d. Ecological and s\stematic notes on Cae-

cidae from St. Croix, U.S. Virgin Islands. Bulletin ot Ma-

rine Science 22(4):881-.899, figs. 1-12. [with abstract in

Spanish] [December].

Moore, D. R. 1973. Mollusks from a small landlocked Mexican

lagoon. Bulletin of the American Malacological Union tor

19V2. pages 5-6. Thiit\ -eighth Annual Meeting of the

American Malacological Union, GaKeston, Texas, 9-14

July 1972. [Abstract] [March].

Moore, D. R. 1974. Remarks on the molluscan tauna ot north-

em South America. Bulletin of the American Malacolog-

ical Union for 197.3, page 17, Thirt\'-ninth .\nnual Meeting

of the American Malacological Union, Newark an Green-

ville, Delaware, 24-28 June 1973. [Abstract] [22 Mav]

Moore. D. R. 1975. Philobr^idae in the Northern Hemi-

sphere. Bulletin of the American Malacological Union for

1974. pages .34— .35. Fortieth Annual Meeting ot the Amer-

ican Malacological Union. Spnngfield, Massachusetts, 4-

8 August 1974. [Abstract] [May]

Moore, D. R. 1976a. Is Meioceras living in the Indo-Pacif-

ic? (Gastropoda: Caecidae). Bulletin ot the American

Malacological Union for 1975, pages 19-20, Joint Meet-

ing of the Western Society of .Malacologists and Amer-

ican Malacological Union [Fort\-first .\nnual .Meeting].

San Diego, California. 22-26 June 1975. [Abstract] [30

Januar)]

Moore, D. R. 1976b. .Vlicromolluscs ot the continental shell,

northeastern Gulf of Mexico. Bulletin of the American

Malacological Union for 1976, page 49, ForU-second An-

nual Meeting of the American Malacological Union, Co-

lumbus, Ohio, 2-6 August 1976. [Abstract] 130 Decem-

ber]

Moore, D. R. 1977. Small species of Nuculidae (Bi\al\ia) trom

the tropical western Atlantic. The Nautilus 91{4);119-128.

figs. 1-7. J27 October].

Moore, D. R 1978a. The western Atlantic Dim\idae. Bulletin

of the American .Malacolf)gical Union tor 1976, pages 41-

43, Forts-third .\nnual Meeting of the American Mala-

cological Union, Naples, Flonda, 10-15 July 1977, ]Ab-

stract] [April]

Moore, D. R. 1978b. From the first cniise of the 'Blake" to

the present day-one hundred years of progress in the

study of Gulf and Caribbean marine mollusks. Bulletin of

the American Malacological Union, Inc., tor 1977, pages

63-68, Fortrv-third Annual Meeting of the American Mal-

acological Union, Naples, Flonda, l0-15 July 1977. jApril]

Moore, d" R. 197Sc. A.xel A, Olsson— An obituary The Nau-

tilus 92(2);.59-6]. 1 portrait. [27 April]

Moore, D. R. 1978d. The Caecidae of Brazil. Bulletin of the

American Malacological Union for 1978, page 56, Forty-

fourth Annual Meeting of the American Malacological

Union, Wilmington, North Carohna, 16-21 July 1978,

[Abstract]

Moore, D, R, 1979. On proper citations. The Mollusk JGreater

Miami Shell Club. Inc.] 17(2.):4, [2 Febnian ]

Moore. D. R, and M, F Miller 1979. Discovery ot living bi-

valved gastropods in the Florida Keys. The Nautilus 93(2-

3): 106. [23 April]

Moore. D. R 1980a. Molluscan exploration and research in

and around the Gulf of Mexico. Bulletin of the Amer-

ican Malacological Union, Inc. for 1979, pages 1-5.

Forty-fifth Annual Meeting of the American Malacolog-

ical Union, Corpus Christi, Texas. 5-11 August 1979.

[24 Marcli]

Moore, D R 1980b. Distnbution of some small micromol-

liisks in the northeastern Gulf of Me.xico. Bulletin of the

American .Malacological Union tor 1979, page 64, Forty-

fifth Annual Meeting of the American Nlalacological

Union, Corpus Christi, Texas, 5-11 August 1979. [Ab-

stract] [24 March]

Moore, D. R 1980c. The sliallow water fauna of Sanibel and

its relationship to upper Cenozoic fossils in South Florida.

In: Gleason, P, J, (ed,) Water, oil, and the geology of Col-

lier, Lee, and Hendry counties, pages 57-59, .Miami Geo-

logical Societv', Miami. ]May]

Moore, D, R. 1980d. A long lost (extinct?) mollusc of the Ohio

River. Bulletin of the American Malacological Union for

1980, page 70, Forty-sLxth Annual Meeting of the Amer-

ican Malacological Union, Louisville. Kentiick-v, 19-25

July 1980,

Meeder, J, F,, D. R, Moore and P, Harlem, (organizers).

1981a. Survey of central Florida geology, Miami Geolog-

ical Society 1981 Field Tnp, Tnp! 24-26 April 1981. Mi-

ami Geological Society, Miami, 44 pages. [24 .'\pnl]

Meeder. J. F, D. R. Moore and P Harlem. 1981b. Road

log and narrative. In: Meeder, J. F., D. R. Moore and

P. Harlem, (organizers). Survey of central Florida ge-

ology, pages 6-15, Miami Geological Society 1981 Field

Trip, Trip, 24-26 April 1981, Miami Geological Society,

Miami. [24 April]

Moore, D, R, 1983. The little bivalve Plnnktomija un-

masked. Anales del Instituto de Investigaciones Mari-

nas de Punta de Betin [Santa Marta. Colombia] 13:123-

132, figs. 1-9,

Romer, N. S. and D. R. Moore. 1988. A new species of Al-

lania (Rissoidae) from the West Indian region. The Nau-

tilus 102(4);131-1.33. 5 figs. [21 December]

Leal. J. H, and D, R. Moore, 1989. Ri.ssoina indiscrcta. a new

rissoid species from the tropical southwestern Atlantic

with Indo-Pacific affinities. Bulletin ot Marine Science

45(1): 139-147. figures 1-12, table 1, [March]

Leal, J H. and D, R, Moore, 1993. Thnia esperanza. a new

Costellariidae (Mollusca: Gastropoda) from northern

Puerto Rico. The Nautilus 107(2):58-62. figs. 1-11. ]28

July]

Moore, D, R, 1997. Close encounters with the nurse shark.

In: Feitschrift for C Richard Robins in honor of his re-

tirement trom the University of Miami. Bulletin of Marine

Science 60(3): 101.5-1016, JMay]

ACKNOWLEDGMENTS

I am indebted to Cynthia Moore, Nancv Voss, Kav Hale,

Avis Miller, Helen All^ertson (all from RSMAS, Univer-

sity of Miami), and M.G. Harasewych (USNM) for pro-

viding materials and information about Donald R.

Moore, and/or comments on the manuscript. The fol-

lowing curators/museum specialists are gratefullv ac-

knowledged for assistance with type material: .^dam Bal-

dinger (MCZ), Phillipe Bouchet (MNHN), M.G. Hara-

sevvvch, C\mthia .^hem. Rave Germon (USNM), Paula

Page 40

THE NAUTILUS, Vol. 112, No. 1

Mikkelsen (AMNH), Tim Pearce (DMNH), Gar\- Ro-

senberg (ANSP), Edmund Gittenherger and Jeroen

Goud (NNM), John Tavlor and Joan Pickering (BMNH),

Nancy Voss (RSMASAJMML).

LITERATURE CITED

AnoiiMiious. 1955. Beachcomber. Shell News 2.3(2):4-6. [pub-

lished b\ Shell Oil Coinpanv. New York]

Downey, .VI. E, 196S. A note on the Atlantic species of the

starfish genus Linckia. Proceedings of the Biological So-

ciety' of Washington 81{l):41-44.

Ponder, W. F. and A. Waren. 1988. Classification of the Caen-

ogastropoda and Heterostropha - a list of family-group

names and higher taxa. Malacological Review, Supplement

4:2SSU32S.

Robertson, R. 197.3. Cijclostrcinclla. a planispiral pyramidellid.

The Nautilus 87(,3):88.

THE NAUTILUS 112(1):41, 1998

Pase 41

Notices

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TH Et^NAUTI LUS

CONTENTS

Volume 112, Number 2

September 22, 1998

ISSN 0028-1344

E. Suiirez-Morales

R. Casca

Theeosome pteropod (Gastropoda) assemblages of the

Mexican Carilihean Sea (1991)

43

BiTice A. Marshall

A new deep-sea limpet of the genus Prctiiunloiitd Dall,

18S2 from New Zealand, and new distribution records for

P. aupoiiria antl P Diorioiia Marshall, 1985 (Gastropoda;

Acmaeidae) 52

Rediscoven' oi Discus l)ninsoni Bern', 1955 and Orniliclix

alpina (Eirod, 1901) in the Mission Mountains, Montana,

with comments on Oivohclix clrodi (Pilsbrv, 1900) 58

EpHoiiitiDi fahriziiu (Gastropoda: Epitoniidae), a new

species from Patagonia, Argentina 63

Two new species oi Pciiplomo (Bivalvia; Anomalodesmata:

Periplomatidae) from the southern Caribbean 69

Paul Hendricks

Guido Pastorino

Pablo Penchaszadeh

Nestor A. Ardila

Juan M. Diaz

THE NAUTILUS 112(2):43-51, 1998

Paee 43

Thecosome Pteropod (Gastropoda) Assemblages of the Mexican

Caribbean Sea (1991)

E. Suarez-Morales

El Colegio de la Frontera Sur

(ECOSUR), Uiiidad Chetiinuil,

Apartado Postal 424,

Chetmnal, Quintana Roo 77000,

MEXICO

esuarez®'nicte-lia.ecosur-qroo,nix

R. Gasca

El Colegio de la Frontera Sur

(ECOSUR). Unidad Chetmnal,

Apartado Postal 424,

Chetiniial, Quintana Roo 77000,

MEXICO,

ABSTRACT

The composition, distnbution, and abundance of the theco-

some pteropods collected off the Mexican coast of the Carib-

bean Sea were anal\-zed. Samples were collected during four

months (Febniar\-, March, May, and August) of 1991, Highest

mean thecosome pteropod density was recorded in Ma\ (329

org./IO'm"), while lowest mean densit)' occun'ed in March (71

organism.s/IO'm'). Seventeen species and subspecific tonus

were identified. Creseis acicula aciaila. Limacina troclufonnis.

L inflata, Clio pijramidata pijramidata and Cavolinia longi-

rostris longirostris were the more abundant species, accounting

for 84% of the overall thecosome pteropod counts. The first

two species represent the influence of Caribbean surface wa-

ters in the sur\e\ed area. The local thecosome pteropod fauna

also shows some affinits with that from the southeni Gulf of

Mexico, Cluster analysis oi the stations re\ealed two tvpes of

assemblages of low and high densities, with distributions that

showed month-to-month \ariations. Circadian \anations were

well defined for two of the more abundant species; the other

two species were nearly equally abundant in day and night

samples. The thecosome pteropod communitv' along this oce-

anic area seems to be represented b\' a homogeneous complex

dominated by neritic-oceanic species with a strong influence of

C aciaila acicula even in fully oceanic areas, Intermi.xing of

these assemblages and the co-occurrence of neritic and oceanic

species near the coast could be attributed to the narrow witltli

of the continental shelf and to local circulation patterns.

INTRODUCTION

Thecosome pteropod molluscs are one of the most abun-

tlant holoplaiiktonic groups in neritic and oceanic envi-

ronments. As a group, thev show an adaptive trend to-

ward shell reduction that results in less densih' and in-

creased ability for vertical displacement in the water col-

umn (Van der Spoel, 1996). The Thecosomata is one of

the two holoplaiiktonic orders of Opisthobranchia (the

other one is Gyinnosomata) that were lormerK' united

under the name Pteropoda. Todav we know that these

two orders are not closely related (Lalli and Gilmer,

1989). The term "pteropod" is used herein niidnlv as an

easy reference name based on widespread, common

practice. Thecosome pteropods are mainly herbivorous,

and feed upon microplankton and cUatoms using drifting

mucous webs to capture food (Lalh and Gilmer, 19S9).

Most species inhabit surface waters, but some e.xhibit

large migration patterns from depths of 1000-2000 m.

Some species are useful as hydrological, geological and

ecological indicators (Biekart, 1989).

In the northwestern Atlantic Ocean thecosome pter-

opods have been surveyed mainh' in the Gulf of Mexico

and along the East Coast of the United States (Wor-

melle. 1962; Chen and Hillman. 1970; Matsubara, 1975;

Michel and Michel, 1991; Suarez-Morales and Gasca,

1992). Some studies ha\e been made in the Caribbean

Sea about general aspects of their abundance and their

horizontal and vertical distribution (Wells, 1975, 1976;

Haagensen, 1976; Gasca and Suarez-Morales, 1992). A

general review of the thecosome pteropod records in the

Gulf of Mexico and adjacent areas is presented by Sua-

rez-Morales (1994). However, knowledge ol the com-

position, distribution and abundance of the thecosome

pteropod fauna in the Caribbean Sea off Me.xico, the

westernmost portion of the Caribbean Sea, is practically

limited to the innermost reaches of the coast (Gasca and

Suarez-Morales, 1990). In the present work, the com-

position, distribution and abundance of thecosomes

pteropods collected in the oceanic area off the Mexican

coast of the Caribbean during four oceanographic cruis-

es are anal\zed.

Sti'dv Are.\

The surveyed area hes behveen 18° and 21°30'N and

86°20' and 87°40'W off the eastern coast of the Yucatan

Peninsula, Mexico (Figure 1). The area is under the in-

fluence of the northward flow of Caribbean surface wa-

ter on its way into the Gulf of Me.xico through the Yu-

catan Channel. The continental shelf is narrow along this

section of the coast and depth drops \'ery quickly off-

shore (Merino and Otero, 1991), H\drographic condi-

Pai^e 44

THE NAUTILUS, Vol. 112. No. 2

CAR/BE l-IV

Figure 1. Siineved area with zooplankton sampling stations

off the Mexican coast of the Caribbean .Sea (1991).

tions along the Caribbean coa.st of Me.vito are deter-

mined bv the main northward flow of the Yucatan Cur-

rent and by a southward-flowing coastal countercurrent

(Merino, 1986). The Yucatan Current shows seasonal

variation in strength; its miwimum and mininumi inten-

sities occur respectively during spring and winter (Me-

rino 1992). Three main climatic periods can be distin-

guished: drv (March-|une), rainv (July-October), and

the "nortes" perifxl, with northern domiTiant winds, from

November to Februaiy.

MATERIALS AND METHODS

Four oceanographic cruises were carried out ilurmg

February, March, May, and August 1991, on board dif-

ferent "Dragaminas" (minesweeper) ships (jf the Mexi-

can Secretan'a dc Marina (Februar): D-12\ March: D-

04; May and August: D-OS). The sampling plan mchuled

22 stations (Figure 1).

Zooplankton was coUectetl bv surface ()l)li(|uc hauls

(0-10 m) using a sijuare-mouth (0.45 m side) standard

plankton net (0..3 nun mesh size). A digital flowiueter

was attached to the net Tnoutli to estimate the amount

of water filtered i)\ the net (mean volume: 160 m' per

haul). Zooplankton samples were fi.xed and preserved in

a buffered 4% formalin solution (Smith and Richardson,

1979). Thecosome pti'r()i)ods wi'ri' sorted from the en-

tire samples, identified (Van der Spoel, i9Si), and

counted. Voucher material of the Mexican Caribbean

thecososme pteropods is deposited in the zooplankton

collection of El Colegio de la Frontera Sur (ECOSUR)

under catalog numbers ECO-CH-Z0073 through ECO-

CH-Z0090. Densih,' data were integrated into four data

sets, one for each cruise. Shannon's Index of Di\'ersity

was calculated (in bits/indi\idual, which ex-presses the

average degree of uncertaintv in predicting what species

an individual chosen at random will belong to; Lndwig

and Reynolds, 1988). The Index of Importance Value

(IIV) was used as a dominance measurement. The IIV

accounts for relative (percentual) density and frequency

of the species. In one standard data matrix, IIV would

have a maximum value of 200 ( 1009c oi densitx' plus

100% of frequency) (Brower and Zar, 1977). The IIV

has been used more often by botanists, but the index

can also be used in faunal studies (De la Cruz, 1994).

The Bra\-Curtis Similaritx Index was used to identifv'

station clusters and anaKze the thecosome pteropod

communitA stnicture within indi\idual cruises.

RESULTS

Mean temperature dunng the four-month perioil was

28°C, with a minimum of 23°C in Febmary and maxi-

mum of 29°C in August. Salinitx' averaged .'35.2 %c. with

a maximum of .3.5.6 %c in Mav and a minimum of .34.5

%o in Febniarv.

Space distribution of thecosome pteropod densities in

the surveyed area showed monthly variations (Table 1).

Highest total mean densities were recorded during May

(296.25 organism.s/10'm'), followed by Februaiy (158

org./lO^m'), August (141.79 org./10'm'), and March (71

org./10'm'). During February, thecosome pteropods

showed txvo high-densitv zones, one in the northern por-

tion of the area, and the other in the central zone. No

high-density zones were detected during March. In May,

thecosome pteropods showed high densities along the

entire coast, and in August two groups of stations with

high densities were obseiyed, one in the north and the

other in the central portion. Most high-densitx- stations

were sampled at night (Figure 2).

A total of 17 species and subspecific forms were col-

lected in the suneved area (Table 1). Four species, Cic-

sci.s (Uivula acinila (Rang, 1828), Liniaciua inflata

(d'Orbignv, 1836), L. trorhifonnis (d'Orbigny, 1836), and

Clio piirainidala j)i/raini(latti Linnaeus, 1767 were re-

cordeil as the douunant thecosome pteropods in the area

throughout the surveyed peiioil. Together they consti-

tuted 86.6% of the total overall pteropod catch (83% in

FebmaiA', 81% in March; 95.6% during May; 74.6% in

.August). The relati\e abundance (%) and mean density

(org./lOin') of each species during the four cruises is

shown in Table 1 .

Overall thecosome pteropod densities were 2.2 times

higher during the night (mean density 298.75 org./

lO'nr) than during da\time (mean den.sitx' 133 org./

lO'm). During the nigtit samplings, theco.some ptei'o-

pods were most abundant during May (499 org./10'm').

E. Siuirez-Morales ami R. Gasca. 19yS

Paiie 45

Table 1. Mean deiisih (ofg./lO'i and felati\e abundance i7c) ot pteropod

Mexitan Canhbean Sea 11991'

species dniin2 eacli iit tonr CARIBE cnii.ses in tin

followed by Febraan.- (332), August (295), and March

(69). Daytime densities were highest during Mav (264

org./10'm'), followed bv Febmarv and August (97), and

bv March (76).

0\erall mean densits' of C. acicula acicula at night

(59.5 org./10'm') was similar to that of davtime samples

(61.9). The same was recorded for Limaeina trochifur-

mis (20.8 (day) vs. 14.5 (night) org./lO'm'). Linuicina

inflata was clearlv most abundant at night (137.2 t.s. 9.8

org./lO'm'), as was C. pi/raniidata pi/rauiiclafa (23 r.s.

9.4 org./lOhn').

In Febnian- the most abundant species was Limaeina

inflata. This species represented 41.7% of the total the-

cosome pteropods, and was mostly captured at night on

the central portion oi the surveyed area (Figure 3e). Li-

maeina inflata was followed bv Creseis aeicula acicula

(24.1%), which shows a broader (hstribiition (Figure 3a),

and by Cavolinia longirostris louoiro.stiis (11.8%). Cre-

seis aeicula acicula occurred in 80% of the stations,

while the other tvvo species occurred in less than 50%.

Limaeina troehifonnis and Clio pijramidata pi/ramidata

occurred onl\' at a fev\' stations (Figure 4 a, e). About

77% of the total thecosome pteropod density' was col-

lected in nighttime samples (da\1:ime mean density': 97

(dav) IS. 332 org./lO'm' at night). The Index of Impor-

tance Value (IIV) showed a shared dominance of L. in-

flata and C. acicula acicula (IIV > 53), followed by C.

pijramidata pijramidata (IIV = 24). Although diversity

was variable (0.6-1.4), on average it was moderately high

(1.22 bits/ind.). Clustering revealed tsvo station groups

(Figure 5e). The first one included stations with less

than 45 org./lO'm', and contained onlv 11.7%' of the the-

cosome pteropod numbers in this month. In most of

these stations Creseis aeicula acicula was the onl\' spe-

Page 46

THE NAUTILUS, Vol. 112, No. 2

CAR/BE/

(February, 1991

CAR/BE//

(March. 1991)

CARIBEIII

(May, 1991)

CARIBE IV

(August, 1 991)

Figure 2. Monthly deiisitie.'i (org./lO'ni') of thecosome pteropods for each sampling; station dnrincr the snneyed period

Me,\ican Caribbean Sea, Black circles indicate night samples.

the

cies present, but with densities below 30 org,/10'ml The

group was distributed in the fully oceanic zone and in

the shelf border area. The second assemblage showed

the same general distribution as the first one; it included

stations with the highest density' values (most over 100

org,/10'm^) (Figure 5a), Liinacina inflata was character-

istic of this group, which comprised 100% of the occur-

rence of the species.

In March, Limacina trochifonnis represented up to

35,6% of the total thecosome pteropod numbers, fol-

lowed by C, ptjramidata pyramidata (27%) and by C,

virgithi conica (18%), Clio piiramidafa ptjramidata and

Crcscis acicula acicula occurred in more than (S0% ot

the stations, the former with higher densities (Figures

3b, 4f), Both species oi Limacina showed a limited dis-

tribution, and were more frequent in night samples (Fig-

ures 3f, 4b), Nighttime densit\- represented 47%- ot the

total catch (mean densit)': 76 (day) us-, 69 (night) org./

lO^m'), Clio pyramidata pyramidata was dominant (IIV

= 56,3), followed by L, trocliiformi.s (49) and C. vir^ula

conica (36), Diversity was moderate (mean 1,14), slightly

lower than in February. Two station clusters were re-

vealed (Figure 5f), with a distribution similar to that oi

February (Figure 5b), The first group was characterized

by low to moderate thecosome pteropod di'usities (2S-

128 org,/10'm'), and mcludetl only 38% ol' the total the-

cosome pteropod density. Up to 75% of C. pyramidata

pyramidata and 96% of L. inflata numbers were includ-

ed in this group, A second group showed low and high

densities (5-344 org,/10'm'), and was characterized In'

the highest densities of L, trochifonnis; this group in-

cluded up to 84% of the total number of C, vir^ula

conica.

In May the most abuuilant species was Crcscis acic-

ula acicula, representing about 54%- ol the total the-

cosome pteropod catch; it was followed by L. injlata

(30%) and by L. trochifonnis (4,4%), Only C, acicula

acicula occurred at all stations, with medium and high

densities (Figure 3c); C, pyramidata pyramidata was

collected at most stations with medium densities (Fig-

in"e 4g), and L trocliiformis was collected at a few sta-

tions (Figure 4c). Limacina inflata was abundant in

nighttime samples on the southern portion of the area

(Figure 3g). Nighttime overall densit\' represented 65%

of the total catch and was 1.8 times higher than davtime

density (499 vs. 264 org./10'm'). The IIV showed the

dominance of C, acicula acicula (80) and of L, injlata

(42,1) in this month. Mean diversit\ was moderate

(1.10 bit,s/ind.), but slightly lower than in the tvvo pre-

vious months. Two clusters were defined (Figure 5 g).

The first one included only four stations with the lowest

densities (9-29 org,/10'm'). Only 0,1% of the theco-

some pteropod numbers occurred in this cluster; these

stations were made in the oceanic area and near the

coast (Figure 5c), Crcscis acicula acicula and Clio pyr-

amidata pyramidata were present in this group. The

second cluster showed the highest total densities (over

98 and up to 1578 org,/10in'), with more than 98% ot

C, acicula acicula and 1009( of L. inflata numbers. This

group showed a variable (h\ersity.

In August, Crcscis acicula acicula was again the most

abundant species (39% relati\e abundance), followed

by Caioliuia low^irostris limhata (19. 3%), by L. inflata

( i6,9%) and by L. trochifonnis ( 16,3%), Crcscis acicula

acicula occurred at all stations with medium and high

densities (Figure 3 d), only both species of Limacina

occurred at more than 50% with medium density \alues

(Figures 3 h, 4 d), while the remaining species each

occurred at less than 20% of the sampling sites, Clio

pyramidata pipamidata was scarce in this cruise (Fig-

ure 4 h). Nearly 75%' of the total densitv' was collected

in nighttime samples; mean density was about 3 times

higlier duruig night collections (295 vs. 97 org,/10''m'),

C'rcscis acicula acicula sliov\ed the highest dominance

E. Suarez-Morales and R. Gasca, 1998

Pa^e 47

CARIBE I

Limacina inflata

Figure 3. Distribution and densit^ of the most ubumlant species during the sunesed period, a-d. Crc:

Liiiiticiiui inflatd. Black circles indicate night samples.

,s'('(.s acicula acicula. e-h.

(IIV = 61.6), while L inflata. C. lonfiirostris limhata

were less dominant. Di\ersit:v fluctuated (0.67-1. 28)

and was in average (1.16 bits/ind.) the second highest

in the survey. Again, t\vo clusters were defined, as

shown in Figure 5h. The first assemblage, with stations

distributed at the middle and southern portions of the

area, was characterized by the lowest total thecosome

pteropod densities (below 20 org./10'm') and contained

onh' 1.49( of the total thecosome pteropod counts. In

this four-station cluster, C. acicula acicula was either

absent or showed its lowest densities. Diversity was low.

The second assemblage was distributed all along the

surveyed area and showed the highest total densities

(40-450 org./10'm') (Figure 5d). More than 98% of C.

acicula acicula. 100% of L trcchifonnifi and of L. in-

flata occurred in this assemblage. Diversity was vari-

able.

DISCUSSION

Previous records ot thecosome pteropods from the

tropical northwestern Atlantic report 51 species and

subspecific forms (Wells, 1975, 1976; Haagensen, 1976:

Suarez-Morales and Gasca, 1990; Michel and Michel,

1991; Suarez-Morales, 1994). Most of these records are

from the Gulf of Mexico and the central and eastern

portions of the Caribbean Sea. All the species identified

here have been reported pre\iouslv from adjacent areas

of the Northwestern Atlantic (Suarez-Morales, 1994).

In a coastal system adjacent to the surveyed area, only

6 species have been recorded (Gasca and Suarez-Mo-

rales, 1990). However the present study shows for the

first time faunal records of thecosome pteropods in the

oceanic area of the Caribbean Sea off Mexico. More-

over, occurrence of Limacina Ic.sucuri. Creseis acicula

Page 48

THE NAUTILUS, Vol. 112. No. 2

Figure 4. Distnliiitioii and ilt'iLsih ot the nuxst aluiiKlaiit .species duniig the suneved period, a-d. Liinacina trochijoniiis: e-h.

Clio ]njniini(lafn jii/rdiiiidata Black circles indicate niijht sanijiles.

clava and Ci/iiihiilia pcroni in the Caribbean Sea is

herein recorded for the first time. These species were

pre\i()uslv known onl\ from the central and sonthern

portions of the Gulf of Me.xico (Suarez-Morales, 1994)

The main branch of the Yucatan Current, which trans-

ports Caribliean surface waters northward, iiffects offsliort^

conditions in the Mexican C>'aribliean. The current repre-

sents an important factor that influences the fauuiil tlistii-

butions and the dynamics of the entire ecosystem. The

pteropod fauna of tropical surface waters of tlie C;iribbean

is dominatefl bv a group of sevenil species (Haaiienseu,

1976), wluch include Cirsci.s (iciailu, Liiiwrina tiDcliiJor-

mis, Creseis virffila and CavoUnia lon^roatris. Only part

of this group of species w;ls found to be dominant in tlic

surface layer of the Caribbean Sea off Me.xico during the

present shidv. The lormer two .species indicate the prinian

influence of die (Caribbean suriace water oii the study ;uea.

The low abundance of C. vArfftla conira and C. langirosth-s

might result from specific, unknown local conditions. The

niiiin species group of the Mexican (Caribbean is similar to

that reported bv Suarez-Moniles (1992> for the southern

Gulf of Mexico (C. aciaila aciaila, L. injiata. L. trochijor-

ini-s, C. lonjS,ir('stri.s lonprostris).

The near-shore and continent;il shelf oceaiiographic dv-

uamics along the (Caribbean coast ot Mexico are related

to the strength of a coast;il coimtercurrent moxing soudi-

wards (Merino, 1986) from the northernmost edge of the

(Caribbean coast, which is iilso the easternmost portion of

the (^ampi'che Bank. Its influence would explain, at least

partialK, a certain affinitx of the local thecosome pteropod

iaiuia with that of the southern (iulf of Mexico, with at

least 60% of species in conunou behveiMi the 2 areas

(Le;il, 1965; Matsubara, 1975; Suarez-Moniles and Gasca,

1992). The species composition of the loc;il thecosome

pteropod fauna seems to represent a mi.xed faunal com-

plex of Caribbean and Gulf of Mexico ;iffinities,

E. Suarez-Morales and R. Gasca, 1998

Pas^e 49

CARIBE I

(February)

fl

14

7

24

23

10

21

15

6

5

20

12

19

11

t^

CARIBE II |_

(March) U

H-

r20

•-19

16

■15

13

21

10

^

14

CARIBE IV

(August)

Figure 5. Distribution of pteropod assemblages a-d. Obtained from the Bray-Curtis Index dunuij; the

drogrums resultinsl from the Bra\-Curtis Index during the tour months suneved.

tour monttis

ths e-h. Den-

In thf surveyed area, the oceanic thecosome pteropod

Limocina hulimoidcs was collected rjuite close to the

coast, as most of the species (either neritic-oceanic or

neritic) recorded here. In Bahia de Ascension, a coastal-

estuarine s\steni on the central coast ot the Mexican

Caribbean (Figure 1), species such as C. acicnia acicnla.

L. trocliifonui.s and C. lougiro.stiif; loiiaimstiis have been

recorded well inside this system (Suarez-Morales and

Gasca, 1990). The occurrence of neritic/oceanic zoo-

plankton within the estuarine systems on this coast has

been studied by Zamponi and Suarez-Morales (1991)

and by Suarez-Morales and Gasca (199fi). According to

local studies of coastal suriace currents (Merino, 1986),

the plankton carried northward by the western edge ot

the Yucatan Current tend to drift inshore. This niiglit

explain the presence of oceanic species yen near the

coast, over the narrow shelf, antl in the estuarine sys-

tems. The inshore drift might also promote the hydro-

graphic homogeneity represented bv the salinity and

temperature values recorded during the sur\e\' period

It is generally accepted that the low variability of these

factors does not have a pronounced effect on the zoo-

plankton distribution in the Caribbean (Hubbard et al.,

1991). Another possible consecjuence of the narrow shelf

present in this area is the presence of deep-sea species

near the shelf break (Segura-Puertas and Ordonez-

Lopez, 1994). Howexer, no deep-sea thecosome ptero-

pods were recoriled during this sunex'.

Most species have been reported as common in ne-

ritic ami oceanic waters, antl represent more than 85%

of the recorded species; C. acicnla acicida. the most

abundant thecosome in the area, has been considered to

be the most aiumdant thecosome in Caribbean neritic-

coastal waters. (Haagensen, 1976). The influence of ne-

ritic yvaters on these otherwise mainly oceanic assem-

blages is represented bv the occurrence of high densities

of C. acicnla acicnla. The study area clearly shows a

strong oceanic affinitx' with a variable neritic influence.

Neritic-oceanic species were mmiericallv dominant or

subdominant durinsi; the four-month survey and can be

Paee 50

THE NAUTILUS. Vol. 112. No. 2

regarded as the m;iiii group ol repre.sentative local pter-

opod.s. Thi.s is probably a result of local mixing processes

within the boundaries of the main current and the coast-

al countercurrent. In a survey of fish larvae, Sanchez-

Velasco and Flores-Coto (1994) found that the Carib-

bean Sea off Mexico is characterized mainly by oceanic

species intermixed with neridc-coastal species.

The Brav-Curtis Index clusters show a similar pattern

throughout the area during the tour sui"veved periods,

with overlapping distributions and irregular z(jnation

patterns. Clusters were sorted m;iinly by whether sta-

tions showed low or high densities. However, day-night

results were partialK' affected bv the migratory circadian

behavior of the most abundant species. This was evident

for Limacina inflata and C. pi/ranudata pi/ramklata.

known to be present during the dav below the 10()-2()()

m, moving up to the surface layer at night (Haagensen,

1976; Wormuth, 1981; Gilmer and Harbison, 1986). In

the sur(.eved area, up to 85% of the local densitv ot L.

inflata and over 70% of C. pijramidata pi/ramidata den-

sit)' was recorded in night trawls.

Clearly, the high-densit\' assemblages were deter-

mined mostly by the night samples; two (C acicula acic-

ida and L. trochiforims) out of four dominant species

were represented bv similar densities during both day

and night samples, as recorded bv Haagensen (1976) in

the Caribbean. This, and the fact that overall dav-night

differences were not significant, would buffer the effect

of circadian variations in our cluster patterns. It appears

that, even with the migration effect, the species com-

position did not varv significantlv, which suggests that

the local thecosome pteropod community is more or less

homogeneous throughout the surveyed area. High and

low densitv assemblages overlapped in several areas din-

ing the four cruises, probably as a result of strong and

continuous mixing processes between the oceanic and

the neritic environments. Monthly variations in theco-

some pteropod density and distribution may be related

to the influence of the Yucatan Current, which increases

in the northern portion of the suivey area during late

spring upwelhng (May). During winter and early spring

(March-Februarv), the current is weak and the environ-

ment becomes oligotrophic (Merino, 1992). This could

cau.se a slight decrease in density (as obsened iluring

March), but the community' stnicture remains othenvise

relatively stable throughout the four-month period.

The studied area represents a nearly homogeneous

mixing zone in which oceanic and neritic-oceanic species

were represented tluring the survey periods by variable

and mixed cbstributional patterns of distriliution. This

could be related to the presence of different environ-

ments (coastal, neritic, oceanic! within a relatively small

area, and to the dispersal and transport caused the local

hydrologic dviiamics (mixing processes, eddies).

ACKNOWLEDGEMENTS

We received financial support from Mexico's (X3NAC\T

(Projects D112-9()452() and 1189-N9203). We gratefully

acknowledge the authorities of the Mexico's 11th Naval

Zone and of the Secretaria de Marina (Depannent of

the Navy) for access to the cruises of the "Dragaminas'

ships. Rosa Maria Hernandez Flores and I. Castellanos

participated actively in the project and in the laboraton'

work.

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drological antl paleoecological indicators in the Tvrrenian

deep-sea core. Palaeogeograpliv, Palaeocliniatologv,

Palaeoecologv' 71:205-224.

Brower, J. E. and J. H. Zar 1977. Field and Laboratory' Meth-

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pp.

Chen, C. and N. Hiilman. 1970. Shell bearing pteropods as

indicators of water masses off Cape Hatferas, North Car-

olma. Bulletin of Marine Science of the Gulf and Carib-

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De la Cruz, G. 1994. ANACOM. Sistema para el analisis de

comunidades. Version 3.0, Manual del usuario. CINMilS-

TA\-IPN, Merida. 99 pp.

Gasca, R. v E. Suarez-Morales. 1992. Pteropodos (Mollusca:

Gastropoda: Thecosomata) de la Bahi'a de la Ascension,

Resen'a de la Biosfera de Sian Ka'an. Quintana Roo, Mex-

ico. In: D. Navarro v E. Suarez-Morales (eds. ) Diversidad

Biologica en la Reserva de la Biosfera de Sian Ka'an,

Qmntana Roo, Mexico. II. Centro de Investigaciones de

Qmntana Roo/SEDESOL. Mexico. 115-122.

Gilmer R. W. and R. G. Harbi.son. 1986. Moq:)hologv and field

behavior of pteropod molluscs: feeding methods in the

families Cavoliniidae, Limacinidae and Peraclididae (Gas-

tropoda: Thecosomata). Marine Biology 91(1 ):47-57.

Haagensen, D. A. 1976. Caribbean zooplankton. Part 11, The-

cosomata. Office of Naval Research, Department of the

Navy. Washington, pp. 551-712.

Hubbard, R. H., ). B. R. Agard and S. N. Gajbhiye. 1991.

Groups of zooplankion along environmental gradients in

the Caribbean. Caribbean M^irine Studies 2(r-2):54-59.

Lalli. C. M. and R. W. Gilmer 19S9. Pelagic snails. The Biologv'

of holoplanktonic gastropod molluscs. Stanford tlniversiU'

Press, Stanford, 259 pp.

Leal, D. G. 1965, Distnhucion de los pteropodos de \'eracnr/,

\'er Anales del Instituto de Biologia de la Universidad

Nacional Autonoma de Mexico 36:249-251.

Ludwig, J. A. and J. F. Reynolds. 1988. Statistical ecology. A

primer on methods and computing. John Wiley & Sons,

New York, 337 pp,

Matsubara, ]. A. 1975, Sistematica, distribucion, ahundancia y

relaciones ambientales de los pteropodos tecosomados de

la Bahia de Campeche. Mexico. Tesis |irotesional, Fac. de

Ciencias, Univi-rsidatl Nacional Autonoma de Mexico,

Mexico, 52 pp.

Menuo. M 1986. Aspectos de la circulacion costera superficial

del Caribe Mexicano con base en obsenaciones utilizando

larjetas de deriva. Anales del Instituto de Ciencias del Mar

\ Limnologia 13:31—46

.Vleriiio, M, 1992, Afloramiento en la plal.itorma di- Yucatan:

estnictura v fertili/aiion, Tesis doctoral, Instituto de Cien-

cias del Mar \ Linmologia. lhii\ersida(l Nacional .Auton-

oma de Mexico, Mexico, 255 pp.

Menno. M, and I„ Otero, 1991. Atlas amhu-ntal costero de

Puerto Morelos Centro de Investigac ioenes de Quintana

E. Suarez-Morules ant! R. Giisca, 1998

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ersidad Nacional Autononia de Mexico, Mexico. 80 pp.

Michel, H. B. and J. F Michel, 1991. Heteropod and theco-

sonie (Molln.sca: Gastropoda) niacroplankton in the Flor-

ida Straits, Bulletin of Manne Science 49:563-574,

SanchezA'elasco, L, and C, Flores-Coto. 1994. Larval fish as-

semblages at the Yucatan Shelf and in the Mexican Carib-

bean Sea during the upwelling period (spring, 1985).

Scientia Marina 58:289-297.

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la coinunidad de medusas (Cnidaria) de la region oriental

del Banco de Campeche v el Caribe Mexicano, Caribbean

Journal of Science 30(1-2):104-115.

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prospecciones de huexos v larvas de peces pelagicos.

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tropoda: Thecosomata v Pseudothecosomata) del Golfo de

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42(3):523-530.

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THE NAUTILUS 112(2):52-57. 1998

Paw 52

A New Deep-sea Limpet of the Genus Pectinodonta Dall, 1882

from New Zealand, and New Distribution Records for P.

aupouria and P. mohoria Marshall, 1985

(Gastropoda: Acmaeidae)

Bruce A. Marshall

Museum of New Zealand Te Papa

Tongarewa

P.O. Box 467

Wellington, NEW ZEALAND

[email protected]\t,nz

ABSTRACT

Pectiiuxltitilii iiuiriiioiichi n. sp, is ile.senbetl antl illustrated.

and new distrihution data are pro\ided tor P. aupouria Mar-

shall, 19S5 and P moriorui Marshall. 1985.

Kei/ wonts: Mollusca, Acmaeidae, Pccthuxhrnta. tieep-sea, new

species. New Zealand

INTRODUCTION

Since the (li.sc()ver\' of two Recent Prctinixlonta species

in the New Zealand region {Marshall, 1985), a third spe-

cies has been recognized, and considerable additional

material of the other two species has come to hand. The

opportunit\' is taken to record this additional material

and to plot the distributions. Note that New Zealand

Pectinodonta species have onlv been found in areas that

have been extensivelv worked by commercial trawlers,

so their tnie distributions are likelv to be more extensive

than implied bv the cbstribution maps presented here

(Figures 6-8). Acronyms: NMNZ, Museum of New Zea-

land, Welhngt(m; NZ'OI, National Institute of Water and

Atmosphi'ric Research, Wellington.

SYSTEMATICS

Superfamily Acmaeoidea Forbes. I S5()

Family Acmaeidae Forbes, 1850

Subfamily Pectinodontinae Pilsbn. 1891

Genus Pcdinoilonta Dall, 1882

Pectinodonta Dall. 18S2: 409. T\])e species (In onginal desig-

natioii); Pectinodonta arcnata Dall, 18cS2; Recent. C!:aril)-

bean

Rcmark.s: Liudberg (198fi) rcdclincd .\cniaeidae, re-

stricted it to include Aciiuieti iniira f-Jathke, 18.3.3 (Ac-

maeinae) and Pcctinodoida species (Pectinodontinae)

alone, and referred most other ta\a hitherto referred

there to Lottiidae. More recently Okiitani ct aJ. (1992)

have introduced two new genera (Scrradonta and Beith-

i/avniaca) for pectinodontines with highly chstincti\e rad-

ulae that live on vestimentiferan tubes and Calijptogena

\'aK'es from a bathval cold seep in Sagami Bay Japan.

Worldwide Recent and Tertian' Pectinodonta species

were listed by Marshall (1985). Liudberg and Hedegaard

(1996) discussed pectinodontine shell stnicture, reciiiit-

ment patterns and ph\logenetic relationships.

Pectinodonta maiinovichi new species

(Figures 1-6)

Pectinodonta aupouria Marsliall. 1985: 277 (in part: largest

measnreil parahpe).

Description: Shell (Figures 1, 2) up to 25.4 mm long,

strongK arched, of moderate thickness, white, exterior

dull, interior porcellanous. Anterior end occupying 20-

■5] % of shell length; anterior slope weakly concave in

most specimens, in others almost flat or weakly con\-ex;

posterior slope rather strongK convex: lateral slopes

wt'akly convex. Aperture elliptical, shallowlv or \er\' shal-

lowly concave from side to side, sides broadly rounded,

anterior end more broadK rounded than posterior. With

growth, shell gradually t^\isting clockA\ise about 10° be-

tween ju\enile and adult. Teleoconch at 1.6-1.8 mm

length translucent and glossy, smooth apart from \ery

fine growth lines: fine radial hues show through outer

shell layer but do not resolve as surface features. Sub-

siMjueiit teleoconch chalk-\- white, with low, rounded ra-

dial ribs, interspaces wider than each rib: and stronger,

rounded or roundK augulate concentric ridges. Fine,

crowded, concentric growth lines throughout. Myostrac-

um clearly defined. Animal with eyes lacking. Foot and

ui:inlle edge cream wliiti'. Large, tapered ctenidium ex-

teuilmg antero-lateralK from left to right behmd head.

Mantle edge weakK and irregularly serrate. Muzzle lap-

B, A. Marshall, 1998

Paee 53

Figures 1-5. Pcctiiuxlcntd inuriiiovichi new species 1, 2. Holohpe, ott Cape Kidnappers, 1100 in. length 25.4 nun. 3-5. Radula

of holot)pe. 3. Inner side of single tooth (length 370 |jLm). 4. Part of ratlnlar rihhon showing teeth in situ attached to basal plates

(width .3.30 (jLni)- 5. Part ot radnlar ribbon showing basal plates from which teeth ha\e been renio\ed (shrinkage is minimal) (width

370 fjLm).

pets thin, broad. Cephalic tentacles small, circular in sec-

tion, tapered, tips rountled. Radular nhbon about 409^

longer than shell. Radular formula 0+3 + 0+3 + 0, each

tooth (as defined here) comprising 3 (used laterals, in-

dividual units ot which are defined bv fine gro(j\'es (Fig-

ures 3-5). Teeth arranged in a posteriorK' tli\erging \'-

shape; each with 13 or 14 conical cusps, innermost cusp

largest, next 3 cusps smaller, outer cusps yet smaller and

similar.

Type data: Holobvpe, NMNZ M. 127085 (length 25.4

mm, width 19.4 mm. height 15.0 mm) and 3 paratopes,

NMNZ M. 127023: Off Cape Kidnappers, 39° 50' S.177°

39' E, ;ilive on wood with P. aupouria, 1100 m, 10 Au-

gust 1995, F.V. Petersen, coll. M. Marino\ich.

Other material examined: Off White Island, Bav of

PlentA-, New Zealand, 37° 23.7' S, 177° 39.5' E, alive on

wood together with P aupouria. 1075-1 10() m, 23 No-

vember 1981, coll. O.M. Moore and P.J. McMillan, F.V.

Kalinovo (stn. KOl/19/81) (3, NMNZ M. 126299); off

East Cape. 37° 32.6' S. 179° 19.3' E, ali\-e on wood with

Page 54

THE NAUTILUS, Vol. 112, No. 2

Figure 6. Map of New Zealand showing distribution of Pectinodonta nmrinovichi new species: 200 and 1000 ni isobaths indicated.

P. aupotiria, 1280-1215 ni, 4 August 1974, R.V. Taiijia-

roa (2, NZOI, stii. KS73/2); N of Cape Runaway, 37°

19.46' S, 178° 04.16' E, alive on wood with P. aupouria,

1124-1169 m, 14 Mav 1995, F.R.V. Tangaroa (stn. 9506/

42) (1, NMNZ M. 126152); off Ea.st Cape, 37° 47.95' S,

179° 04.35' E, alive on wood, 1289-1364 m, 23 March

1993, F.R.V. Tangaroa (stn. 9.303/57) (33, NMNZ

M. 118799); off Gi.sbome, 38° 49.14' S, 178° 34.81' E,

alive on wood, 736-760 m, 25 March 1993, FR.V Tan-

garoa (stn. 9.30.3/69) (6, NMNZ M. 117958); off Cape

Egmont, 38° 58.5' S, 172° 10.2' E, alive on wood, 1045-

1055 m, 3 June 1986, FV Wanaka (stn. WK5/17/86) (1,

NMNZ M.86819); off Mahia Peninsula, 39° 25.7' S, 178°

25.3' E, alive on wood, 921-939 m, 24 June 1986, FV.

Otago CallianI (stn. (JC2/31/86) (20, NMNZ

M. 126569); off Hokitika, 43° 27' S, 168° 47' E, alive on

wood, 910-987 m, 17 May 1985 (3, NMNZ M. 118017).

Distribution (figure 6): Off White Island to off Hok-

itika. New Zealand, lixing and ieethng on wood, 736-

1364 m.

Remarks: Pectinodonta nmiinovichi attains larger size

than P. aupouria and smaller size than P. morioria

(length up to 25.4 mm against 18.2 mm and 14.7 mm

respecti\e]\) It differs further from lioth these species

in that the radial ribs are nnich narrower, and the inter-

spaces are broader than each rib instead of much nar-

rower. The anterior slope is weakly concave in most of

specimens of P. marinoiichi, but convex in most speci-

mens of /' (ntponria The nuintle edge is more weakly

B. A. Marshall 1998

Page 55

Figure 7. Map of New Zealand showing distiilnition oi Pcctinoclonta aiipoiiria Murshall, 19.S5: 200 and 1000 ni isoliatlis indicated.

serrate than in P aiipoiiria. Tlie railiila resembles that

of F aupouiia and has more numerous cusps than that

of P. morioria. P. imirinovichi has been taken twite li\ing

together on the same piece of wood as P. aiipoiiria.

Three specimens of P. marinovichi were found among

the 1232 parat\pes of P. aiipoiiria Marshall, 1985, the

largest of which (length 20.2 mm) was imtortunatelv in-

cluded in a table of shell measurements lor that species

(Marshall, 1985, Table 1).

Etymology: After Miro Marino\ich, F.V. Petersen (Si-

munovich Fisheries, Auckland), who collected the ho-

lotype and who has acquired tremendous quantities of

bvcatch for the Museum of New Zealantl.

Pectimnlonta anpoiiria Marshall. 1985

(Figure 7)

Pcctinoclonta tjiipouria Marsliall, 198.5: 277. fig. 1. 3F-I. 4A,

D (in part; largest paratope = P. marinovichi n. sp.; spec-

imen identified h\ Powell (1979) as Maoricrntcr cxplorata

= P morioria).

Type data: Holotspe, M. 76089 and 1229 paratvpes,

NMNZ; Off White Island, Bav of Plenty-, New Zealand,

37° 23.7' S, 177° 39.5' E, alive on wood together with

P. marinovichi n. sp., 1075-1100 m, 23 Noxember 1981,

coll. O.M. Moore and P.J. McMillan, F.V. Kalinovo (stn.

KO 1/19/81).

Other material examined: N of Cape Runaway, 37°

Page 56

THE NAUTILUS, Vol. 112. No. 2

Figure 8. .\I;i]-) of New Zealand showing ilistnhiition uf Pcctiiiixloiitti inorior'ui Marsiiall. 1985: 2(10 ami lOOO ni isohatlis indicated.

19.46' S, 17(S° 04.16' E, alive on wood witli /' marinov-

ichi, 1124-1169 m, 14 Mav 1995. F.R.V. Tanoowa (stn.

9506/42) (1, NMNZ M. 126151 ); off East Cape, 37° 32.6'

S, 179° 19.3' E, alhe on wood with P. iniraiuiiiclu n.

sp., 1280-1215 m, 4 Angnst 1974. R.V. Taii<ifiroa (9,

NZOI, stn. K873/2); off Gishorne, 38° 39.00' S, 178°

43.5' E, alive on wood, S85-1000 ni, 17 (Ktoher 1988,

F.R.V. James Cook (stn. J 12/45/88) (14, NMNZ

M.92452); off Mahia Peninsula, 39° 05' S, 178° 10' E,

alive on wood with P. inorioria, 900-1000 ni, June 1994,

coll. M. Marinoviih, F.V. Petersen (28, NMNZ

M. 118319); E of Mahia Peninsula, 39° 24.64' S, 178°

27.68' E, alive on wood, 1271-1352 m, 25 Mareh 1993,

F.R.V Tanffiroa (stn. 9303/73) (30, NMNZ Ml 17964);

off Cape Kidnappers, 39° 42.91' S, 178° 10.15' E, alive

on wood with /' inoriorin. 818-952 ni. 8 June 1993. toll.

M. Clark, FR.V Tanzaroa (stn. 9306/210) (4. NMNZ

M. 117997); off Cape Kiduiippers, 39° 50' S, 177° 39' E,

ali\e on wood with P manuovichi, 1100 m, 10 August

1995, FV Petersen, coll. M. Marino\ich (18, NMNZ

M. 127022); N of" Chatham Islands, 42° 45.37' S, 177°

09.74' VV, alive on wood. 1064-1067 m, 23 Jnl\- 1994,

FR.V Tanofiroa (.stn. 9406/383) (16. NMNZ M.ri8817).

Distribution (figure 7): W'lnte Islaiul to oil Cape

Kidnappers, north-eastern North Island, and oil Chat-

ham Islanils. New Ze;iland, lixing and leeding on wood,

818-1280 m.

Remarks: In tlie original description of Pcctiiunlonta

(iiipiHiriu I stated that the shell attains a length of 20.2

nun. This measurement, howe\er. refers to a specimen

of /' iiKiriiioi iciii. uliich w;is iiuludi'd with the t\pe ina-

B. A. Marshall, 199S

Page 57

terial, and tlif largest specimen ol F aupourid seen (a

parahpe) is 18.2 mm long, I was also mistaken in con-

clutling that the Pcctinodonta species misidentified h\-

Povveli (1979) as "Notocratcr cxplomta Dell, 1953" (a

lepetid) was P. aitpouria, for examination of the actual

specimen reveals that it is P. nuniDria. P. anpotiria has

been taken ali\e on two occasions \\ith P iinuioria and

on foiu' occasions with P. inariiiovichi.

Pcctituxlonta inarioiia Marshall, 1985

(Figure 8)

MaorUiYitcr cxplorafa — Powell, 1979: 50, fig, 5: 1, 2 (not No-

toacinca cxplomta Dell, 19.5.3),

Fectinodonta aiipoiiria Marshall. 1985: 277 (in part; Povvell's

(1979) record as Maoiicrntcr cxplomta onl\ )

Fectinodonta niorioria Marshall, 1985: 279, fig, .3]. K, 4B, E,

T>pe data: Hok>t^pe, NMNZ M. 76090 and mam

paratopes, NMNZ NI, 75107: NE of Menioo Bank, Chat-

ham Rise, New Zealand, 42° 43,9' S, 176° 08' E, alive

on wood, 800-810 m, 28 September 1982, F.V. Krdtan

(KTN/152/82). Parat^pes (4, NMNZ M.75242): Off Ti-

maru, 44° 33.2' S. 17.3° 42.3' E, alive on wood, 750-738

m, 8 June 1984, F.R.V. James Cook (stn. JlO/4/84).

Other material examined: Oft Aldermen Islands,

366—176 m, on wood (1, Auckland Institute and Muse-

um AK131610); off Aldermen Islands, alive on wood,

400 m, April 1988, pres. D. Gibbs (2, NMNZ M.95302);

off Cape Runawav, 37° 29.9' S, 177° 47' E. alive on

wood, 450-481 m,'8 December 1985, FV. Wanoka (stn.

\\'K3/19/85) (ca. 1000, NMNZ M.84252); off East Cape,

37° 47.64' S, 179° 04.45' E, ahve on wood, 1289-1342

m. 24 March 1992, FR.V. Taiii^ama (stn. 9203/133) (27.

NMNZ M. 117898); off Maliia Peninsula, 39° 05' S, 178°

10' E, ahve on wood with P. atipoiiria. 900-1000 m, June

1994, coll. M. Manno\ich, FV. Petersen (18, NMNZ

M. 118321); off Cape Kidnappers, 39° 42.91' S. 178°

10.15' E, alive on wood with P. aiipoiiria. 818-952 m, 8

June 1993, coll. M. Clark, FR.V. Tan^aiva (stn. 9306/

210) (.several hundred, NMNZ M. 117996); off Hokitika,

42° 37.6' S, 170° 02.8' E, alive on wood, 490-439 m, 13

October 1988, F.V. Diashin Mam 22 (stn. 269/24) (sev-

eral hundred, NMNZ M. 92467); slope of Mernoo Bank,

Chatham Ri.se, 42° 58.06' S. 175° 01.03' E, N alue on

wood, 545 m, 15 Januarv 1992, FR.V. Tanzania (stn.

9106/91) (manv hundreds, NMNZ M. 126570); E of

Mernoo Bank, Chatham Rise, 42° 53'- 43° 08' S, 176°

04'-177° 00' E, alive on wood, 370-420 m, December

1994, coll. M. Marinovich, FV Petersen (50, NMNZ

M. 119153); off Puv.segur Point, 46° 29' S, 166° 14' E,

alive on wood, 560 m, 20 October 1987, F.V. Cdiii/o

Mant 5 (.52, NMNZ M.90196).

Distribution (figure 8): Aldermen Islands to off Puv-

segur Point, New Zealand, h\ing and feethng on wood,

366-1342 m.

Remarks: Pcctinodonta niorioria proves to attain a

shell length of 14.7 mm, which is considerably larger

than the type material (length of largest paratype 8.30

mm). Apart from smaller size, P. morioria differs from

P. anpoiiria in having more numerous radial ribs, rountl-

ed instead of adapicallv shelved concentric ribs, finer,

more numerous cusps on the radular teeth, and a

smooth instead of serrate mantle edge. P. rnin-ioria and

P. aitpouria have been taken li\ing together on the same

piece of wood on two occasions.

ACKNOWLEDGEMENTS

I am grateful to the numerous fisheries scientists and

obseners (MAFFish, Welhngton) and particularK' Miro

Marinovich (Simuno\ich Fisheries, Auckland), who pro-

vided the material. Thanks to Bmce Hawvard (Auckland

Institute and Museum) for the loan of specimens, to

James McLean (Los Angeles Countv Museum of Natu-

ral Histoi-y) for comments on the manuscript, to Norman

Heke (Museum of New Zealand, WelUngton) for the

photographv and to Wendv St George (Institute of Geo-

logical and Nuclear Sciences, Lower Hutt) for scanning

electron microscope facihties.

LITERATURE CITED

Dall, \\'.H. lSiS2. On certain limpets and chitons from the deep

waters off the eastern coast of the United States. Pro-

ceedings of the United States National M\iseimi 4:400-

414.

Lindberg, D,R, 1986, Name changes in the "Acmaeidae". The

Veliger 29(2): 142-148.

Lindberg. D.R, and C, Hedegaard, 1996, A deep water patel-

logastropod from Oligocene water-logged wood of Wash-

ington State, USA (Acniaeoidea; Pcctinodonta). Jonnial of

Molluscan Stndies 62(3):299-314,

Marshall, B.A, 1985. Recent and Tertiarv' deep-sea Hnipets of

the genus Pcctinodonta Dall (Mollusca: Gastropoda) from

New Zealand and New South Wales. New Zealand Journal

of Zoolog)- 12: 27.3-282.

Okutani. T. E. Tsuchida, and K. F'tijikura, 1992. Fi\e hatl]\al

gastropods li\ing within or near the Cali/ptof^cna-com-

niunit\ of the Hats\ishima Islet, Sagami Bd\. \'enus 51(3):

137-148.

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

freshwater shells. Collins, Auckland, .500 pp.

THE NAUTILUS 112(2):58-62, 199S

Page 58

Rediscovery of Discus hninsoni Beriy, 1955 and Oreohelix

alpina (Elrod, 1901) in the Mission Mountains, Montana, with

Comments on Oreohelix elrocli (Pilsbry, 1900)

Paul Hendricks

Montana Natural Heritage Program

1515 East Sixth A\enue

Helena, MT 596:20

phen(lricks(S'nris.nit.gov

ABSTRACT

During sunniier 1997, extant populations ot the endemic land

snails Discus brtinsoni and Oreohelix alpina were rediscovered

near the type localities in the Mission Mountains, Montana;

last known collections were made 33 and 48 years ago, re-

specti\elv. Discus hniusttiii and Oreohelix elrodi (also endemic

to the Mission Mountains and nearhv Swan Range) were pre-

viously reported to be svnipatric in subalpine limestone talus.

Both .species were found in 1997 in diorite talus; the latter

species also occupied argillite talus with little, if any, limestone

present at any site. Live D. hninsoni [n = S) were foinid near

the talus surface only on bare rock or foliose lichen on rock

during cool, wet conditions. Live O. elrodi were found near

the talus surface on rock (n = 9) or organic litter accunuilations

{n = 55) under all conditions, although in reduced number

during warmer and drier weather Above treelme, live O. al-

pina were found exclusively in exposed locations under lime-

stone talus, sometimes attached to rock surfaces {n = 7) but

more often present in organic detritus {n = 10). Lack of col-

lecting records of D. hninsoni and O. alpina in recent decades

probablv resulted from 1) extremely restricted distributions, 2)

limited knowledge of habitat requirements, and 3) .scarcitv of

active collectors in the region.

Key ivords: Discus hninsoni, Oreotntix nlpina, Oreohelix el-

rodi. Montana, distribution. ecolog\'.

INTRODUCTION

The Mission Mountains of northwestern Montana are

recognized as an area of significant snail endemicitv in

the Interior Columhia Ri\-er Basin (Frest and Joluinnes.

1995). Tile inoimtains extend about SO km north to

south and 19 km east to west, rising ahruptK from a

ha.se at 1036 m in the Swan and Flathead valleys to a

crest of nearly .3()I<S m. The range is composed of Pre-

cambrian Belt rocks (mostly hmestone and argillite) that

were deeply dissected by alpine glaciers during the

Pleistocene (Alden, 1953). The Mission Mountains still

support small circiue glaciers and permanent ice patches.

The portion ot the mountain range within Lake Countv

contains the t\pc localities lor three narrowK endemic

terrestrial suiiil species, none of which has yet been doc-

umented at more than two sites.

Oreohelix elrodi (Pilshn, 1900) was discovered in

1S99 in talus slopes between 1067-1524 ni above the

north side of McDonald Lake (Pilsbn; 1900; Elrod,

1903a), where it was most common, and also found rare-

K' along the south side of the lake. The amphitheater

around McDonald Lake remains the only knowai locahtv

for O. elrodi in the Mission Mountains, but the species

has been found more recently above Lion Creek in the

nearhv Swan Range (Fairbanks, 1984). Oreohelix alpina

(Elrod, 1901) was first found in 1900 on "SinNaleamin

Mountain" (Elrod, 1901; 1902; 19()3b) and again later

that year on the west ridge of McDonald Peak; both

localities were above treehne beKveen 2377-2743 ni and

remain the only sites where this species has been locat-

ed. A single shell of Discus bntusoui Berrv, 1955, was

first encountered in 1948 in talus slopes above the north

side of McDonald Lake, at an estinuited elevation of

1067 m. Additional material was collected there in 1950

(BeriT, 1955; Bnuison, 1956). The type locality rem;uns

the onK known occurrence site for D. hninsoni. Inter-

estingly, this talus slope is also the tvpe locality for O.

elrodi, which had been collected several times between

1899 and 1948 (Brunson, 1956); D. hninsoni. however,

was not discovered there until almost half a century lat-

er Known localities tor the three snail species in the

Mission Mountains are within the Mission Mount;iins

Tribal Wilderness of the Confederated Salish and Koo-

ten;ii Tribes and are relatively secme from human chs-

turbance. None of the snail species has an\' special fed-

eral designation hut all three are on the state fist of An-

imal Species of Special Concern maintained by the Mon-

tana Natmal Heritage Program

Live Oreohelix elrodi were repcuied at the tvpe local-

itv as recentk as 1993 (Frest and Johannes, 1995). There

are no known records in recent years, however, for O.

alpina ;ind Di.scus hnin.'i(nii (Frest and Johannes. 1995).

App;n"enlK, O alpina was last collected on 2 August

1949 on McDonald Peak, ;ind /) hvunsoni was last col-

p. Hendricks, 1998

Page 59

lected on 29 May 1964 at McDonald Lake (R. B. Brun-

son, pers. conini. for both records). Prinian ohjecthes

ot this studv are to 1) re\isit the t\pe localities ot Orco-

Iwlix alpina and Discus bninsoni and, il possible, locate

extant populations of each species, and 2) supplement

previous descnptions of the habitats associated with each

species.

MATERIAL AND METHODS

I usetl a variety- of sources to obtain relati\'el\ precise

descriptions of tvpe localities and habitat use by Orco-

hclix alpina and Discus bninsoni, inclucbng pubhshed

hterature, conversations uath Dr R. B. Brunson (who

made the last kniown collections of each species), and

examination of Brunson's specimen catalog. The type lo-

cality of D. brunsoni was easily identified. Berry (1955)

provided a photograph of the site and Brunson (1956)

added considerable det;ul of its conditions and physical

setting. The site ( Site 1 ) is an extensive talus slope on

the north side of McDonald Lake (T19N R19\\' Sll)

that is accessible by trail. Trips to Site 1 were made on

10 June, 1 Julv, 4 jiiK' and 10 JuK- 1997. Rock at Site 1

was composed predominantly of bk)cks of dioiite about

1 m' in size, intermixed with less than 5% ot argillite

fragments. Neither rock material reacted with acid,

which indicates the absence of calcium carbonate. Can-

opy coyer was 0% in the search area, slope = 36" facing

SSE. Forest at the margins of the talus slope was com-

posed of Douglas fir (Pseudotsuga menzicsii) and pon-

derosa pine (Pinus pondcrosa). Pockets of water birch

(Betula occidcntalis). quaking aspen (Populus trcmuloi-

dcs) and mock orange {Philadclpluis Icuisii) were scat-

tered in and near the margms of the talus. Their leaves

comprised much of the organic litter among the talus

fragments. Talus wns inspected to a maximimi depth of

1 m in an area of 25 X 40 m. Additional searches during

other visits to Site 1 were focused in this area. Site 2

(about 100 m east of Site 1) was composed of about

equal amounts of smaller-sized diorite and argiHite; av-

erage talus fragment size was about 30 X 30 cm. Canopy

cover (surrounding species composition as at Site 1) was

0%, slope = 32" facing SSE, search area was 15 X 20

m. Site 3 was about 500 m east of Site 2. Search area

at Site 3 was 10 X 25 m, rock was 100% argillite with

average fragment size about 10 X 20 cm. Slope, aspect,

cover and surrounding vegetation at Site 3 were similar

to the other two sites.

Access to the historical sites of Orcoliclix alpina was

not so straightfori.vard. Neither location can be reached

by trail. The McDonald Peak site is in the Cirizzlv Bear

Conservation Zone of the Mission Mountams Tribal Wil-

derness and is closed to all recreational use from 15

Julv-1 October. Snovvpack often precludes trips into the

alpine areas before mid-Julv, so the McDonalil Peak site

was not visited in 1997. Elrod (1901; 1902; 1903b), Pils-

bry (1939) and Frest and Johannes (1995) describe the

type locality as above treeline on "Sinyaleamin Moun-

tain". There is no Sinyaleamin Mountain on current

uses, topographic maps. Details of the 1900 trip ilur-

ing which the species was discovered (Elrod, 1902) in-

dicate, however, that Sinyaleamin Mountain is now

known as East St. Marys Peak; O. alpina was collected

by Elrod on the southwest ridge of that mountiiin (TISN

R18W S20).

I climbed East St. Mans Peak on 27 August 1997 and

searched for Orculwlix alpina at two sites (Sites 1 and

2) along the southeast ridge (T18N R18W S21) and also

along the top of the southwest ridge (Site 3), which in-

cluded the sinnmit. Site 1 was a broad level area atop

the southeast ridge at approximately 2637 m; the ground

was covered with a varietv of alpine forbs and grasses

(vegetation cover = 60-70%) in which were open patch-

es of limestone scree (fragment diameter about 2-4 cm).

Site 2 was farther north about 1 km, where the ridge

narrowed between 2713-2774 m. Exposure at this site

was to the southwest but near the ridge crest. Shallow

limestone talus became predominant with only scattered

patches of alpine vegetation present (cover = 8.5-95%

rock). Plants m the area mcluded snow cinquefoil iPo-

tcntilla nivea). mountain avens (Dn/as octopctala), al-

pine sorrel (Oxiiria difii/iia) and moss campion iSilcnc

acaulis). Site 3 included the summit (2873 m) and top

of the southwest ridge. Plant ccjmposition and percent-

cover at this site were similar to Site 2.

Time spent searching for each species was recorded

as a measure of search effort (Ausden, 1996); time of

day refers to Mountain Daylight Time. The substrate on

which each live snail was found ("rock" or "organic lit-

ter") was noted, and shell diameter of live individuals

was measured with a dial ciiliper. Photographs ot live

sn;iils and habitats for each species were taken and are

available from the Montana Natural Heritage Program,

Helena, Montana. Voucher specimens of Discus bnin-

soni (MTHP 4069) and Orcohclix alpina (MTHP 4071)

were verified bv R. B. Brunson and will be deposited at

the U.S. National Museum following anatomical study

by H. L. F;urbanks. Only empty shells of O. clrodi were

collected for reference material, as the species is very

distinctive and readily recognized from photographs. G-

tests (Sokal and Rohlf, 1981) were used to analyze fre-

quency distributions of substrate use; statistical signifi-

cance was assumed when F<0.05.

RESULTS

DlSClS BRiSSOM WD OrEOUEUX ELRODI

Presence at different sites: Three talus sites within

the general tvpe localitv of both species were examined

during four different visits (Table 1). All sites were at

1128 m along the trail parallehng McDonald Lake on its

north side. On 10 June and 4 July, weather was sunny

and warm (21-23"C) at the time of the searches (10:15-

12:00); no precipitation had fallen in the previous 48

hours. On both dates only Site 1 was checked. Four live

estivating Orcohclix clrodi and no Discus bninsoni were

found during 75 min of searching bv two persons on 10

Page 60

THE NAUTILUS, Vol. 112, No. 2

Table 1. Maxiniuin diameter (iiieaii ± SD) of live Orcolielix clrodi ami Discus hninsoni in three talus sites on 1 and 10 JuK 1997.

Mission Mountains. .Montana. Ail measurements are in nun. .Sample sizes in I are in parentheses.

Site 1

Site

Site .3

Rock t\pe

Axerage fragment size rn'

O. elrocli

mean

range

D bninsoni

mean

range

diorite, argillite

0.09 nr

14.0 ± 4.7 (IS)

7..5-22.3

argillite

0.02

12.0 ± 4.2 (7)

6.,5-17.7

June. Tlie 4 juK- \isit vva.s devoted to studv of the rocks

and no Ii\e sn;iiis of either .species were noted.

On 1 JuK' weather was wet and cool (10"C) during the

search period (10:00-13:00); the area had received

steadv precipitation during the previous 4S h. At Site 1

I found 20 li\'e Oivohclix clrodi and 5 h\e Diaciis bnin-

soni in 120 min. At Site 2 I found 11 O. clrodi and no

D. brtimoni during 60 min. Under similar conditions on

10 Julv (wet, 14-i7"C) I found 19 live O. clrodi and .3

Lve D.' bninsoni at Site 1 during 120 min (11:00-13:00).

At Site 2 I found 7 live O. clrodi and no D. bninsoni in

30 min (13:30-14:00). At Site 3 I found 7 live O. clrodi

and no D. bninsoni in 15 min (15:35-15:50).

Substrate .selection within talus: Sulistrate of occur-

rence (rock, organic litter) for live snails foiuid at all

three sites was documented on 1 and 10 Julv. All li\e

Discus bninsoni (n = 8) were found on bare or lichen-

covered rock. Nine of 64 live Oreohclix clrodi were

found on similar substrate, 55 were found on organic

htter accumulations (bark, Kvigs, leaf fragments, nee-

dles) or h\e moss among the talus. The difference in

substrate selection between the two is statisticallv sig-

nificant (G = 26.724, df = 1, P < 0.001).

Shell diameter: Discus bninsoni measured in average

9.5 mm diameter and ranged from 6.7-10.5 mm (Table

I). F(jr Oreohclix clrodi at Site 1, 28 (71.8%) of 39 live

indivaduals were > 15 mm diameter. At Sites 2 and 3

the respective ninnbers of live snails in this category

were 6 (33.3%) of 18, and 2 (28.6%) of 7. Sample,s\m-

doubtedlv contained members of more than one cohort

The presence of inchviduals < 7.5 mm diameter in each

sample suggests reproduction is occurring at each site.

Oiu'.ouiLix \i.ri\-\

Presence at different sites and .substrate selection:

Weather on East St. Mans Peak was clear and cool

(12.5"(; at the smnmit) on 27 August. I touuil no snails

in 30 min (12:15-12:45) at Site 1 on the .southea.st ridge.

At Site 2 I found 16 live O. (dpina in 45 min (13:30-

14:15) in three areas searched along 300 m of ridge. All

live snails were imder limestone blocks about 20 X 30

ciri scpiare and 4-8 cm thick. Six snails were attached to

the undersiiles ol linicstone Iragmenls or atop iiare rock

beneath (deriving blocks, one group of 10 li\e snails was

founil in leaf litter (area = 9 cm-) accumulated near the

base of snow cinquefoil. Some soil development was

present at each site where li\e suiiils were found. Dead

shells were found with little effort on open ground.

I searched the summit area (Site 3) for 15 min (14:

30-14:45) and found one live snail at 2865 m near the

top of the southwest ridge. This individual was on bare

rock under a limestone block near a patch of snow

cinquetoil.

Shell diameter: Diameter of h\e shells (mean ± SD,

n = 16) was 5.7 ±1.9 mm; range was 2.5-8.5 mm. The

smaller individuals indicate that reproduction is probably

occurring at this location. Mean diameter of dead shells

(n =9) was 8.9 ± 0.5 mm; range was 8.0-9.6 nun.

DISCUSSION

Extant populations of Discus bninsoni and Oreohclix al-

pina were located in 1997 at or near the t\pe localities

33 and 48 vears, respectively, after the last docinnented

collections (R. B. Brunson, pers. comm.; T J. Frest,

pers. comm.). The range in shell size of live individuals

at each site (including O. clrodi sites) indicates the pres-

ence of multiple cohorts and likelihood of continuing

reproduction. The sites where O. alpina was found on

the southeast ridge of East St. Marvs Peak represent a

slight range e.xpansion. (The type locality is on the south-

west ridge.) I have foimd no evidence that O. alpina has

been collected on East St. Mans Peak since 1900, when

Elrod discovered the species. The last documented col-

lection of O. alpina (Bninson specimen cat;ilog) was

made on McDonald Peak on 2 August 1949. Correspon-

dence from Stillman Bern (21 .\ugust 1951: Bnm.son

pers. comm.) indicates that another collection mav have

been UKide on McDonald Peak in 1950 or 1951. but

tliere is no evidence of this in Bnmson's c;it;il()g

Several intenicting factors prob;iblv contributed to the

failure to find Discus bninsoni and Oreohclix alpina dur-

ing the last several decades. First, I know of no active

resident collectors of terrestrial mollusks. Non-resident

collectors passing through the area m;iy luive visited sites

when I'onditions were not espei i;illv tavonible tor finchng

these siiccies ( D bninsoni in particular) near the surface

p. Hendricks. 1998

Paee 61

of tains slopes. The nearlv halt-centim' span liehveen

first tliscoveries of O cirodi and D hniiisoui at tlie same

site supports tliis contention. Differences in lialiitat se-

lection mav have contributed to this lapse, as will he

discussed below. Furthermore, it seems unlikely that

\er\' nianv non-resident collectors would attempt to xdsit

the relati\elv inaccessible localities of O. ulphia. Second,

the known ihstributions of both species are (|uite re-

stricted. It would be easv for a collector to o\erlook

them without knowledge of the geographical area. For

example, one cannot identifv "Sin\aleamin Mountain" or

find the exact location of O. aljiina on this moimtain or

on McDouiild Peak without a c(jpv of Elrod's (1902) de-

scription of his collecting trip. Third, habitat retjuire-

ments, especiallv for D hniiisiyni. mav be narrower than

pre\iouslv appreciated. Information gleaned from Beny

(15^55) and Brunson (1956) suggests that D. bninsoiii

and O. cirocli are ,s\nnpatric in talus slopes on the north

side of McDonald Lake. Howexer. it ma\ be that both

species co-occur in onlv a verx' restricted zone defined

bv the narrower habitat preference of D Ijnnisoui Ev-

idence to date indicates that O. el rod i is significantK

more widespread, even in the McDcjnald Lake cinjue.

All three sn;iil species favor exposed talus habitats.

The t\pe of talus in which each species is found, how-

ever, appears to differ among species. Orcohclix alpina

is found exclusivelv above treeline in shallow limestone

talus on mount;un ridges. The descriptions proxided by

Elrod (1901; 1902; i903b) and Frest and Joliannes

(1995) generallv coincide with my 1997 findings. Elrod

(1902) commented that the snails were found among

and imder rocks with little \egetation nearbv. All five

imlixiduals I located were under the protection of stones

in areas of scant \egetation cover, Init usnalK with some

soil and litter accumulation nearby. Ten (62.5%) of 16

li\e indi\iduals that I could measure (one shell was bro-

ken during handling) were sm;iller than the minimum

diameter (7 mm) described for this species (Elrod,

1903b; Pilsbr)', 1939). However, size range of empty

shells found mostly on open ground closelv matched the

published range and mean. Perhaps adults are more Uke-

1\' to be caught awav from refuges in aiKerse conditions,

with dead adult shells washing onto open ground

Discus hninsoui and O. cirodi are found well below

the treeline in t;ilus slopes surrounded bv closed- and

open-canopv forest (Biimson, 1956), but the rock txpes

comprising the talus inhabited hv the snails are infre-

(juentK mentioned in pubfished reports. Berrv' (1955)

and Frest and Johannes (1995) identified the talus as

limestone; Elrod (1901; 1902; 1903a) and Bmnson

(1956) failed to mention rock composition. I did not de-

tect limestone at the three sites where I foimd one or

both species. Discus bninsmii was found exclusivelv in

talus where diorite boulders predominate. Orcoliclix ci-

rodi was found at this site as well as in smaller-sized talus

of argillite (Table 1). Discus bninsoni may have narrower

habitat recjuirements than expected, associating only

with a particular subset of a\ailable rock types.

The biologv and ecological requirements of Discus

hninsoui and Orcohclix cirodi remain largelv unknowii,

antl the following explanations for patterns of presence

or absence near the surface of talus slopes are specula-

ti\e. Orcohclix cirodi was more abundant (4-6 fold dur-

ing mv searches) than D hruusoni near the surface of

talus in wet and cool conditions, and a few indi\iduals

could still be found when it was warmer and drier. Sev-

eral factors could contnbute to this pattern. First, ab-

solute population size of O. cirodi at Site 1 may be great-

er than that of D, hruusoni. Ratios of each species near

the talus surface could be representative for all depths

in talus. Second, I found significant tlifferences in sub-

strate selection between the hvo species. Some D. hnin-

soui were found on foliose lichen (tentatively identified

as Arcfopunncliii suhccntrifu^a) that grows on the dio-

rite, but most uuli\iduals were found on bare rock. In

contrast, O. cirodi were most often found on organic

litter and \egetation. Preference by O. cirodi for organic

litter mav keep them nearer the surface of talus slopes

where fitter accinnulations are larger and apparentlv

more numerous. Thnil. larger shell size of O cirodi

might reduce its rate of desiccation bv decreasing the

surface are;i/\'olume ratio (see Goodfriend. 1986), allow-

ing larger individuals to remain nearer the drier talus

surface for longer periods than D. hninsoui and small

O. cirodi. The four live O. cirodi found on 10 June in

warm and diT conditions measured 18-21 mm diameter

and were estivating at that time. Fourth, timing of m\

searches did not allow for detection of this pattern, but

D. hntihsoni may tend to be nocturnal or crepuscular and

make \'ertical migrations to the talus surface during pe-

riods of acti\its' (Bnmson, 1956).

ACKNOW'LEDGM ENTS

Field work was generonsK supported bv a Canon Ex-

ploration Grant aihnmistered by The Nature Consenan-

cy and Canon U.S.A., Inc. I benefited greatly from con-

versations with T. J. Frest and especiallv R. B. Bnmson,

whose knowledge and documentation of the snails of the

Mission Mountains were enthusiasticallv shared when-

ever requested. J. S. Marks and L. M. Hendricks partic-

ipated in the hunt for Discus hninsoui; T Gignoux iden-

tified the rocks at the D. hninsoui site. T Shreve \isited

the tvpe localiU of Orcohclix alpino with me. An earlier

draft of the manuscript benefited greatlv frf)m the com-

ments of Uvo anonvmous reviewers. This paper is decfi-

cated to the memor)' of the late J. R. Reichel of the

Montana Natural Heritage Program, who supported my

efforts even before this project was conceived.

LITERATURE CITED

Alden, W. C. 1953. Phvsiographv and glacial geoIog\- of western

Montana and adjacent regions. U. S. Geological Survey

Professional Paper 231:1-200.

Ausden. M. 1996. Invertebrates. In: .Sutherland, W. J. (ed.)

Ecological census techniques, a liandbook. Cambridge

Universitv Press. Caniliridsie. pp. l.'59-177.

Page 62

THE NAUTILUS, Vol. 112, \o. 2

Berrv', S. S. 1955. An important new land-snail from the Mis-

sion Range, Montana. Bnlletin ot the Sontheni Calitoniia

Acadeniv of Sciences 54:17-19.

Brunson, R. B. 1956. The nivsterv of Di.sais hninsoni. The

Nautilus 70(1):16-21.

Elrod, M. J. 1901. Montana shells. Rocky Mountain Magazine

2:691-697.

Elrod, M. J. 1902. A biological reconnoissance in the \icinit\-

of Flathead Lake. Uni\ersitv of Montana Bulletin No. 10,

Biological Series No. 3.

Elrod, M. J. 190.3a. Notes on Pi/raniirluhi clrodi Pils. The Nau-

tilus 16(10):109-1 12.

Elrod, M. J. 190,3b. Montana shells — Pi/nniiidiilri strii^^osa. The

Nautilus 17(1): 1-6.

Fairbanks, H. L. 1984. A new .species ot Onohclix (Gastrop-

oda: Puhnonata: Oreohelicidae) from the Se\en De\-ils

Mountains, Idaho. Proceedings of the Biological Societ\'

of Washington 97:179-185.

Frest, T |., and E. J. Johannes. 1995. Interior Cohunbia Basin

mollusk species ot special concern. Final Report to Inte-

rior Columbia Basin Ecosystem Management Project.

Dei.\is Consultants, Seattle, 274 pp.

Goodfriend, G. A. 1986. Variation in land-snail shell fonn and

size and its can.ses: a resiew. S\stematic Zoolog\' .35:204-

223.

Pilsbry, H. A. 1939. Land moUnsca o\' North Amenca (north

of Mexico), volume 1 part 1. Monographs of the Academx'

of Natural Sciences of Philadelphia (3), Philadelphia, 57.3

pp.

Sokal, R. R., and F J. Rohlf 1981. BiometrA. Second Edition.

VV. H. Freeman, San Francisco, 859 pp.

THE NAUTILUS 112(2):63-68, 199S

Page 63

Epitonium fabrizioi (Gastropoda: Epitoniidae), a New Species

from Patagonia, Argentina

Cuido Pastorino'

Departaniento Paleozoologi'a

Invertebrados

Museo de La Plata

Paseo del Bosque s/ii

1900 La Plata

ARGENTINA

i"vpastor@criba. edu. ar

Pablo Penchaszadeh

INTECMAR

Universidad Simon Bolivar

1080 Caracas

VENEZUELA

and

Museo Argentino de Ciencias

Naturales "Beniardino Rivada\ia"

Angel Gallardo 470

1405 Buenos Aires

ARGENTINA

ABSTRACT

Epitonium fabrizioi, a new species of gastropod mollusk be-

longing to the family Epitoniidae, is described from shallow

waters of Puerto Piraniides, Chubut Pro\ince, Argentina. This

new species is similar to E. gcorg,fttiniim from the same area,

which was known only from its shell. It can be distinguished

from E, ^eoracttimim by its smaller size, more straight profile

antl, b\' its sharjier, most numerous ribs. In addition, the pro-

toconch of the new species consists of 4.25 whorls. Its radula

has only marginal teeth, each with three cusps. The outermost

cusp is larger and hook-like, the central and inner cusps are

shorter anil similar in size. An additional, ob.solete, cusp is pres-

ent near the base ot the teeth. The egg capsules ha\e a mean

diameter of 76 \).m. The new species is compared with £. ^cor-

^ettinuui. the species with which it co-occurs, and £. albidnm.

INTRODUCTION

Most recent papers about the taiinlv Epitoniidae deal

with .species from the northern hemisphere (CJarihliean:

Robertson, 1983a, 1983b, 1994a, 1994b; northeastern

Pacific: DuShane, 1974, 1979; northeastern Atlantic:

Bouchet and Waren, 1986). An e.xception is the work of

Kiiburn (1985) who studied representatives ot the taniilv

from South Africa and Mozambicjue, and provided a

good account of the subgenera living in that area. In

their classic and comprehensive work. Clench and Tur-

ner (1951, 1952) reported most of the knowii western

Atlantic species. Rios (1994) included a complete list of

the approximately 30 species of Epitoniiilae knowm from

northern South America, together with illustrations and

distribution ranges. However, the material examined in

' Current Address: Department of Invertebrate Zoolog)', Di-

vision of Mollusks, National Museum of Natural History,

Smithsonian Institution. W'ashinsTton, DC 20560, USA.

his work was primarily from Brazil, and only margmally

included representative species from Uruguay and

northern Argentina.

The Magellanic area is poorly represented in mollus-

can faunal reports, and almost never represented in pub-

hshed revisions of Epitoniidae. Strebel (1905) was the

last author to publish a systematic account of the Mag-

ellanic epitoniid fauna. He described one species and

one variety from the Strait of Magellan.

In this paper we describe a new species of Epitonium,

and include illustrations of its radula, operculum and

protoconch. Data on eggs and egg capsules are also in-

cluded.

MATERIALS AND METHODS

Specimens of Epitouiitin fabrizioi were collected from

tidepools near Puerto Piramide, Chubut, Argentina

(42°34'S, 64°17'W) in November, 1995, during .spring

(lower low water) tides, and near Punta Loma (42°49'S,

64°53'W) in February, 1996 (see Map 1). This species

was found only in the intertidal zone around the pedal

discs of sea-anemones (genus Biinndoctis?) on which

they probably feed. In several cases, more than one

specimen was found on the same sea-anemone. Several

animals were dissected and the radulae were prepared

for the SEM obsei'vation. The protoconch whorls were

counted folltming the method of Jung (1986). Institu-

tional abbreviations used are: ANSP, Academy of Natural

Sciences of Philadelphia; MACN, Museo Argentino de

Ciencias Naturales "Bernardino Rivadavda", Buenos Ai-

res, Argentina; MHNC, Museum d'Histoire naturelle de

Geneve, Switzerland; MLP, Departaniento Zoologia In-

vertebrados, Museo de La Plata, La Plata, Argentina;

USNM, National Museum of Natural Histoiy, Smithson-

ian Institution.

Page 64

THE NAUTILUS, Vol. 112. No. 2

Figures 1-9. Epitoninin jahrizioi new .specie.s. 1-3. Holotvpe, MLP 54():i, .Scale bar = 5 nnii. 4. Apical \ie\v, MLP 5.3.3.3. Scale

bar = 200 (xiii 5. Protocoiicli, MLP .5.3.33. Scale bar = 2.50 |j.rri. 6. Operculum of holoUpe, Scale bar = 5 mm. 7. Radula of

liolotype in '.^eiieral \ie\v. Scale bar = 20 |xm. 8. Detail of luar'Tinal teeth. Scale bar = 10 jjim 9. Enn capsules. Scale bar = 1 nun.

SY.STEMATICS

KaiTiilv Epitoniiilac BcriA', 1910

Cenu.s Epitoninin K()cliiii», 1798

Epitoniiini fahrizioi new specie.s

(Figures 1-9)

Epitoninin alhidum (d'OrbifiUN. bS42,). — Ciencli .mil 'runiei,

1951: pi. 114. fi" 3 only. '

Epitonmni '^rorui'ttiiia (Kiener. 1S.39). — Scarabuio, H)7T:1.S3,

pi. 2, fig. 8.

Epitoninm '^eori^fttinai' (Kiener, lS.39i, — Paslonno, 1995:S, pi.

2. fig. 10,

Dcseriptioii: Shell sinali, iij) to 1.3 nun leniitii, clialkT,

thill, opaque, Frotoeoneli iWoni in all but oni' spei-inien)

eon.sisting of 4.25 whorls and nie;isuring 4S5 length X ,390

fjLm width. Protoeoneh whorls smootli imder SEM, tran-

sition to teleoeoneh not vei"V' shaip. Teleoeoneh with S

gentK' convex, smooth whorls. Spire angle 40°. profile

strmght. Suture crossed li\ tenuination of a\i;il ribs (fenes-

trate). A.\ial ornamentation cousi.sting of vei"v shiup ribs, up

to IS on last wiiorl, hut usuiillv l.'3-14 (Tal)le 1). A.\i;il ribs

slighlK o!)li(ine in relation to shell a\is, with iiTegularlv de-

fined edges near sntine, and ri'mnants from breakage in

the r("st. .\.\ial ribs gentlv ri'fleeted on last whorl. y\ll ribs

perteitK iiligned witli overlapping ribs from preceding

whorl; attachment erect. Ajiei+nrt' o\al, penstome with one

la\'er, forming a getitle basal expansion (aunculate). Colu-

mell.ir callus weakK' deNcloiied. L'mbilicns i-lo.si'd.

G. Pastorino and P. Penchaszadeh, 199S

Page 65

Puerto

Madrvtf*-mf Lonia

-42 S

P Ninfas

to 5 0 10

' ■ '

Map 1. Records o( Epitoniuin fabrizioi (® = t\pe localit)')

Operculum oval, paucispiral, thin, translucent, hrown-

i.sh, entire surface co\'ered b\' growth lines.

Radula lacldiig central and lateral teeth, but with nu-

merous tricuspid marginal teeth, identical in shape. Out-

ermost cusps large and hook-hke, central and inner

cusps similar in size and shoiter Secondar\' cusp present

centrally on teeth near base.

Egg masses always found near (but not attached to)

adults. Egg masses composed of 180-320 egg capsules

attached to each other b\- tough elastic string as in other

Table 1. Epitnitiiiin fabrizioi Shell measurements and number of axial ribs per whorl. Numbered column headings refer to whorl

munber, Duslies indicate worn nbs on tliat wliorl.

Shell

No

Lens

\N'idt!i

Teleo-

concli

whorls

16

IS

Page 66

THE NAUTILUS, Vol. 112, No. 2

Figures 1()-16, 17-21. EjnUiminn ficor<^cttiiiiiiii (Kit-iu-r, 1S.391, 10-12. A[ii-rtunil, basal, ami a]iic.il \it-\\s. MLP 54(14. Puerto

Piranuiles. Scale bar = 5 niiii. 13. Operculum of a 24.72 mm length specimen. Scale bar = 10 nnii 14. Scalaria ^corgeftina,

paralectohpe, MHNG 98.3/111/2, "Ocean Atlantique". Scale bar = 10 mm. 15. Raduia of the specimen shown in F'igiires 10-12.

Scale bar = 20 (xni. 16. Detail of the railula. Scale bar = 10 jjim. 17-21. Epihmhim albidum (d'Orbigiiy, 1842). 17^ 18. USNM

4.39556, Baie Anglai.se, Haiti. 17. Apertural \iew. Scale bar = 1 nnn 18. Detail of the shell surface. Scale bar = 10 (jLm. 19. Detail

of the same area under higher magnification. Scale bar = 2 fim 20. Protoconch. Scale bar = 40 jjim. 21. Apical view. Scale bar

= 40 |xm.

species of sanK' gi'inis. Largest egg capsules iwaminci!

measured 1..35X1.1 nmi (ii = 2(); x= 1 .19X0.94; Sn =

0.()S()X().()69). Egg capsules pyramidal or polyhedral in

.shape and covered with sand grain.s. Nmnher of eggs per

capsule ranged 69-141 (n=l(); x= 123.9; SD = 2().12).

Uncleaved eggs measured 76 \i.m in diameti'r.

Type locality: Puerto I'inimides. (-'huhut Pro\iuci-,

Argentina (42'°34'S, 64°]7'VV).

T\pc material: nolot\pe. MLP 5102; 10 paratopes,

USNM SS0252; I parat\pi>, MLP 5.33.3; 4 parahpes.

ANSP A1S86(); 10 paratypes MLP 5403; all from the

t\pe loc;ilit\-, (;. Pastorino coll., November 1995; MLP

5333, 1 specimen (protoconch), from near Punta Lonia,

Chubut Province. Argentina, 42°49'S, 64°53'\V, J. Mer-

moz coll., Februarv 1996.

Other material e.xamined: Five specinien.s. MACN

muuunbered, trom near Punta Loma. Chubut Province,

Argentina, 42°49'S, 64°53'W, J. Mermoz coll.. Feb.

199(i.

Etviiioloijy : Dedicated to Fabiizio Scar;ibino, voung

niiihii'ologrst ;uul Iriend, who called our attention to the

new siH'cies.

G. Pastorino and P. Pencha.szadeh, 199.S

Page 67

Table 2. Epitoniiii}! acorvcttiiiiim (Kieiiei). Shell measurenient.s and number ot axial nbs per whoil. Nuniberetl tohuini headings

refer to whorl rnniibei' Dashes indicate uoni ribs oir that uhorl.

Epitoiiitim gcorgetfiuuin (Kiener, 1839) is the most sim-

ilar species to E. fabrizioi. It also occurs in the httoral

zone in the Puerto Pinimides area, where it lives near

sea anemones on hard sulistrate, but, unlike E. jahhzioi.

it can also be found on sandv bottom.

The type specimens of Epifoniitm gcorgcttiiuiin

(MNHG' 983/11 1/1-3) were illustrated bv Clench and

Turner (1951, Pi. 117, fig. 1). Figure 14 herein repre-

sents one of the paralectotvpes (MHNG 98.3/111/2). The

main conchological differences between E. gcorgettiiunn

and the new species is the profile of the shell, which is

straighter in £ gcorgcttiniiiii. and the whorls, which are

more con\ex and detached in this latter species. The rilis

in E. fabrizioi are sharper, thinner, and increase in num-

ber with growth (Tables 1, 2). Epitonium gcorgcttiniim

has a multispiral, thick, and opaque operculum (Figure

13), whereas that of E. fabrizioi is paucispiral, thin, and

translucent (Figure 6). The radula of E. georgcttinitm is

also distinct. It has one sharp terminal cusp per marginal

tooth, and two lilimt cusps, one almost obsolete (Figures

15-16).

The eggs and egg capsules ol these two species are

quite similar (Pastorino and Penchaszadeh, in press).

However, based on the material available, £. fabrizioi

has smaller egg capsules, each containing fewer eggs al-

though egg diameter is nearh' identical to that in E

gcorgctiiniiiu Bell (1985) has indicated the lar\al fife-

span and size in E (//;/ Pilsbrv, 1921 (26 days; 3 whorls,

390 |jLm). The larger number of protoconch whorls and

size oi Epitonium fabrizioi (4.25 whorls, 485X390 |xm)

suggests that E. fabrizioi spends more time than E. (//(/

in the planktotrophic lanal stage. In comparison with

other known species of Epitonium, the new species has

the largest protoconch (Table 3).

Epitonium albidum (d'(Jrliigny, 1842) is suuilar in

shell shape, liut its protoconch shows a deficate (visible

under SEM) ornamentation that contrasts with that in

the new species (Figure 20), In adcfition, the entire sur-

face of the teleoconch in E. albidum shows a character-

istic pitted microsculpture (visible under SEM; Figures

18, 19). This character is not t)pical of the genus Epi-

tonium and may support a new generic allocation for E.

gcorgettiiunn, as suggested bv Bouchet and Waren

(1986) for E. albidum^ In contrast, E. fabrizioi lacks this

h-pe of microsculpture. Robertson (I983ii) pointed out

that Clench and Turner's (1951) Argentinean record of

E. all'idum from Bahia San Bias was zoogeographicallv

anomalous. Their illustration (Clench and Turner, 1951;

Plate 114, fig. 3) represents a specimen of E. fabrizioi.

Kilburn (1985) proposed a subgeneric arrangement

for the South African .species of Epitoniidae. He did not

consider, however, that radular morpholog)' could pro-

\ide rehable taxonomic characters to his arrangement. If

we followed Kilbuni's classification, E. fabrizioi could be

allocated in the subgenus Hirioscala Monterosato, 1890.

However, E. fabrizioi does nf)t have corcjnate lamellae

and a duplicate peristome that are characteristic of that

subgenus.

Including E. fabrizioi. there are 5 named species of

Epitoniidae living in the Magellanic prn\ince. Keen

(1971) illustrated 76 Epitoniidae from the tropical east-

em Pacific; Kilburn (1985) cited 80 species from south-

em Africa and Mozamlii(|ue; Rios (1994) recorded 30

Table 3. Measurements of protoconchs of Epitonium species in relation to the adult size (in mm'

Epitoniiiiii speci

Protoconcl

Whorls

Aduli

Protoconch size

leirgth-width

Source

19941)

Page 68

THE NAUTILUS. Vol. 112. No. 2

from Brazil, while Diaz-Merlano and Pu\ana-Hegedii,s

(1994* listed 22 from the Colombian Carihliean. Based

on pubhshed records ot the known distribution of Epi-

toniidae this famiK' is more speciose in tropical than in

temperate and cold regions.

ACKNOWLEDGMENTS

Special thanks are due to f. Mermoz, wht) pr()\ided

specimens of Epitoninin from Punta Loma. Yves Finet,

Museum d'Histoire naturelle de Geneve, Switzerland,

kindly sent photographs of the paralectotype of Epiton-

ium ^corin'ffinum. M. G. Harasewvch, Nation;il Museum

of Natural HistoiT, Smithsonian Institution, gave good

ad\ace on earlv stages of the manuscript. Two rcNdewers

improved considerablv the original manuscript. This

work was made possible b\' an external fellowship grant-

ed bv the Consejo Nacional de Investigaciones Cienti-

ficas'y Tecnicas (CONICET), Argentina, to G. P and a

grant from Funilacion Antorchas, Argentina to P. P.

LITERATURE CITED

Bell, ]. L, 1985. Lanai growth and metanioqihosis of a pro-

sobranch gastropod associated with a solitary coral. Pro-

ceedings of the Fittli Inteniational Coral Reef Congress

5:159-163.

Bouchet. P and A. Waren, A. 1986. Re\ision of the Northeast

Atlantic bathyal and abyssal Aclididae, Euiimidae, Epito-

niidae (Mollusca, Gastropoda), Boilettino Malacologico,

Supplemento 2:498-523.

Clench, W. J. and R. D. Turner. 1951. The genus Epitonium

in the western Atlantic (Part 1). Johnsonia 2(301:249-288.

Clench, W. J. and R. D. Turner. 1952. The genus Epitonium.

(Part II), Depressisc/ila. Ci/lindriscala. Ni/stiella and So-

hiti.sailii in the western Atlantic. Johnsonia 2(31 ):289-35fi.

Dfaz-Merlano, J. VI. and M. Pnvana-Hegediis. 1994. Molusc<w

del Caribe Colombiano. Un catalogo ilustrado. Fmidacion

Natura, Bogota, 291 pp., 78 pis.

DuShane, H. 1974. The Panamic-Galapagan Epitoniidae. The

Wliger 12(Supplenient):l-S4.

DuShane, H 1979 The faniih Epitoniidae (Mollusca: Gas-

tropoda) in the northeastern Pacific. The Wliger 22(2):

91- 1.34.

|ung, P. 1986. Neogene paleontology in the northern Domin-

ican Republic. 2. The genus Strombina (Gastropoda: Col-

umbellidae). Bulletin of American Paleontology 324:1^2.

Keen, M. 1971. Seashells of Tropical West American, Second

Edition. Stanford University' Press, Stanford, 626 pp.

Kilbuni, R. N. 1985. The fiunily Epitoniidae (Mollusca: Gas-

tropoda) in southern Africa and Mozambique. Annals of

the Natal Museum 27(l):239-337.

Pastorino, G. 1995. Moluscos costeros Recieutes de Puerto Pi-

ramide, Chubut, Argentina. Academia Nacional de Cien-

cias, Cordoba, Miscelanea, 93:1-30.

Pastorino, G. and P. Penchaszadeh . In press. On the egg cap-

sules of Epitonium ^eorgcttinum (Kiener, 1839) (Gastrop-

oda: Epitoniidae) from Patagonian shalloyv waters. The

\eliger

Rios, E. C. 1994. Seashells of Brazil. Editora da Fimdavao

Universidade do Rio Grande. Rio Grande, 328 pp.

Robertson, R. 1981. Epitonium miUccostatum and Cornllio-

phila clathrata: two prosobranch gastropods symbiotic

yvith Indo-Pacific Pahjthoa (Coelenterata: Zoanthidae).

Pacific Science 34(1): 1-17.

Robertson, R. 1983a. Observations on the life historv of the

yventletrap Epitonium echinaticostum in the Bahamas.

The Nautilus 97(3):9S-103.

Robertson, R. 1983b Obsenations on the life historv of the

wentletrap Epitonium albidum in the \\'est Indies. .Amer-

ican Malacological Bulletin 1(1): 1-12.

Robertson, R. 1994a. Protoconch size variation along depth

gradients in a planktotrophic Epitonium. The Nautilus

107(4):107-112.

Robertson, R. 1994b. Tyvo neyv tropical yvesteni Atlantic spe-

cies of Epitonium. yvith notes on similar global species and

natural histoiy. The Nautilus 107(3):81-93.

Scarabino, \". 1977. Moluscos del Golfo San Matias (Pro\incia

de Ri'o Negro, Repi'iblica Argentina. Inventario v Claves

para su identificacion. Conumicaciones de la Sociedad

Malacologica del Unignav 4(31-321:177-285.

Strebel, H. 1905. Beitriige zur Keimtnis der Mollusken-fauna

der Magalhaen-Proyinz. 3. Zoologischen Jahrbiichem. Ab-

teilung fiir Systematik, Geographic, und Biologic der Ti-

ere 22(6):575-666.

THE NAUTILUS 112(2):69-72, 1998

Paae 69

Two New Species o{ Pchploma (Bivalvia: Anomalodesmata:

Periplomatidae) from the Southern Caribbean

Nestor E. Ardila

Institiito de Iinestigaciones Marinas v

Costeras, INN'EMAR

A.A. 1016

Santa Marta. COLOMBIA

[email protected]

Juan M. Diaz

Institute) lie Investigaciones Marinas y

Costeras, INVJEMAR

A.A. 1016

Santa Marta, COLOMBIA

[email protected]. CO

ABSTRACT

Periploiim coseli new species and P. sutuiamiiilliacusis new

species are described from near Santa Marta, Caribbean coast

of Colombia. The overall shape of the shell, the outline ot the

pallial sinus and the shape and/or orientation of the resiliter

are the main features distinguishing the two new titxa from

each otlier, and from other congeneric species trom the west-

ern Atlantic, eastern Atlantic, or eastern Pacific regions.

Kei/ uanls: Santa Marta, Colombia, Caribbean Sea, western

Atlantic, eastern Pacific, eastern Atlantic, Thracioidea.

INTRODUCTION

The familv PeriploniatRlae (Anomalodesmata: Thracioi-

dea) is a relatively small taxon consisting of about 30

living recognized species arranged in 7 genera (see

Rosewater, 1968; Bernard, 1989). The group is best rep-

resented in the eastern Pacific and the western Atlantic

species were fouiu! so

sp.

regions. Notwithstanding, onl'

far in the Caribbean Sea, both belonging to the genus

Pcriploina (s.s.) Schumacher, 1817: P. inorfiaritacciiin

(Lamarck, 1801) and P. coqitcttac Altena, 1968. In ad-

dition to these 2 ta-xa, 2 other unnamed species of the

genus have been collected along the C>'aribbean coast of

Colombia. Cosel (1978, 1986) and Diaz (1985, 1990),

who studied the molluscan fauna from ibfferent localities

of the Colombian coast, first noticed the occurrence of

these unnamed Pcriploina. Finther material of one of

these species has become availalile to the authors in re-

cent vears through intensive collecting near Santa Marta.

Thcv are herewith described as new species. Institution-

al abbreviations are: BMSM, The Baile\-Matthews Shell

Museum, Sanibel, Florida; INVEMAR MOL, Collection

of Mollusks, Instituto de Investigaciones Marinas y Cos-

teras, Santa Marta, Colombia; SMF, Senckenberg Mu-

seum Frankfurt, Cermanv.

SYSTEM ATICS

Class Bivahia Linnaeus, 17.58

Subclass Anomalodesmata Dall, 1889

Superfamily Thracioidea E.A. Smith, 1885

Familv Periplomatidae Dall, 1895

Genus Paiploma Schumacher, 1817

Periploma [Pcriploina) coseli new species

(Figures I, 2, 5)

Periploma spec. — Cosel, 1978:161. pi, 4, figs. 12,1.3.

Pcriploina spec, II. — Diaz, 198.5:84, erroneously figured in

pi. 12, fig.l.

Periploma sp".— Cosel, 1986: 199, fig. 11.3

Periploma sp, 1.— Diaz & Puyana, 1994:104, pl.:31, fig. :30.3.

Description: Shell merhum-sized (length up to 29.3

mm), subcircular-oval, ven- thin and fragile, moderately

convex, antero-ventral margin evenlv circular posterior

margin scjmewliat projected upward and subtnmcate.

Subecjuilateral, anterior end slightly larger, beaks with

characteristic tran.sversal slit. Outer surface white, with

irregular growth lines and numerous, extremely fine, ir-

regular radial striae more apparent in the central area.

Periostracum diity gray, with comarginal rows of micro-

scopic granules. Inner surface weaklv nacreous, smooth

but with weak impressions of growth lines. Palfial sinus

broad and short, reaching to about 1/3 of shell length.

Ligament internal; resiliter conspicuous, spoon-shaped,

thrected vertically toward center of valve and weakly re-

inforced at its basis b\ cui-vetl buttress pointing to pos-

terior muscle scar

Type material: Ilolohpe, SMF 311857, complete

shell (left valve slightly broken), 29.3x25.2 mm

(lengthxheight), trawleil h\ RA' Ancdn at tspe locality,

C.P Arango coll. September 1995. Paratvpes: INVE-

MAR MOL-1151, 1 complete shell (left valve fractured

near midline), 24.8X20.0 mm, Golfo de Salamanca, Co-

lombia, ir07'N, 74°20'W, trawled hv WW Ancdn, 59 m,

muddv sand, C.P Arango coll. September 1995; INVE-

MAR MOL 1152, 1 complete shell (left valve damaged),

10.5X8.5 mm and BMSM 2361, 10.9X8.8 mm, 1 com-

plete shell (left valve damaged), off mouth of Toribio

Page 70

THE NAUTILUS. Vol. 112. No. 2

^

/

Figures 1—1. 1-2, Pcriploma roscli. Hol()t)pe, riglit \al\e, 29.3 mm length, 25.2 mm lielglit, 3—4, Frriphtuia sanrtmimiihncnsis.

HoloUpf. left \alve. 21.9 mm length, 17.5 nnii height.

river. ir04'N, 74°15'W, taken by bottom grab, 6 in.

mud, N. Ardila coll. Octolier 1996;' BMSM 23fi(). 1 com-

plete .shell, 16.8X14.2 mm. off mouth of Toiihio Ri\er,

1I°()4'N, 74°15'\V, bottom grab, fi m. imid. N. Ardila

coll. Jamiar)' 1998.

Type locality: Colfo dc .Salamanca, (>'olond>ia.

ini'N, 74°15'W, 59 m, muddy .sand.

Etymology: Named altci- I-tiido xon Cose), who fiist

noticed the existence ol this species in his comprehen-

sive study of the molhiscan fauna of the ('aril)bean coast

of Colombia.

Remarks: PcripUiinu roseli new species is similar to

F. discus Stearns, 1890, from the eastern Pacific, and

to P. camerunensis Cosel, 1995, from off west Africa.

The postero-dorsal margin of P. discus is almost

straiglit (somev\hat projected dorsalK' in Pcoscli). and

its posterior margin more angulale ibroadK rounded

in the new species) (compare figures 5 and 7). Pcri-

ploma caincnincnsis exhibits a somewhat more elon-

gatetl shell, with the umbones situated posteriorh' to

the vertical midline. In addition, the resilifer in P. ca-

incnincnsis is projecteil obliquelv. in relation to the

lunge line and toward the antero-\entral margin,

whereas it is almost perpendicular in relation to the

hinge line in P coscli (compare Figures 5 and 8).

These 3 species may be regarded as members of a

group ol similar taxa that show Recent disjunct distri-

bution. Isolation ol eastern Pacific and eastern Atlan-

tic anci'Stral populations could ha\e occurred respec-

ti\('K' alter establishment of a continuous Isthmus of

Panama and a lull-Hedged mid-Atlantic oceanic bar-

rier. Absence ol planktotrophie de\elopment in the

genus Pciiploina (see (ioodsell ct al. 198.3) ma\' ha\e

laeilitated isolation and subse(juent speciation.

P CDipicHiic .\ltena. 19tiS Irom the iiortlu'in and

N. E. Ardila and J. M. Diaz, 1998

Page 71

Figures 5-11. Diagrams ot'tlie inner surfaces of left \;il\es of several species o( Pciiploinu 5. f ((iscli Holohpe, 6. P snmiaiiuiiihiwnsis:.

Holohpe^ 7. P disriis Steams, 1890. retlrawii from Keen (lOThfig. 750). 8. P caiiirnincmis Cosel. 1995. Holohpe reth-awii from Cosel

(1995:fig. 144). 9. P coqucttac Altena. 1967. Holohpe redrawn from Altena (1967:fig-147a). 10. P filnnUiscuhnn ,Sowerb>, 18:34, retb'awi

from Keen (1971;fig, 7.52). 11. P hii^aiHUa Olsson, 1961. Holohpe reclrawii from cils.son (196!:pl. 82, fig. .5h).

northeastern coa.st.s of South America (off Suriname

and Colombia), i.s another vaguely similar specie.s, hut

it has definitely a more elongated and inequilateral

shell (compare Figures 5 and 9).

P. coscli seems to he \\idel\' tlistrihuted ;dong the Carih-

bean coast of Cok)mhia. Cosel (1986) recorded diis species

as "Pcriploum spec." from off Punta Broijueles (approxi-

mateh- 9°2()'N, 76°1.5'W) and die Santa Mmta Baw and

Diaz (1990) found it iJ.so ;ls north as Poitete Ba\- (12°18'N.

71°55'W)

Page 72

THE NAUTILUS, Vol. 112. No. 2

Periplonta (Pcriploina) sanctanuuihacims new specie.s

{Figures 3, 4, 6)

Periplomn spec. I. — Di'az, 1985:84, pi. 12, fig. 2.

Periplomti .sp. 2.— Diaz & Puvana, 1994:104, pl.32, fig. .304.

Description: Shell niedimn-.sized (length to 22 mm),

.semicircular-ov;il, equilateral, convex, very fragile; an-

tero-ventral margin semicircular; posterior margin sub-

truncate, postero-dorsal margin strtiight. Beaks with a

characteristic tran.s\ersal slit. Outer surface milk\' white,

with irregular growth lines, some of which appear as

shallow grooves under magnification. Radial ridge nms

from beaks to postero-ventral margin. Periostracum vel-

lowish-gray, with numerous microscopic granules which

are arranged comarginalK- and are more aliuntlant within

growth lines and posterior slope. Inner surface dirts-

white and weakl\' nacreous, showing impressions of ir-

regularities on outer surface. Pallial sinus broadK trian-

gular in outline and rather deep, reaching to about Vi of

shell length. Ligament internal, in well-defined, scjuarish

resililer. Resiliter oblique in relation to hinge line, ori-

ented toward ventral margin.

Type material: Holot>pe, SMF 311S58, 21.9X17.5

mm (lengthxheight), 1 left valve, from t\pe locality,

SCUBA, J.M. Diaz coll. Febman 1983; Parat\pe, IN-

VEMAR MOL 1153, 13.4X11.3 mm, 1 right valve (bro-

ken in two pieces after measurements), from type local-

itv-, SCUBA, J.M. Diaz coll. Februaiy 1983.

T^pe localit>': Bahia Nenguange. Ta\Tona Natural Na-

tional Park. Colombia, ir20'N'," 74°()5'VV, 7 m, coarse

coralline sand with broken shells.

Etymology: The species is named after the citv oi

Santa Marta. near which the Tavrona National Natural

Park, the type locality, is located.

Remarks: The shell of Periplomn saiictamaiihaciisi.s

differs from that of P. coseli in being smaller and more

inflated. In addition, the outline of the antero-\entral

margin is gentlv rounded, has a strmght postero-dorsal

margin that does is not projected dorsallv. and a railial

ridge nnming obliquel)' along the postero-ventral slope.

Additional differences between these species do exist in

the outline of the pallial sinus and the shape of the re-

silifer (compare Figures 5 and 6).

Pcriploma planiiisciilutn Sowerbv. 1834. from the

eastern Pacific, has a larger and more elongated shell

with a short, rounded pallial sinus (broadly triangular in

P. saiirtaiiuu-tliaciiMis) (compare Figures fi and 10). Pir-

iploina laoarlilla Olsson, 1961, from the Pacific coast of

Panama, is also more elongated, and exhibits a rather

pointed posterior margin and a short and narrow pallial

sinus (compare Figures 6 and 1 1 ).

P. stmrtaiiwiiluiciisis new species is so iai' known oni\

from the t>pc locality.

ACKNOWLEDGMENTS

We wish to express our gratitude to Claudia P. Arango

for providing the type material of Pcriploma coseli new

species, and to Dr. Eugene V'. Coan for his kind help

with the literature search. The personnel and facilities

of the Reference Collection at INVEMAR, Santa Marta,

made possible the assemblv and improvement of pho-

tographs and drawings. Two anonvinous reviewers con-

tributed to the improvement of the maiuiscript. We ac-

knowledge the economic support of COLCIENCIAS —

BID (grant No. 2105-13-079-97). This is contribution

No. 601 of INVEMAR.

LITERATURE CITED

Alteiia, C. O. van Regteren. 1968. The holocene and recent

marine bivalve Mollusca of Surinam. Studies on the Fauna

of Suriname and other Guyanas 10(42):15.3-179.

Bernard. F.R. 1989. Living Periplomatidae of the Pacific and

Indo-Pacific regions, \emis 48(1): 1-11.

Cosel, R. \on. 1978, Die .Mollusken der Cieuaga Grande de

Santa Marta (Kolumhien) uud ihre Lebensgemeuischaften

im Wechsel der Jahreszeiten. Ph.D. Dissertation. Univer-

sity of Giessen, West Germany. 348 pp.

Cosel, R. von. 1986. Moluscos de la region de la Cienaga Gran-

de de Santa Marta (costa del Caribe de Colombia). Anales

del Institute) de Investigaciones Marinas de Punta de Be-

ti'n (15-16):79-^370.

Cosel, R. von. 1995. Fiftv-one new species of marine bivalves

from tropical West Africa. Ibenis 13(1): 1-1 15.

Diaz, J.M. 1985. Mollusken iind ihre Gemeinschaften in der

Bahi'a de Nenguange (Karibik. Kolumbieni. Ph.D. Disser-

tation, Uui\'ersitv of Giessen, West Germany. 305 pp., 29

pis.

I^iaz. |,M. 1990. Malacofauna subfosil v reciente de la Bahi'a

tie Portete, Caribe colombiauo, con iiotas sobre algunos

fosiles del Terciaiio. Boletin Ecotropica 23:1-22.

Diaz, J. M. y M. Puvaua. 1994. Moluscos del Caribe Colom-

biano, un catalogo ilustrado. COLCIENCI.^S-Fuudacion

Natura-IN\'EMAR. Santafe de Bogota. 291 pp., 74 pis.

Goodsell, J.G.. R A, Lutz. M Castagna and J. Kraeuter 198.3.

Nonplanktotropliic development of two species of conti-

ueiitnl shell bi\alves Journal of Shellfish Research 3(1):

91

Keen, M. 1971. Sea shells of Tropical West .Vmeiica. Stanford

University Press, Staufoixl. 1074 pp.

Olssou. A. A. 1961. Mollusks of the eastern tropical Pacific,

particularly from the soutlieni halt of the Pauamic-Pacific

laiinal province (Panama to Peru). Pananiic-Pacific Pele-

cvpoda. Paleontological Research Institution, Ithaca. 574

pp., 86 pis,

Rosewater |. 1968. Notes on Periplomatidae iPelecvpoda: .\ii-

omalodesmata), vritli a geographical checklist. Annual Re-

]icirt 111 llic .\mi-ncaii .Malacologic.il t'niou tor 1968: 37-

39.

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TH EfyNAUTI LUS

CONTENTS

Vohimi' 112, Number 3

October 15, 1998

ISSN 0028-1344

Noah Feinstein

Stephen D. Cairns

Learning from the collector: a .sur\e\' of azooxanthellate

corals affixed hx Xcnopluva (Gastropoda: Xenophoridae),

with an anaK'sis and discussion of attachment patterns 73

Shell microstructure of mvtilids ( Bivalvia) from deep-sea

hvdrothermal vent and cold-water sulfide/methane seep

environments 84

Molluscan taxa and bibliography of Henry van der Schalie 90

A new Favdiiia (Miurxiclla) from the Panamic Province

(Gastropoda: Muricidae) and designation of a lectotvpe for

F. (M.) cxiffLia (Broderip, 1833) . ." ' 95

Michael J. Kennish

Antonieto S. Tan

Richard A. Lutz

Mark E. Gordon

Carole M. Hertz

Barbara W. Myers

THE NAUTILUS 112(3):73-83, 1998

Page 73

Learning from the Collector: A Sui-vey of Azooxanthellate

Corals Affixed by Xenopliora (Gastropoda: Xenophoridae), with

an Analysis and Discussion of Attachment Patterns

Noah Feinstein

Hanard Lliiiveisih'

Cambridge, MA 02138, USA

and

National Musenni of Natural Historv

Research Training Program

Smithsonian Institution

Washington, DC 20560 USA

Stephen D. Cairns'

Department of Invertebrate Zoolog\'

National Museum ot Natural Historv

Smith.sonian Institution

Washington, DC 20560 USA

[email protected]

ABSTRACT

AU species of die gastropod familv Xenophoridae affix foreign ob-

jects to the upper shell surface. Affixed objects ma\' include bi-

valve shells, smaller gastropod shells, sheO fragments, and conJ

skeletons, as well as a wide array of inorganic materiid. In deep

water, Xcnophom im\ ser\'e as useful prox\' collectors ot bentluc

organisms. Since 1S42, coral skeletons have been noted among

the attached objects, but this association has never been studied

in detail. Tliis paper survevs 227 Xenophora shells, comprising 8

species from 69 stations, for affixed azooxandieUate corals. Fi\e

hundred and eightv-one coralla were foimd, representing 74 coral

species, 2 of wliich remain undescribed. Twenh-tour of the affixed

coralla were alive at tlie time die Xcnophom hosts were collected;

Xenophora not onlv collect live conils, but corals can remain iJive

long after being iiffixed. Corals were found at 6 sites where the\'

had pre\iousK' been unknown, iind the geographic riuiges of 29

species of coral were exjiiuided as a result of specimens found on

Xenophora

This paper luis hvo sections. In the first. quantitati\e obsena-

tions were made on die orientation of affixed conils; statistical

analvsis of these obsenations revealed non-random patterns of

attachment, based on orientation of bodi the long axis of die coral

iuid die coral cahce. Qiuditative obsenations suggest diat species

of Xenophora favor corals ot paiticnku- shapes. In tlie second sec-

tion, die speculations ot previous authors regiii-ding the ecological

basis for attachment behavior are summarized and new dieories

are discussed. Four of diese explanations suggest defensive ad-

aptations, and die remaining 3 are functional support adaptations.

Three of these hvpodieses (annor, tactile camouflage, and snow-

shoeing) are proposed tor the first time in tlus paper

Key iLords: Carrier shells, Mollusca, gastropods, aliermat\pic

corals.

INTRODUCTION

According to Ponder (1983), the nionotxpic gastropod

familv Xenophoridae contains 25 Recent species of AV/i-

' Corresponding author.

ophorei. marine gastropods that inhabit the continental

shell and slope regions of tropical and temperate oceans.

Xenophora have dra\Mi the attention of naturalists and

svstematists since the earK' 1800s. This was caused pri-

mariK' because of a peculiar behavioral pattern: all spe-

cies affix objects to the upper surface of the shell

throughout some or all of its growth (see Shank, 1969

for a detailed description of the affixing procedure in X.

conchi/Iiophora (Bom, 1780)). Although objects are onlv

affixed at the growing edge of the whorl, older attach-

ments remain on the perimeter of earlier whorls as the

shell increases in size. Older attachments often become

secondariK' affixed to the \oimger whorl that is ft)rmed

beneath them. One of the traits used to distinguish spe-

cies of Xenophora is the degree to which the shell sur-

face is obscured by attachments (Ponder, 1983).

A great diversih' of material has been found affixed to

Xenophora: among the objects we obserxed in the

course of this studv were coral skeletons, bivalve and

other niollusk shells, brachiopod shells, echinoderm

spines and skeletal fragments, brvozoans, sponges,

sharks' teeth, and a wide assortment of inorganic frag-

ments.

Although corals tvpicalK' comprise less than 10% of

the affixed objects, their presence on Xenophora shells

was first recorded a centurv and a half ago (Reeve,

1842). Pourtales (1871) was the first to report a deep-

sea (azooxanthellate) coral, Can/oplii/Uia cornufonnis

Pourtales, affixed to a Xenophora shell, collected from

the Straits of Florida at 433—454 m. In this case, the

coral remained alive after attachment. Morton (1958)

Listed living, solitarv flabellid corals affixed to A', corru-

gata (Reeve, 1843) from New Zealand, and Kawase

(1996) identified 4 species of azooxanthellate corals on

X. paUidula (Reeve, 1842) from Japanese waters. Cairns

(in press) reported 19 azooxanthellate species affi.xed to

xenopliorid shells collected from the slope region of

Page 74

THE NAUTILUS. Vol. 112. No. 3

Vanuatu ( = New Hebrides) (Figure 6), prompting the

research presented here. In summary, there have been

sever;il pubUshed observations of deep-water corals af-

fixed to Xcnopliora, but no comprehensive review of the

association.

Xeiiophora are not highly mobile (Berg, 1975). It is

therefore possible for one to learn something about the

benthic fauna of a particular region b\' examining the

local Xcnophi)ra. Although some species are found only

in shiillow water, most Xcnophora inhabit regions well

below the penetration depth of photosyntheticalK' active

radiation, and specimens have been dredged from

depths exceeding lOOO m (Ponder, 1983). Because rel-

ativelv little is known about the deep-water bentlios.

deep-dwelling Xcnophora are potentialK' useful as proxT

collectors.

In the process of examining Xcnophora shells for af-

fixed conJs, we noticed that there appeared to be some

regularity in orientation of affixed objects with respect

to the shell. These apparent trends, in the conte.xt of the

pre\iousK documented observation that Xcnophora at-

tach lamellibranch valves with the concave side up (Pon-

der, 1983; Morton, 1958; Linsle)- and Yochelson, 1973;

Shank, 1969). led us to look for statistical patterns in the

manner and orientation of coral attachment. The results

of that analysis are presented in this report.

MATERIAL AND METHODS

Two hundred and twentv-seven coral-bearing A'riio/j/iora

shells were examined: 145 of these are from the collec-

tions of the National Museum of Natural History

(USNM), Washington, DC; 42 from the Museum na-

tional d'Histoire naturelle (MNHN), Paris; 38 from the

Delaware Museum of Natural Histon, (DMNH), Wil-

mington; and 2 from the Museum of Comparative Zo-

ology' (MCZ), Cambridge. Appendix 1 lists the stations

at which coral-bearing Xcnophora were collected, the

station data, and the coral species collected at those sta-

tions. The study material includes shells collected

throughout the Indo-West Pacific, Hawaiian Islands, and

the Gulf of Guinea. Eight coral-bearing species of Xcn-

ophora were examined, a large majorih of the speci-

mens (166/227, including all shells borrowed from the

MNHN and DMNH) belonging to the species X. palli-

dula. The other 7 species, in order of abundance were:

X. Japonica Kuroda and Habe, 1971, X pcroniana kon-

cloi Ponder. 1983, X cornifSfita, X. ccrca (Ree\e, 1845),

X. ncozcahmka Snter, X. cti.spa (KiJuig, 1831), and X.

granulosa Ponder, 1983. The 145 USNM specimens in-

cluded all coral-bearing shells in the USNM collections,

with the exception of those belonging to X. conchi/lio-

phora, which is tyjiicallv found in shallow water (Ponder.

1983), and certain lots of X. pcroniana and X. japonica.

which preliminarv' examination showed to be redimdant

in terms of sites sampled and corals collected.

When possible, corals were identified to tiic species

level; when onlv a coral fragment or badlv eroded or

damaged coralluiu was present, it was identified to the

Figure 1. Diagram of the apical view ot a Xcnophora shell,

illustrating the various tvpes of orientatious in which a solitary

coral may be affixed: radial (up, down), lateral (out, in), and

svinmetrical (facing up, facing down). Note: Number of apical

whorls underrepresented in figure.

lowest possible taxonomic level (tvpically genus). Un-

described species were designated bv a letter. If a well-

presened but unidentifiable phenotvpe appeared re-

peatedly, it was given the designation n. sp. Anv corallum

that contained remnants of dried coral tissue or was e.x-

ceptionallv well preserved was considered to have been

alive when the Xcnopliora was collected.

Everv coral found was classified as a primarv or sec-

ondary attachment. Primary attachments were those that

were embedded in the shell and had clearlv been affixed

bv the Xcnophora: secondarv attachments included

those conds growing on a substrate which was snbse-

(juentlv affi.xed to the Xcnophora and those that settled

upon the surface through no action of the Xcnophora.

The length of the long a.xis (maxinuun dimension, re-

gardless of the orientation of nioqihological features) of

each coral skeleton was recorded. The orientation of the

long axis was classified as radial, meaning roughly per-

pendicular to the arc of the whorl: lateral, meaning

roughlv parallel to the arc of the whorl: or svmmetrical,

meaning that all axes of the coral v\ere e<ju;d. The ori-

entation of the calice (Figure 1) was also noted as being:

up or down (a subset of radial orientation), in or out (a

subset of lateral orientation) or facing up or facing down

(a subset of svnuuetrical orientation). We applied statis-

tical hvpothesis testing (chi-s(juare test) to the orienta-

tion data, using as our null hvpothesis the assumption

that each hpe of orientation would have the same

chance of occurring.

We measured each Xcnojihora shell across the shell

base, and took the maximum diameter as a proxy for

shell size. For selecti'd X paUidula. we measured the

diameter of each whorl l)otli with ;md v\ithont attach-

ments. Using a circular approximation lor area, we cal-

culated the ratio of shell basal area (rachus measured,

not including attachments) to expanded shell basal area

N. Feinstein and S. D. Cairns, 1998

Page 75

Table 1. Nuiiierical disfribiition of azooxanthellate scleracti-

nian genera, species, and indi\iduals per families of eonJ-bear-

ing Xfitophoni.

(radius measured, including attachments) for each shell

whorl. These ratios were analvTied for statistical trends:

Pearson product-moment correlation coefficients and

the respective tests of significance were computed with

the aid of Staf(.iew SE statisticiil sofWare. For selected

Xenophora solans (Linnaeus), we countetl the number

of spines per whorl and analyzed these data for an\

trends, again using Stat\iew SE.

Lists of coral species previcnisK' reported were avail-

able for 2 series of stations {Allmtmss Expedition, Caims

and Zibrowius (1997); MUSORSTOM S E.\i:)edition

(Calms, in press)). We compared the list for each station

with the corals that we fountl affixed to tiie Xiiiophora

at that site, and noted an\' instances of affixed corals not

previously- knowii from that location, as well as which

corals were potentialh- a\ailable for fixation but were not

affixed.

RESULTS

A total of 581 azooxanthellate coralla were found affixed

to Xctuiphoni shells, 511 of which were identifiable to

genus, and 411 to species. The affixed coral fauna rep-

resents a diverse taxonomic distribution, including spe-

cies from all 5 suborders in 11 famifies, 29 genera, and

74 species. The onK zoo.xanthellate coral species en-

countered was Fuitf^ia (Ci/chisciis) laufihani, found on

X. pcroniaiKi trom the Hawaiian Lslands. Table 1 shows

the numerical distribution of genera, species, and indi-

viduals among families of corals and the Xenophora spe-

cies tliat affixed them. A taxonomic list of all azoo.xan-

tlieliate coral species is included in Appendix 2.

SLxts-seven of the 74 species were recognizable as de-

scribed species of Scleractinia. Of the remaining 7. 5

(designated as sp. A or sp. B) are represented bv single

specimens and may be aberrant examples of described

species. The remaining 2 are considered undescribed

species. One of the undescribed species had previousK-

been observed in dredged samples from the same re-

gion, antl illustrated 1)\- Stolarski (1996) as "Garchneri-

idae gen. n." Sexen well-preserwd specimens of this tax-

on were found on 3 X. palliditia shells from MUSOR-

STOM stations 1008 (Figures 8-9). 1087, and 1088 off

the coast of Vanuatu ( = New Hebrides). The second un-

described species belongs to the genus Placotrocltidcs

and is represented b\' 4 specimens affixed to 2 X. pcr-

oniana shells from 2 stations in the Hawaiian Islands.

The genus Placotrochidcs had not pre\iouslv been col-

lected near the Hawaiian Islands, and no other truncate

Habellids (transverseK' dividing members of the family

Flabellidae) are known from the area. Even though none

of the specimens of Placotrochidcs n. sp. is particularly

well preserved (Figure 10), thev are clearly not members

of other described Placotrochidcs species.

Large specimens of Truiicatoflabclhim fiardincri

Caims and Keller proved to be new size records for that

species. The previously listed maximum length for T.

f^ardiiicii vyas 18.7 mm. We discovered 8 specimens larg-

er than that, the largest (USNM M818232) measuring

27.5 mm from base to calice (Figure 7).

Xenophora frequently collected corals that were not

previously kTiown from their source station: in 74 in-

stances, a coral species found affixed to Xenophora from

a given site had not previously been found at that site.

These discoveries expanded the known geographic rang-

es of 29 coral species. Coral-bearing Xenophora were

also found at 6 sites (Albatross 5121, 5394, 5395, 5416,

5661; MUSORSTOM 8-1008) where dredging had not

previousK produced aii)' coriils at all.

There are definite patterns in the orientation of af-

fi.xed corals. Seventv-two percent, 359 of 498 non-sym-

metric coralla, were positioned with the long axis radial

rather than lateral (Figure I); occurrence of such a high

percentage of radiallv' oriented coralla is extremely un-

likely (p<C0.01) if Xenophora affix corals at random,

without regard to orientation of the long axis. Calice ori-

entation was also cfistinctly non-random. The down and

out orientations were encountered significantly more

times than it would have been if Xenophora ignored cal-

ice orientation (p<CO.OI), whereas the up and in orien-

tations were encountered significantly fewer times. Cer-

tain shape classes were attached in a specific manner:

conical corals were typically oriented radially witli the

cahce in the down position (Figures 1, 4), whereas Ha-

bellids were tvpicallv oriented laterally with the calice in

the out position (Figures 1, 3, 5). Symmetrical flat corals

were found (with onlv' 1 exception among 92 Hat corals)

with the calice facing up (i.e., with the base cemented

to the shell).

Early observations led us to hypothesize that Xeno-

phora select objects preferentially based on shape. The

shells from MNHN frequently collected BoKnicofroc/ius

stcllulatus Caims, a small, coin-shaped, spinose coral,

but none of these moiphological traits appeared in sig-

nificant numbers at other regions or in other collections.

By contrast, we noted a high frecjuencv of attachment

for both Habellate and comute coralla at widely distrib-

uted sites. It is difficult, however, to test these frecjuen-

Page 76

THE NAUTILUS. Vol. 112. No. 3

Figure 2. XcnojiliDrn solari.s

off Zululand (DMNH 20120)

roiii MaqiK'il.i B.i\. I'liilippiiies {DMNH 5173): apical \iew, X ().S2

apical and lateral \ie\vs showiiii; liw Tnincalofliilnlluiu i^anlini'ii

piillidiild Iroiii

iiul out latenil

Figures 3, 5. .\

ilfi.\cd in the in

positions, X 0.7.5, X 0.5<S, res])ectiveK-. Figure 4. .V, pallidiilii iroin Sulu Sea (\\'49S)(US.\M MS0776.3), showing two li\e coralla

of Trochoq/athns cooperi affixed in the radial (dovvii) position. Figure 6. Stereo apical view of X. pallklitln from MUSORSTOM

8-963 (MNHN), illustrating a heavily "encrusted" shell with 6 species of solitan,- coral. Figure 7. Largest knovvii (length 27.5 mm)

corallum of Tntncatofiahcllum nardineri affixed to X. paUiduhi from off Diu-han (V\'320) (USNM M,S1S232), X 1.4. Figures 8-9.

Coralla of three live "Gardineriidae. n. gen. n. sp." affi.xed to X pallidula from MUSORSTOM S-IOOS (MNHN). figure 9 showing

rejuvenescence from presumed growth trauma scar following (ixation, X 2.8, X 6.5. respectivelv. Figure 10. Corallum of Placo-

trochidea n. sp. affixed to X. pcroniann from David Stiirr ji»d<in T(:.32-2 (USNM M807659). x'T.I.

N. Feinstein and S. D. Caims, 1998

Page 77

cies for statistical significance, as eacli Xeiioplioia has a

different range of shapes axailahle, and it would he nec-

essarv' to know if the frequencies of attachment were

significantK' different from the frequencies at which the

various shapes occur unattaclied. The latter figures are

not available due to lack of data.

A small number of affixed corals (24 specimens, ap-

proximately 4%) were alive at the time their Xenophora

hosts were collected. Only 3 of these uere secondare

attachments, allowing us to confirm that Xenophora ac-

tiveK' affi.x living corals, if onlv at a low trequencv. Living

corals were often affixed to earlier whorls, and tspicallv

bore a visible growth-trauma scar on the theca (outer

surface). The presence of live-at-collection corals on ear-

lier whorls indicates that corals mav survive after attacli-

ment, and a growth-trauma scar (Figure 9) suggests that

they also continue to grow. Relative frequencies of "live"

and "dead" corals on Xenophora shells cannot, unfortu-

natelv, be compared with relative frequencies of living

corals and coral skeletons in situ because the latter fig-

ures are unknowTi. It is therefore impossible at this time

to state whether Xenophora preferentiallv selects or

avoids living corals, although the small number of living

corals suggests that it is imlikelv that live specimens are

preferred.

DISCUSSION

Descriptive D.at.v

The examination of 227 Xenophora shells brought to

light 2 undescribed species of coral, increased the size

range for one described species, and extended the geo-

graphic ranges for 29. Had the shells been examined

immediatelv upon collection, the results would have

been even more impressive: 215 of the 581 coralla were

found on Xenophora collected by the Albatross cruises

before 1910, at which time 24 of the collected species

(32%) were undescribed.

Admittedlv, Xenophora-ha.sed samphng has limita-

tions. There was not a single case in which we found

every coral knowai to exist at a given site (i.e., at a station

where coral was previouslv recorded) affixed to Xeno-

phora shells. Altogether, 41 species of coral known to

co-occur with Xenophora were not affixed bv' the gastro-

pod. This mav be an artifact of the small number of

Xenophora hpicallv kiiowii from each site, but, given the

broad expanse of the bentliic plain and the unknown

population densities of Xenophora, samples are likelv to

remain small. Many of the non-collected coral species

were onlv passed over 1 time (i.e., were onlv present at

1 Xenophora source station where thev were not affixed).

This single occurrence makes their omission statisHcallv

negligible, but others were conspicuousl)- absent from

Xenophora shells, suggesting that there are groups with-

in azooxanthellate ScleracHnia which Xcnopliora does

not affix. The reasons behind this are often clear: for

example, Xenophora is unlikely to affix any coral that

settles and anchors firmlv onto hard substrate. Similarlv,

colonial corals are tvpically too bulk-y and irregular to be

affixed. There is a strong possibility' of a selection mech-

anism in Xenophora that causes them to choose certain

more suitablv shaped corals over others.

Still, it is difficult to dispute the usefulness of Xeno-

phora as prox\-collectors. Examination of anv Xenopho-

ra collecteil in a dredge sample mav reveal attached spe-

cies that were not otherwise collected in that sample.

Because bhnd dredging is still the most common method

for sampling the benthos, anv method that complements

or makes more complete use of the dredged sample

should not be overlooked.

Attachment Beiiavk )r

Introduction: Arthur Adams (1848:248), the natural-

ist and assistant-surgeon on board the H.M.S. Samarang,

observed Xenophora (species unknown) during an 1842

vovage from Singapore to Java. Adams remarked: "In

order for them to escape from their enemies, nature has

instructed them to cover dieir shells with the same ma-

terial as those of the banks which thev inhabit." Adams'

rationale is understandable. Manv people, both shell-col-

lectors and scientists, have made the same assumption

about Xenophora In the modem era of deep-sea explo-

ration, however, it is unreasonable to accept the visual

camouflage hvpothesis for those species of Xenophora

that live below the depth of light penetration.

Although manv of the frequently attaching species can

be found in relativelv shallow water, 2 of die most spec-

tacularlv encmsted species (X. paUiduIa and X japonica)

range deeper than their congeners (Pf)nder, 1983). Ob-

ject attachment is an energeticallv e.\q")ensive behavior

(Shank, 1969). For Xenophora to attach an object to the

growing edge of the shell it must first select and properly

position the object (see Shank, 1969), and then affix it

bv secreting shell material around it, using more calcium

carbonate than would be retjuired to secrete an equiv-

alent section of shell. Given that members of the familv

Xenophoridae are found in the fossil record as far back

as the Cretaceous, it is unlikelv that such an energeticallv

expensive behavior would persist if it was purelv non-

adaptive.

A number of hvpotheses have been suggested to ex-

plain attachment behavior. These fall into 2 broad,

mechanistic categories: defense and functional support.

Table 2 lists the various hvpotheses and the authors that

have supported them in the past. It is rare for anv author

to cite just one hvpothesis, but no recent efforts have

been made to collect and review all the hvpotheses, al-

though Linslev and Yochelson (1973) and St. Jean (1977,

1983) provided the most comprehensive review previous

to this report.

Defense Theories: Most authors still regard camou-

flage as the most likelv ex}:)lanati()n tor object fi.xation bv

Xenophora. Camouflage usuallv implies visual conceal-

ment, but in the case of deep-water Xenophora, it must

be expanded to include olfacton/ concealment as well.

Shank (1969: 5), one of the few people to maintain Xen-

Page 78

THE NAUTILUS, Vol. 112, No, 3

Table 2. Outline of possible adaptive advantages of attach-

ment behavior in Xenophora, and those audiors who have dis-

cussed those views.

I. Defensive

A. Camouflage

l.\'isual (.\dams, 1848; Morton, 1958; Shank, 1969; Lin-

slev and Yochelson, 197.3; Berg, 1975; St. Jean, 1977,

and 1983; Das, et al., 1981; Ponder, 1983)

2. Olfacton- (Shank, 1969; Linslev and Yochelson, 1973;

St. Jean' 1977, 1983: Ponder, 1983)

3. Tactile

B. Armor Hvpotliesis

II. Functional Support

A. Snowshoeing

B. Increased Shell Stabilitv' (Berg, 1975; St. Jean, 1983)

C. Feeding Cone Hvpothesis (Shank, 1969; Linslev and

Yochelson, 1973; Berg, 1975; St. Jean, 1977, 1983)

ophora alive in aquaria for any length of time, remarked

that "evers •thing [Xenophora] do points to a means of

eluding detection." While obsening captive specimens

of Xenophora conchyliophora, a species hpicaJh' found

in shallower water. Shank witnessed the animals burying

their feces, a behavioral trait generalK- associated with

predator evasion (Shank, 1969; Linslev and Yochelson,

1973). Shank and others have also observed that the pe-

cuhar and discontinuous motion oi Xenophora would be

less likely to attract the attention of a predator, and

might create less of a chemical trail than the "crawling"

movement of many gastropods (Shank, 1969; Linslev

and Yochelson, 1973; St. Jean, 1977, 1983).

These observations are of value for understanding the

ecologv' of a shallow-water species of Xenopht)ra, but

may not be generalizable to all species in the family.

Linsley and Yochelson (1973) observed (Linsley's obser-

vations) live X. neozealanica for several months and cbd

not witness the feces-burving behavior recorded by

Shank. Additionally, they noted that, despite the char-

acteristically discontinuous motion of X. neozealanica.

the animal's foot was in contact with the sechment fre-

quently, impKing that the discontinuous trail hypothesis

is less hkely for that species.

Tactile canwuflage is also a possible advantage of af-

fixing objects. If animals that find prey by touch en-

counter Xenophora. the peculiar assortment of attach-

ments might confuse them. We consider this hvpothesis

unlikely due to the lack of complete shell-coverage in

many species. Although X. conchyliophora and X.japon-

ica are often completely covered, other species that fre-

quently .show attachments (e.g., X. pallidula. X. peroni-

ana. and X crispa) do not coat their entire shells with

affixed objects. The attachment pattern of this latter

group would probably only deter a cursorv chemo-tactile

search, and, in the ca,se of shallow-water specimens,

would be unlikely to deceive a visually orienting preda-

tor

To our knowledge, the suggestion that objects are at-

tached as a means of annorin'^ shells against predators

has not previously been made. Ponder (1983), however,

noted that the shells of some Xenophora species are

quite thin. Considering the extent to which some spe-

cies, such as X. conchi/liophora. are encrusted, one may

theorize that objects are affi.xed to enhance the protec-

tive value of the shell. There are a number of arguments

that can be made against this hvpothesis. W'hereas it is

not known what Upe of animal prevs upon Xenophora,

both Berg (1975) and Linsley and Yochelson (1973) at-

tempted to elicit an escape response from Xenophora

(X. conchyliophora and X. neozealanica, respectively) by

confronting them with starfish and predaton- gastropods.

Linsley and Yochelson (1973: 5) reported that "the gen-

eral reaction to these predators was one of apparent un-

concern.' Berg (1975) fists several species offish as prin-

ciple predators. Affixed objects mav have made Xeno-

phora immune to predation from drilling gastropods;

however, armor of this tvpe would provide little defense

against predation by malacophagous fish, which routinely

crush far more durable shells (Berg, 1975). As noted

above, few Xenophora are as thoroughly covered as X.

conchyliopliora. The mechanism of attachment is argu-

ably inefficient for producing armor: any object affixed

in such a way that it projects out from the shell margin

is largely useless as armor until the next whorl is formecl

under it. Finally, the species of Xenophora that do not

frequently affix foreign objects produce either a pro-

jecting, unbroken flange, or a fringe of radial spines or

digitations (Figure 2), which suggests a function analo-

gous to that of the projecting fringe of attachments, but

lack any armoring potential. It has lieen argued that oth-

er gastropods, such as the genus Miircx, produce spines

to e.xpand their effective size and make themselves less

vTilnerable to predators (oral pers. conini., M. G. Har-

asewvch). Berg (1975) has observed that the affixed ob-

jects quadruple the basal area of X. conchyliophora, so

Xenophora may also be employing this mechanism.

Functional Support Theories: The gross morpho-

logical similarih,' between fretjuentlv attaching and spine/

flange-producing species of Xenophora (Figure 2) is the

source of several structural theories. That certain struc-

tund features appear to be present in iill species argues

powerfully for a common functional basis. Ponder (1983:

43) has noted this siinilarih remarking for X. pallidula

that "this species habitudlv attaches large, often elon-

gate shells in a radial fashion, so diat they presumably

function in the same way as digitations on some other

species." We believe that the analog) can be e.xpanded

in some respects to all known species of Xenophora.

.\s mentioned above, the flange or spines produced

by some Xenophora expand the effective radius of the

shell in much the same way as the projecting fringe of

attachments. One possible purpose for this projecting

"sldrt ' is suggested by the material nature of the benthic

plain: if Xenophora commonly inhabit soft (i.e.. high wa-

ter content) substrates, the\ run the risk of sinking and

suffocation. Thayer (1975:185) has assembled a list of

the various methods used by invertebrates to confront

N. Feinstein and S. D. Cairns, 1998

Page 79

this danger, among which is tlie so-called "snowshoc"

effect: "The potential \alne of a hroad. Hat form to dis-

tribute the weight of an organism has long been recog-

nized. In 1909, Homell (p. 92) suggested that the win-

dowpane ovster, Placuna placenta, was able to live on

soft mud In- emphning 'the same principle as is embod-

ied in the use of snowshoe '. Althougii snowshtjeing has

not been mentioned in the Hterature on Xenophora.

both attachments and fianges seems to have the stnic-

tural potential to ser\e that puipose.

More careful scrutin\' reveals significant problems

\\ith this h\pothesis. In order to snowshoe ef f ecti\ elw

Xenophora would be expected to select for broad, light-

weight objects that expand the contact profile as much

as possible while minimizing the addition of weight. But

Xenophora from certain regions manufacture an attach-

ment fringe entirelv from elongate gastropod shells,

which would penetrate the surface of soft substrate with

little resistance. Also, several species oi Xenophora often

bear more than dieir owii weight in attachments (Shank,

1969; Linsley and Yochelson, 1973). Our own observa-

tions suggest that Xenophora attach objects with little

regard to weight.

Because mass increases in proportion to \olume, with

the cube of length, we e.xpected that the expanded basal

area produced bv the attachments would increase rela-

tive to the basal area of the shell as the animal grew to

mature size. We found the opposite to be tnie for A'.

pallklnia: the ratio of ex]3anded shell basal area with at-

tachments to shell basal area without attachments de-

creased significantly (p<0.01) in the larger whorls. In

other words, as Xenophora pallkluJa grow larger, their

hypothetical snowshoe becomes relativeK smaller.

Though the same test could not be performed on the

spines of X. Solaris due to the large number of broken

spines, we found that the number of spines per whorl

increased statisticalh' (p<0.()l). Thus, X, solans is in-

creasing the area of contact with the substrate b\' de-

creasing the angular interval between spine production

(Figure 2). Although this itself is not comincing evi-

dence of snowshoeing, it suggests that more thorough

moqihometric analvsis of X. solans is needed.

Multiple authors ha\e noted that tlie expanded base

provides added stability to the shell (Shank, 1969; Berg,

1975; St. Jean, 198.3). Although Shank (1969) and Berg

(1975) observed that Xenophora is capable of righting

itself even if cjvertumed in soft sediment, both also not-

ed that the righting procedure is time-consuming. Xen-

ophora may be in greater danger of predation when

overturned, either due to lack of camouflage or to ex-

posure of the apertural surface. If this is the case, re-

ducing the amount of time spent overturned and in the

righting process would give attaching and flange/spine-

producing Xenophora an advantage over less derived

forms. We consider this stabilits' hvpothesis to be rela-

tively unimportant: although it is empiricallv' true that

Xenophora are more stable than thev would be without

the spines, flange, or attachments, without further eco-

logical knowledge it is impossible to sav whether this

provides a selective advantage. Some potential predators

of Xenophora, such as fish, would certainly be capable

of flipping the shell, even assuming that an upright shell

would be less vulnerable than an overturned shell.

One significant structural feature is conserved among

all species of Xenophora: the apertural surface of the

shell is raised off the substrate, although different spe-

cies may achieve that using different structures (e.g.,

spines, flange, attachments). This has been observed bv

Shank (1969), Unslev and Yochelson (1973), and St.

Jean (1977, 1983). Linslev and Yochelson place partic-

ular emphasis on this trait, suggesting that the "stilt"

effect provides a means of olfactorv camouflage by al-

lowing the Xenophora's bodv to remain suspended above

the substrate and thus leaving a cfiscontinuous scent trail.

We believe that the "stilts ' may also aid in the feeding

process bv providing a broad-based feeding cone under

which the animal is able to graze. This hvpothesis is sup-

ported by its sedentarv- lifestvle. As Shank (1969:5) has

commented, "when there is plentv of food it never

reaches beyond its shefl, but feeds entirely on the ma-

terial beneath . . . '. Whether or not this "feeding cone"

provides Xenophora with some safetv while feeding is

subject to debate.

A number of more or less convincing dieories have

been presented as to the function and origin of the at-

tachment behavior in Xenophora. Our observations of

coral-bearing specimens have led us to conclude that

there is some measure of truth in several of the various

theories. We also suggest that different species of Xen-

ophora, subject to different predation pressure and dif-

ferent environmental conditions, mav have adapted the

basic attachment behavior to serve different purposes.

For e.xample, it seems likely that shallow-dwelling, highly

encrusted .species, such as X. concht/liophora. camou-

flages itself against visually orienting predators, whereas

the deep-water, spinose X. Solaris ma)' use the snowshoe

effect to prevent suffocation. We conjecture that visual

camouflage was the original function. The degree to

which the other functions are derived presents an inter-

esting evolutionarv puzzle, but the attachment and

flange/spine-producing behaviors can certainlv be used

to help clarif}' the phvlogenetic relationships between

the various species of Xenophora.

To achieve a more satisfacton- understanding of the

attachment behavior detailed moq;)hometric an;ilvsis

should be performed on the size, shape, spatial density,

and orientation of attached objects. Ultimatelv, however,

we need to known more about the life historv of Xeno-

phora. The differences and similarities in the biology,

ecologv, and behavior of the various species should pro-

vide important clues, and perhaps eventuallv' answer the

riddles posed bv the remarkable carrier shells.

ACKNOWLEDGEMENTS

Both M. G. Harasewych and Ralph Chapman (USNM)

provided vital support and advice throughout the course

of this prcjject. We are ;ilso grateful to Ellis Yochelson

Page 80

THE NAUTILUS, Vol. 112, No. 3

(USGS, ret.), Winston Ponder (Australian Museum),

Kenneth Boss (MCZ), and Beatrice Burch (Bishop Mu-

seum) for offering advice and insight. F'inallw we thank

Alain Crosnier and Philippe Bouchet (MNHN), Tim

Pearce (DMNH), and Dan Graf (MCZ) for the loan of

Xenophora samples. This work was initiated while the

first author was a participant in the 1997 Research Train-

ing Program at the National Museum of Natural Histor\',

Smithsonian Institution. Partial funding for this research

was provided b\- a grant from the National Science

Foundation to support the Research Tniining Program.

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Berg Jr. C. ]. 1974. A comparative ethological study of strom-

bid gastropods. Behaviour 51(.3— 4):.31.3-.321.

Berg Jr, C. J. 1975. Behavior and ecology of conch (Superfani-

ilv Strombacea) on a deep subtidal algal plain. Bulletin of

Marine Science 2.5(3)::307-.317.

Cainis, S. D. In press. Cnidaria Anthozoa: deep-water azooxan-

thellate Scleractinia from Vanuatu and VVallis antl Futuna

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Calms, S. D. and H. Zibrowius. 1997. Cnidaria Anthozoa:

Azooxanthellate Scleractinia from the Philippine and In-

donesian regions. Memoires du Museum national

d'Histoire naturelie 172: 27-243.

Das, A. K., S. C. Mitra and S. Mukhopadhyava. 1981. Studies

on some molluscan collections by the "Golden Crown"

from the Bay of Bengal with a note oti the camouflage

habit of a gastropod, Xenophora pallidiila (Reeve). Pro-

ceedings of the Zoological Societv Calcutta 32:79-87.

Honiell, J. 1909. Report on the anatomv of Placunta placenta

with notes upon its distribution and economic uses. Re-

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in Kattiawar India l:4."3-97.

Kawase, M. 1996. The bathval shell Xcnophorii paUidurn. from

the sea bottom near the Koshiki Island west of Kagoshinia

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seum of Natural Historv 6:21-2.5.

Linslev, R. M. and E. L. Yochelson. 1973. Devonian carrier

shells (Euomphalidae) from North America and Geniianv:

a study of a behavior pattern in which foreign matter is

attached to the shell of living and Devonian gastropods.

U. S. Geological Survev Professional Paper 824:1-26.

Morton, J. E. 19.58. The adaptations and relationships of the

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Ponder, W. F. 1983. Xenophoridae of the world. Memoirs of

the Australian Museum 17:1-126.

Pourtales, L. F. de. 1871. Deep-sea corals. Memoirs of the

Museum of Comparative Zoology 2:1-93.

Reeve, L. 1842. On the genus Phonis. a group of agglutinating

mollusks of the familv Turfiinacea. Proceedings of the

Zoological Socieh' of London 10:160-161.

St. Jean, K. 1977. The Xenophora -how and whv thev collect:

some new insights. The Western Societv of Malacologists

Annual Report 10:11.

St. Jean. K, 1983. Xenophora digitata \on Martens -the missing

' link. Of Sea and Shore I3(l):17-I9.

Shank, P. 1969. The Hmorous carrier shell. New York Shell

Club Notes 15I:.5-7.

Stolarsld, J. 1996. Gardineria -a scleractmian living fossil. .■Kcta

Paleontologica Polonica 41(4):339-.367,

Thaver, C. W. 1975. ,\daptations of benthic invertebrates to

soft substrata. Journal of Marine Research 33(2): 177-189.

Appendix 1. Names and localities of azooxanthellate coral species affixed to Xenophora shells.

MNHN

.VIUSORSTOM .V963 i20"20'S, 169-49'E, 400-140 m), on .V. palhdida. MNHN: F variegatus (I), S. complicata (I), CanjophjlUa

sp. A (1), C ahnipta (1), T brevispUm (1), B. stelhdnliis (2). D cornioatits (1). H' sulcatus (2). H. sp. (1): I lahidni U),

Flabelluin sp. (2), F aotearoa {\),T. kenuach-eensis (1).

.MUSORSTO.M 8-1008 (18°.53'S. 168°.53'E. 919-1000 m), on .V. paUidnla. MNHN: TmncatojiaheUum sp. (I), gardineriid, n. gen.

n. sp. (3).

MUSORSTOM 8-1016 (17°.53'S, 168°28'E, 291-300 m), on .V. pallidula. MNHN: C ahnipta (1), Conotrochiis sp. (1). C. astjm-

metros (I), 7! dens (1).

MUSORSTOM 8-1017 (17°53'S, 168°26'E, 294-295 m), on .V palhdida. MNHN: Canjoplitjlha sp. (1), C a.sipnnietros (11, Con-

olrorhti.s sp. (1), E. graiji (1).

MUSORSTO.M .8-1018 (ir.53'S, 168°25'E. .300-301 m), on X. pallidula. MNHN; B .'^telhdatu.s (1), N. ronieiis (\).

MUSORSTOM .8-1023 (17°48'S. 16.8°49'E. .321 m), on X pallidula. MNHN: Canjophiillia sp. (1), Canjophi/llia sp. B (1). B.

stelhdatus (I), Deltoetjuthus sp. (1), Conotroehus sp. (2). // suleafiis (I), Tnineatofl(dnlluiu sp. (I), Flabelluin sp. (1). T kcr-

nutdeeen.'iis (2).

MUSORSTOM 8-10S7 (I.5°I0'S, 167°14'E, .394-421 m). on .V pallidula. MNHN: Canjophjllia sp. (1), C. abnipta i\). C. deeanwra

(1), Trochoct/athus sp. (I), T. rhombocolumna (1), T va.'-iifonnis (1), T sp. (1), B. stellnlatn.s (2), gardineriid. n. gen. n. sp. (2).

MUSORSTOM 8-1088 (1.5°09'S, 167°15'E. 42.5-4.55 m), on .V pallidula. MNHN: C. abnipta (2). P. follirulus (l)'.'T. discu.1 (2), T.

sp. (] ). Tnincatofiabelluni sp. (1), gardineriid, n. gen. n. .sp. (2).

MUSORSTOM 8-1091 (1,5°1()'S, 167°I3'E, .344-.35() m), on X. pallidula. MNHN: S, eoniplieata (1). C. abnipta (3). C lainellifera

(1). Ileteroei/alhu.s sp. (1), F pavoninuni (I).

MU.SOR.STOM 8-1092 (1.5°10'S, 167°12'E, 314-321 m). on X pallulnla. MNHN: C andnosia (H. C fuuieolnnina (3). Tninea-

toflalxllum sp. (2).

MUSORSTOM 8-1097 (I.5°05'S, 167°10'E, 281-288 m), on X, pallidula. .\INI1N: .S, eoniplieala {\).B. slellulatns (W ). Delloeyallius

sp. (I), D. Stella (1), H. snlcatus (I), C. asijmmctros (1). C bninneus (1). C sp. (1), Flabelluin sp. (1). T piisilluni (1). t s]i .\

(I). G annulata (1).

N. Feinstein and S. D. Cairns, 1998 Page 81

MUSORSTOM S-1106 (15°05'S. 167°11'E, 305-314 m), on X. pnllidula. MNHN: C abniptti (1). B stcllulntus (9). D hetcwclitus

(2), D sp. (1), H mlcatiis (1), T. dens (1), T. ptmllum (3), T. sp. (2), dendrophylliid (2).

DMNH

Marinduque, Philippines, 124-126 ni, on X pallidiila. D.MNH 188571: CaniophijUin sp. A (2), Flal>clluin sp. (1), TnincatoflahcUnm

sp. (1).

Marinduque, Philippines, depth not knowii, on X, pallidula, DMNH 189002 and 188943: A. fni\tuin (1), C spiimcarens (4), H.

sulcatus (1), F deludcns (1), Tnincatoflabelhnn sp. (1), B coniu (2).

Batangas, Philippines, 152-163 m, on X palUdula. DMNH 189854: T. formtmim (3).

Batangas, Philippines, depth unknown, on .V. pnllidula. DMNH 190007: C. spinacnrens (1), H stdcatus (1).

Philippines, depth unknown, on A', pallidula. DMNH 188572: C. spinacarens (7), C. transvcrsali.'i (1).

Philippines, depth unknowni, on .V. pallidida. DMNH 188944: C. spinacarens (1), T. pusillum (1), B cornu (1). B iinpcrialis (1).

Quezon, Philippines, 596 m, on A', pallidida. DMNH 189043: Tnincatoflahelluni sp. (1).

OffZnluland. 293 ni, on A. pallidnla. DMNH 11505-7, 11499, 17902: Madmcis sp. (1). Canjophi/llia sp. (2), FMullum sp. (I ), E

paioninum (5), Tnincatoflahcllum sp. (6), T sflrdincii (6), dendroplnlliid (1).

Off Zululand, 220-293 ni, on A pallidula. DMNH 20120: F paioninum (4), F sp. (1), T sp. (1), 7" cf. fonnosum (1), I g,ardineri

(10).

Off Natal, South Africa, 293-329 in, on X. pallidula. DMNH 186438: Flabcllum sp. (1), F pauon/UHHi (1), I gardincri (2).

off Japan, 110 in, on X. pallidula, DMNH 6180: C quadragcnaria (1), F dentiformis (2), F paioninum (1), Endopachi/s graiji (1).

Sagaini Bay, Japan, depth unknown, on X. pallidula. DMNH 55071: F palifenis (1), H. sulcatus (1), Tmncatoftahcllnm sp. (1).

off Kii, Japan, 183 m, on X. pallidula. DMNH 55070: £. grai// (1), P dentiformis (2).

MCZ

Auckland, New Zealand, depth unknown), on .V, neozcnlandica. MCZ 160266-67: M nibnni} (4).

USNM

Albatross 3810 (.soudieast of OiJui, Hawaiian Islands, 386—463 in), on A, pcroniana. USNM M749844: A, pacifica (1), £. grai/' (!)•

Albatross 3838 (southeast coast of Molokai, Hawaiian Islands, 168-.388 in), on X. peroniana, USNM M335067: E grat/i (5).

Albatross 4079 (Puniawa Point, Maui, Hawaiian Islands. 262-326 in), on X. peroniana. USNM M.3.35063: A macrolobata (3).

Albatro.-is 5117 (13°52'22"N. 120°46'22"E. 216 in), on A. pallidula. USNM M243416: C, spinigcr (1), Flabcllum sp. (1).

Albatross 5131 (off Panabutan Point, Philippines. 49 in), on .\'. pallidula. USNM M243391: C.'secta (1), H. .mlcatus (1).

Albatross 5212 (12°04'15"N, 124°04'36"E, 198 m), on X pallidula. USN.M M277576: F lamellulosum (2), T cf gardincri (1), B.

coniu (3), B. imperialis (1).

Albatross 5213 (12°15'00"N, 123°57'.30"E, 146 ni), on A. pallidula. USNM M243398: Tmncatoflabcllum sp. (1).

Albatross 5265 (13°4I'15"N, 120°00'50"E, 247 m). on X. pallidula, USNM M243347-9, -353^ -356-357, -.361-2. -.366, -.368,

-370, -372, -373, -375, -376, -426, -881: Anthcmiphijllia sp. (I), C. spinnrflredA (1), C. .swta (2). C. graiji (1). T. t/rgnffw

(1), H. sulcatus (1), D. philippinensis (1), C. /j(7e(j.s (I), Flabcllum sp. (3), F pavoninum (I), F lamcllulo.wm (I), T. fonnosum

(12), T f/ms (1), I pusillum (1), I sp. (1), B. co;7i!/ (I), B. imperialis (2). £ grai/i (D. W roWi/ra (1).

Albatross 5273 (13°58'45"N. 120°21'35"E, 209 in), on X. pallidula. USNM M237519: Balanophi/llia sp. (I).

Albatross 527H (I4°00'1()"N, 12(n7'1.5"E. 187 in), on X pallidula. USNM M 24,3411: C. octonaria (1), H. .si//rafi/.v (3). H. altcrnatus

(1), A. nibesccns (1), Tropidocijathus sp. (1).

Albatross 5289 (13°41'.50"N, 120°58'30"E, 315 in), on .\. pallidula. USNM M237585: 7" fonnosum (1). I (;irn/.sto?i/»i (1).

Albatross 5312 (21°30'00"N. 116°32'00"E. 256 in), on A. japonica. USNM M243420: I cf gardincri (1).

Albatross 5391 (12°13'15"N, 124°05'03"E, 216 in), on X. japonica. USNM M238138: H. .siJraffl (1), F po/ifi/»! (14). T candcanum

(1), Balanophijllia sp. (2).

Albatross 5392 (i2°13'35"N, 124°02'48"E, 247 in), on A pallidula. USNM M238192, 243394: F paioninum (2). F /A)/(f!/»i (7), F

sp. (10). B. ronii/ (4),

Albatross 5392 (see above) on X. japonica. USNM M238192: F politum (50). F paioninum (1), F sp. (13), B. foni!/ (6), B. sp.

(5).

Albatross 5394 (12°00'.30"N. 124°05'36"E. 280 m), on A pallidula. USNM M24.34I5: T candcanum (1).

Albatross .5395 (ir56'40"N. 124°14'E. 2.56 in), on .Y, pallidula. USNM M240513: B wr/ii; (1).

Albatross 5405 (10°49'20"N, 124°24'23"E. 479 m). on X pallidula. USNM M238257: Co/iofroc/u/.y sp. (2).

Albatross 5408 (10°40'I5"N. 124°15'00"E. 291 ni). on A. pallidula. USNM M238277: Fimgiacijathus sp. (I). Madrepora oculata

(2), H. .s(//cn?i/.s (4).

Albatross 5416 (I0°11'30"N, 12,3°53'30"E. 274 in), on A pallidula. USNM M238372: A. fnistum (2). //, .si/Zcrtfi/.s (5). Flabcllum sp.

(3). T angustum (1).

Albatross .5417 (10°10'00"N, 12.3°53'I5"E. 302 in), on .V pallidula. USNM M24.3381: Deltocijathus sp. (I). Flabcllum sp. (2). F

paioninum (2).

Albatross 5418 (I0°08'5t)"N. 12:3°52':30"E, 291 in), on X pallidula. USNM M243377: .\/. ori/Z^ffl (1), D rt(u/«/(m/i/c».v (1), Flabcllum

sp. (I), r angustum (1).

Albatross 5592 (4°12'44"N, 118°27'44"E. 558 in), on A. pallidula. USNM .M229317: H. altcrnatus (1).

Albatross 5661 (5°49'40"S, 120°24'30"E, 329 in), on A. pallidula. USNM M239497: Canjophi/llia sp. (1).

Dauirf Starr Jordan, stn TC32-2 (21°21'54"N. 158°I2'24'\V. 119-291 m). on A, pcroniana. USN.M M8076,59: Anthcmiphi/llia sp.

(1), £. grflyi (2). Placotrochides n. sp. (1).

Dnr/f/ Starr Jordan, stn TC3:3-9 (2r00'06"N. I56°45'42"W. 227-234 in), on .V peroniana. USNM M807662: A, pflf(/ir« (1),

Flabcllum sp. (2). F paioninum (1), Placotrochides n. sp. (3).

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THE NAUTILUS, Vol. 112, No. 3

David Starr Jordan, stn TC40-8 (21°09'42"N, 157°24'42"W, 183 ni), on X. penmiana. USNM MS076.31: Balnn<ip}i,jUui sp. (1), E.

grayi (2).

David Starr Jordan, stn TC52-99 (2r02'06"N, 156°47'1.5"W, 223 m), on X. peroninna. USNM M807652: £. grai/i (3).

Off Kisamaya, Somalia, depth unknown, on X pallidiila. USNM M819755: F pavoninitm (6).

16-32 km off sontliem Zuhiland, 293-329 m, on X. paUidula. USNM M6.3.5312-313: H sulcatus (1), F. pavuninuin (1), T gardiiwii

(1), 7! intdti.spiuosum (2), T. sp. (2).

W-320, off Natid, S. .Africa, .37 m, on X pcdiidida. USNM M8I8232: H mlcatm (1), Flalnllum sp. (\),T. gardiiu-ri (2).

W-2.53, off Durban, S. Africa, .366 m, on X. pallidula. USNM M818231: T multi.spino.su m (1).

Sagami Bay, Japan, depth unknown, on X. pallidula. USNM M.346L51: C dentata (1), P. dentifonni.s (1). T candcanum (1), H.

cochlea (3).

Kanabe, Kii, Japan, depth unknown, on .V, pallidula, USNM M27368(): P dcntifonuis (9), Heteroci/athu.s sp. (1), Truncatoflabdlum

sp. (1).

Off Kii, Japan, depth unknown, on X. pallidula. USNM M60.591.5: P. dentifonni.s (1), dendrophvlliid (1).

Minabe, Wakavania, Japan, 64 m, on X, pallidula, USNM M60.578.5: P dentifonni.s (1), H sulcatus (1), dendroplnlliid (1).

Off Tosa, Japan, depth unknown, on X. cerea. USNM M.346L50: C. jogashinmensi.s (1), Truncatofiabellum sp. B (2).

Off Japan (W-976), on X japonica. USNM M.348868; Tmncatoflabellum sp. (2).

Sulu Sea, Philippines (W-498), on X. pallidula. USNM MS07763: T eooperi (3), H. sulcatus (1), P laevi.s (1). H coctdea (2).

Tavabas Bav, Philippines, L5-,30 m, on X. pallidula. USNM M 876993: H sulcatus (4).

Stn CPI/8 ('3°15'S, 128°08'E, 26-55 m), on X. comigata, USNM M746695; H cochlea (1).

Stn KRVI (.5°32'S, 132°41'E, 37 m), on X. comigata. USNM M746948: H. cochlea (1).

22°19'S, 167°11'E, 290-310 m, on paratype of X. granulosa. USNM M842996: D heteroclitus (1).

LaRafalo. stn 7 (.5°18'N, 9°54'.30"W, 200 m), on X. cri.spa, USNM M762004: C. smithii (23).

Appendix 2. Taxonomic list of iizooxanthellate Scleractinia af-

fixed to Xenophora shells, and frequency of occurrence on

those shells (in parentheses). See Appendix 1 for localiU' data

of corals and shells.

Order Scleractinia

Suborder Astrocoeniina

Fainily Pocillopoiidae

Madracis sp. (1)

Suborder Funpina

Family Fungiacyathidae

Fungiactjathus palifenis (Alcock, 1902) (1)

F. variegatu.s Cainis, 1989 (1)

F sp. (i)

FaniiK' Vlicrabaciitlae

Stephanoplujllia eoinplicata Moseley, 1876 (.3)

Suborder Fa\iina

Family Ociilinidae

Madrepora oculata Linnaeus, 1758 (3)

Family Anthemiphvlliidae

Anthemiphijllia fnistum Cairns, 1994 (3)

A. pacijica Vaughan, 1907 (2)

A. nuicrolohata Cairns, in press (.3)

A. sp. (1)

Suborder Caryophylliina

Family Carvophylliidai'

Canjophi/llia (('.) smithii Stokes ;uid Bnxlerip, 1828 (23)

C. (C.) ahntpta Cainis, ms (9)

C. (C.) sixta Cairns and Zibrowius, 1997 (3)

C. (C.) ambrosia Alcock, 1898 (1)

C. (C.) joga.shintaensis Eguchi, 1968 (1)

C. (C.) lamellifera Moseley 1881 {])

C. (C.) octonaria (;airns and Zibrowius, 1997 (1)

C. (C.) (juadragenana Alcock. 1902 (I)

C. (C.) transversalis Moselev, 1881 (1)

C. (C.) sp. A (3)

C. (C.) sp. B (1)

C. (C.) sp. (7)

C. (A.) spinicarens (Moseley, 1881) (14)

C. (A.) decamera Cairns, in press (1)

C. (A.) dentata Moseley 1876 (1)

C. (A.) graiji (Milne Edwards and Hainie, 1848) (1)

C. (A.) spinigera Saville Kent, 1871 (1)

Premocyathns dentiformis (Alcock, 1902) (16)

Trochocijathus (T.) eooperi (Gardiner, 1905) (3)

T. (T.) discus Cainis and Zibrowius, 1997 (2)

T. (T.) rhombocolumna Alcock, 1902 (1)

T. (T.) va^iformis Bonnie, 1903 (1)

T. (T.)sp. (2)

T. (A.) brcvispina Cainis and Zibrowius, 1997 (1)

Tethoct/athus virgatus (Alcock, 1902) (1)

Boumeotrochus stelhdatus (Cainis, 1984) (25)

Deltoctjathus heteroclitus Wells, 1984 (3)

D. andamanicus Alcock, 1898 (1)

D. cornigatiis Cainis, in press (1)

D. philippinensis Cairns and Zibrowius, 1997 (1)

D. Stella Cainis and Zibrowius, 1997 (1)

D. .sp. (4)

Hetcronjathus sulcatus (Verrill, 1866) (31)

H. altematus Verrill, 1865 (2)

H. sp. (3)

Conotrochus astpnmetros Cainis, in press (3)

C./u/»r()/i/m)i(/ (.\lcock, 1902) (3)

C. bnmneus (Moseley 1881) (1)

C. sp. (6)

FaniiK Turbinoliidae

Alatotrochus nd)esecns (Moseley, 1876) (1)

Trojjidocijathus labuhis Cainis and Zibrowius, 1997 (1)

T. sp. (1)

Cyathotroehus pihiis (.\lcock, 1902) (1)

Notoci/athus c<iiucus (.-Vlcock, 1902) (1)

Pejkiuoci/atlius folliculns (Pourt;iles, 1868) (1)

FamiK Flabellidae

Flabellum {F.) polituui Cainis. 1989 (71)

F. [F.) pin'oninum Lesson, 1831 (26)

F (F) lamellulosum Alcock, 1902 (3)

F. (Fj sp. A (1)

R (F.) sp. (43)

F (U.) aotearoa Squires, 1964 (1)

F {V.)deludens Mareu/.eller, 1904 (1)

N. Feinstein and S. D. Cairns, 1998

Page 83

TnmcatoflahcUnm nnrdincri Cainis, 1993 (20)

T sp. ct. T ganlincii (3)

T fonno.siim Cainis, 19S9 (16)

T. sp. ct. T. formusum (I)

7! pusiUiim Cainis, 1989 (6)

T angu.stum Cainis and Zibrowiiis, 1997 (2)

T. candcanum (Milne Edwards and Hainie, 1848) (3)

T den.s (Alcock, 1902) (3)

T. incnistattim Cainis, 1989 (1)

T. inidti.spinosuiu Cainis. 1993 (3)

T. sp. A (1)

T. sp. B (2)

T. sp. (25)

Placotrochides n. sp. (4)

Placotrochiis laevis Milne Edwards luid Hiiinie, LS48 (1)

Monoiiu/ces ndmirn (Qnov and Gainiard, 1833) (4)

Faniilv Gardineriidae

Gardineriid n. gen. sensu Stolarsld (1996) (7)

Family Guyniidae

Gmjnia anmdata Duncan, 1872 (1)

Temnotrochiis kcnnadcccnsis Caims, 199.5 (3)

Suborder Dendroplivlliina

Family Dendrophvlliidae

Dendrophvlliid (5)

Balanophijllia cornti Moseley, 1881 (18)

B. iiiipciialis Kent, 1871 (4)

B. sp. (9)

Endopachys graiji Milne Edwards and Haime, 1848 (17)

Heteropsammia cochlea (Spengler, 1781) (8)

Solitarv corals unidentified to genus (67)

THE NAUTILUS 112(3):84-S9, 1998

Page 84

Shell Microstructure of Mytilids (Bivalvia) from

Deep-Sea Hydrothermal Vent and Cold-Water

Sulfide/Mediane Seep Environments

Michael J. Kennish

Institute of Marine and Coastal

Sciences

Rutgers University

New Bmnswick, NJ 08903 USA

kennishO'ahab. Rutgers.edu

Antonieto S. Tan

Department of Biology'

Worcester State College

Worcester, MA 01602 USA

Richard A. Lutz

Institute of Marine and Coastal

Sciences

Rutgers Universih-

New Bninswick, NJ 08903 USA

ABSTRACT

The shell microstnictures of 6 nutilid species from deep-sea

Indrothernial vent and cold-water sulfide/methane seep sites is

characterized b\' scanning electron microscop\'. Examination of

fractured and sectioned specimens reveals similar shell mi-

crostnictures in distinct arrangements. The shell microstrucct-

ures of the nntilids insestigated in this study consist of outer

fibrovis prismatic and inner nacreous stnictures, which underlie

a periostracal laver Simple prismatic stnicture. comprisnig the

adductor and pallial mvostraca, is also present. The shell mi-

crostnictures of these species exliibit similar complexity' as mi-

crostructures of the previously described vent mussel Bathtj-

mocliolus thermophilus Kenk and Wilson, 1985.

Table 1. Deep-sea mvtilid specimens examined in this studv.

Date

Sampling site

Depth

(m)

Species

Collection

number

Specimen Length

number (cm)

Cold-water Sulfide/Methane Seeps

6/3/92 Florida Escaipnient

26°01.8'N

84°.54.6'W

10/16/86 Florida EscaqJinent

26''01.5'N

84°55.3'W

4/1.3/90 Aiaminos Canvon

26°21.3'N

94°29.7'W

9/15/91 Louisiana Slope

(Bu.sh Hill)

27°46.9'N

9r30.4'W

9/29/91 Louisiana Slope

27°.50'N

92°10'W

Hvdrothermal Vents

6/21/93 Mid-Atlantic Ridge

(Snake Pit)

23°22.1'N

44°57.1'W

5/28/90 Galajiagos Rift

(Mussel Bed)

0()°47.9'N

86°09.2'W

6/7/90 East Pacific Rise

(Mussel Bed)

12°48.6'N

10.3°56.5'W

3313

3300

546

Species A

Species B

Species C

Species D

650 Species E

3521 Species F

ANSP400787

ANSP400788

2515 Bathtjinocliolus thermophilus

2630 Btithi/iiKxhohis thcriniijihihis

8.0

14,1

M. J. Kennish et al.. 1998

Page 85

Figures 1-6. External shell surface of the deep-sea mvtilid specimens examined. 1. Species A, ANSP 400787, right \alve. Length

= 8.0 cm. 2. Species B, ANSP 400788. right \-alve. Length = 14.1 cm. 3. Species C. ANSP 400789, right valve. Length = 11.4

cm. 4. Species D. ANSP 400790. right xake. Length = 8.3 cm. 5. Species E, ANSP 400791, right valve. Length = 8.9 cm. 6.

Species F, ANSP 400792, left valve. Length = 12.5 cm.

INTRODUCTION

Among uniijue faiinal as.semblages disco\'erecl at cold-

water sulfide/methane seep sites in the Gult of Me.xico

as well as at deep-sea hxdrothermal \ent sites in the

north Atlantic are a number of unnamed mvtilids. Ef-

forts are undersva\- to describe both the soft and calcified

tissues of these nntilids v\ith tiie primars' purpose of

providing new information for taxonomic differentiation

of members of the group. Here we report for the first

time on the shell microstnictures obsened within the

shells of 5 mxtilid species collected at cold-water sulfide/

methane seep sites in the Gull ot Mexico and 1 species

from deep-sea hvdrothermal vent sites on the Mid-At-

lantic Ridge (2.3°22.1'N, 44°57.I'W). The shell micro-

stnictures of these specimens are also compared uith

those of the pre\iousl\' described, deep-sea h\tlrother-

mal vent mussel Baflujiiiodiolu.s thcnnophihis Kenk and

Wilson, 19S5 collected at sites on the Galapagos Rift and

East Pacific Rise.

MATERIALS AND METHODS

Mvtilid specimens ranging in length from 3.3-14.9 cm

were collected alive via submersible sampling from hv-

drothermal vent sites at Snake Pit (23°22.1'N,

44°57.rW) on the Mid-Atlantic Ridge (3 specimens), at

Mussel Bed (00°47.9'N, S6°09.2'W^ on the Galapagos

Rift (2 specimens), and on the East Pacific Rise

(12°48.6'N, 103°56.5'W) (2 specimens), as well as from

cold-water sulfide/methane seep sites on the Florida Es-

caq^ment (26°01.S'N, 84°54.6'W: 26°01.5'N, S4°55.3'W)

(2 specimens), Alaminos Canyon (26°21.3'N, 94°29.7'W)

(2 specimens), and Louisiana Slope (27°46.9'N,

91°30.4'W; 27°50'N, 92°10'W) (5 specimens). The depth

of the sampling sites varied from 546 to 3521 m (Table

1). The soft tissues of the mussels were excised and fro-

zen at — 70°C. However, thev were not examined his-

tologicalK', and the sex and stage of dexelopment of the

mussels were not determined. The shells were air-dried,

carefullv packaged, and sent to the laboratorv for anal-

\sis. The taxonom\' and .s\steniatic relationships of these

taxa have not been studied. Voucher specimens of the 6

unnamed species (Table 1) were deposited in the col-

lection at the Department of Malacologw Academv of

Natural Sciences of Philadelphia (ANSPX

Sexenteen specimens were prepared for scanning

electron microscop\- (SEM) following the 2 methods

outlined by Kennish et al. (1996, 1998):

Preparation 1. Shells were fractured, sonicated in chs-

tilled water, deh\drated in 95% ethanol, air-dried,

and coated with gold/palladium.

Preparation 2. Shells were embedded in resin, sec-

tioned, pohshed. treated with sodium hvpochloride,

rinsed with water, deh\drated in 95% ethanol, air

dried, and coated with gold/palladium.

Page 86

THE NAUTILUS, Vol. 112, No. 3

Figures 7-12. Internal shell surface of the deep-sea nntilid specimens examined. 7. Species A, ANSP 400787, same specimen

shown in Figure 1. Length = 8.0 cm. 8. Species B, ANSP 4007SS, same specimen shown in Figure 2. Length = 14.1 cm. 9.

Species C, ANSP 400789, same specimen shown in Figure 3. Length = 11.4 cm. 10. Species D, ANSP 400790, same specimen

shown in Figure 4. Length = 8.3 cm. 11. Species E, ANSP 40079L same specimen shown in Figure 5. Length = 8.9 cm. 12.

Species F, ANSP 400792, same specimen showii in Figure 6. Length = 12. .5 cm.

After preparation, the spedmen.s were observed and

photographed in an Amray 1830 SEM.

SHELL MICROSTRUCTURE

Coi.d-Sef.p M"^Tii,ins

The .shells of the cold-seep nnlilids consist of 4 shell

layers. They are basicalK' composed of Outer fibrous pris-

matic and inner nacreous structures which underlie a

periostracal layer that may be infected with branching

filamentous organisms (Figures 1, 2). Simple prismatic

(aragonite) stnicture is also present, constituting tlie ad-

ductor and pallial myostraca. A thin laver of coarse-

grained homogeneous stnicttire mav be present between

the periostracum and the outer fibrous prismatic la\er

(Figures 1, 3, 4). The first-order prisms comprising the

outer fibrous prismatic layer are composed of laths or

rods of calcite (Figures 1, 5, 6), and they have a much

greater lengtli/width ratio than is hpicallv observed in

simple prismatic structure. In the filtrous prismatic-

structure of the cold-seep mydlids the prisms are usuali\

arranged parallel to each other with the long a.\is in-

clined toward the hinge. However, near the periostra-

cum or at growth bands, the prisms assume either a con-

ical or crossed configuration. The conical arrangement

of the prisms resembles planar sphernlitic prismatic

structure (Figure 7), and the crossed arrangement.

crossed lamellar or comple.x crossed lamellar stmcture

(Figures 8, 9).

The underKing nacreous la\er md\ or ma\- not be sep-

arated from the fibrous prismatic laver hx block\' pris-

matic structure (Figures 10, II). At the boundan' be-

tween the fibrous prismatic and nacreous lavers, the na-

creous tablets are arranged as stacks or sheets (Figure

12, 13). Sheet or row-stack nacre occurs awa\' from the

shell margin (Figure 14-16). The adductor and palliid

myostraca e.xJiibit first-order prisms that are well defined

and generalK' non-interdigitating (Figure 17. 18). Thev

ha\e relativeK low to moderate lengtli/v\idth ratios in

contrast to tliose comprising the fibrous prismatic laver.

lU l>H( (llll'.HM M.-VeNT MvTII.IDS

The microstructures encountered within the shells of the

hvdrothermal-vent mytilids examined in this stud\' ;u-e

hpical of calcified structures pre\ionsI\ described in oth-

er members of the tainiK Mstilidae, including the vent

mussel Baflii/iuodiolus thcniiophilus. From the vacuo-

lated periostracum inwards, the calcified lavers of B.

tliermopliilus consist of (I) fibrous prismatic calcite; (2)

nacre (aragonite): (3) irregular prismatic aragonite (pal-

lial mvostracuin); and (4) nacre (aragt)nite) (Lutz and

Hhoads, 1980). A relatively thick organic periostracum

appears to effectiveK' prevent substantial dissolution of

M. J. Kennish et al., 1998

Page 87

Figures 13-21. Microstmctures of deep-sea mytilids. 13-15. Species A. ANSP 400787. 13. Angular view of periostracum (P)

(ventral margin), coarse grained homogeneous (CGH) (comarginal fracture), and fibrous prismatic (FP) stnictures. Autero-posterior

shell axis from top to bottom of micrograph. Horizontal field width (HF\V) = 240 (xm. Preparation 1. 14. E.xterior \ie\v of

periostracum infested with branching filamentous organisms. Same orientation as Figure 1. HFVV = .320 jim. Preparation 1. 15.

Exterior wew of an oblique fracture of the coarse grained homogeneous (CGH) and fibrous prismatic (FP) structiu'es. Antero-

posterior shell axis from top left to bottom right of micrograph. X'entral shell margin to right. HFVV = 40.5 p-m. Preparation 1, 16-

17. Species B, ANSP 400788, 16. Coarse grained homogeneous (CGH) and fibrous prismatic (FP) structures. Double fracture: (1)

radial fracture towards bottom left of micrograph; (2) horizontal (tangential) fracture towards top right. Autero-posterior shell axis

perpendicular to plane of micrograph. HF\V = 70 \x.m. Preparation 1. 17. Transverse fracture of predominantlv lath-type fibrous

prismatic structure. Outer shell surface towards bottom of micrograph. Antero-posterior sliell axis from left to right of micrograph.

HFW = 35 ^x.m. Preparation I. 18. Species C, ANSP 400789, transverse fracture of predominantlv rod-hpe fibrous prismaHe

structure. Outer shell surface towards bottom of micrograph. ."Vntero-posterior shell axis from left to right of micrograph. HFW =

35 \Lm. Preparation 1. 19. Species D, ANSP 400790. Radial fracture showing conical arrangement of rotl-txpe fibrous prismatic

structure. Outer shell surface towards top of micrograph. Outer shell surface towards bottom of micrograph. Antero-posterior shell

axis fVom left to right of micrograph. HFVV = .305 (jlui. Preparation 2. 20-21. Species E, ANSP 400791. 20. Radial fracture showing

crossed lamellar arrangement of lath-txpe fibrous prismatic structure. Antero-posterior shell axis from bottom left to top right of

micrograph. HFW = 70 \xm. Preparation 1 (Same orientation as Figure 7). 21. Radial fracture showing complex crossed lamellar

arrangement of lath-t>pe fibrous prismatic structure. Antero-posterior shell axis from bottom left to top right of micrograph. HFW

= 170 |a.m. Preparation 1 (Same orientation as Figure 7),

the relativei\' thin and fragile shell of B. thcrmopliilus

throughout its life (Lutz, 1982).

The calcitic fihroiis prismatic laver, a specialized layer

observed in the mxlilids, is characterized by prisms with

high length/width ratios. As reported by Carter (1990),

first-order prisms in fibrous prismatic structures have a

non-spherulitic prismatic and non-composite prismatic

substnicture, but unlike simple prisms, appear as long

fibers. Several investigators have classified fibrous pris-

matic structure as an independent entitx' because of its

unique stmcture (MacCIintock, 1967; Carter, 1980a, b;

Carter and Clark, 1985; Watabe, 1988; Carter, 1990).

Nacreous structures in h\'drothermal-vent niNtilids are

composed of tabular aragonitic crxstals deposited on an

organic matrix. The calcareous tablets, identical to tlie

sheet nacre described by Carter and Clark (1985) and

Carter (1990), are arranged in broadh' continuous, reg-

ular, and mutualK parallel sheets. The\ exhibit similar

Page 88

THE NAUTILUS, Vol. 112. No. 3

' .M-i^^A

Figure 22—30. Microstmctiires of deep-.sea nntilitl.s. 22. Specie.s C, ANSP 4007S9. Radiiil Iractuie ot tlie boiiiRlan- between fibrous

prismatic (FP) and nacreous (N) structures. Dorsal shell surface towiirds bottom of micrograph, .'^ntero-posterior shell axis from bottom

left to top right of micrograph. HFW = 3.5 |xm. Preparation 1. 23. Species A, ANSP 400787. Transverse fracture of the blocky

prismatic (BP) sandwiched between the fibrous prismatic (FP) and nacreous (N) structures. Dorsal shell surface towards top of

micrograph. Antero-posterior shell axis from left to right of micrograph. HFW = .35 \i.m. Preparation 1. 24. Species B, ANSP 400788.

Inner surface view of the nacreous stnictnre, near the boundan,' between fibrovis prismatic and nacreous stnictures. \'entnil shell

surface towards left of micrograph. Antero-posterior shell axis Irom top to bottom ot micrograph. HF^\' = 70 jo-m. Preparation 1. 25-

27. Species D, ANSP 400790. 25. Radial section of columnar nacre (CN) underKing the fibrous prismatic (FP) structure, \entral

shell surface towards bottom ot micrograph. Antero-posterior shell axis trom left to right of micrograph. HFW = 45 jjim. Preparation

2. 26. Inner surface view oi hexagouiil tablets of sheet nacre. Shell margin towards bottom left ot micrograph. HF\\' = 70 jjim.

Preparation 1. 27. Inner surface view ot oval tablets ot sheet nacre. Shell margin towards bottom left of micrograph. HFW = 70 \x.m.

Preparation 1. 28. Species D, ANSP 400790. Inner surface view of rhomboidal tablets of stack nacre. Shell margin towards bottom

left of micrograph. HFW = 70 \i.m. Preparation I. 29. Species C, ANSP 400789. Radial fracture of adductor nivo.stracum (AM)

consisting of simple prismatic stmctme underneath the nacreous laver. Inner shell surface towards bottom left of micrograph. HFW

= 70 \i.n\. Preparation 1. 30. Species B, ANSP 400788. Angular view of pallial mvostracum (PM) consisting of simple prismatic

stnicture underneath the nacreous layer Inner shell surface towards bottom right ot micrograph. HFW = 70 ^.ni. Preparation 1.

complexitv and form as those comprising the inner or

middli' shell laver of other nn+ilid species (Carter, 1990).

Irregular simple prismatic structure occurs under the

areas of muscle attachment: the palliiil and adductor

myostraca. The prism cross sections are highly variable

along their lengths (Carter, 198()a; 1990). This irregular-

ity of the prism cross sections results in a lens-like or

wedge-hke prism shape (Carter and Clark. 1985).

DISCUSSION

SEM examination of fractured and sectioned shells of

deep-.sea mussels targeted in this study reveals similar

microstructures in distinct arrangements. The shells of

mytilitis inhabiting both cold-water sulfiile/methane seep

and deep-sea hvdrothermal vent environments are con-

sistently composed of outer fibrous prismatic and inner

nacreous stnictures. which underlie a periostracal laver.

Simple prismatic structure forming the adductor and

pallial mvostraca are also present. The microstruitures

in these specimens exhibit similar complexity to those of

the previously described vent mussel ButhijiiuHliolus

tlwriitopliihis.

Based on shell microstnicture, Ta\lor et id. (1969)

identified 2 groups of species belonging to the Mvtilacea:

M. T. Kennish et al.. 1998

Page 89

(1) a t\vo-la\'ered, whoIK- nacreous, \\'hol]\' aragonitic

warm-water group (e.g., Pcnw liiidis (Linnaeus, 1758),

Choroimjtihis palliopunctatus (Carpenter, 1857), and

Pcniinytihis puijniratiis (Larmarck, 1819): and (2) a

two- or three-la\ered aragonitic and c;ilcitic, temperate

group (e.g., Mijtihts cduUs (Linnaeus, 1758) and Mytilui

caUforniamts (Conrad, 1837)). In the latter group, the

prismatic lavers consist of calcite, whereas the nacreous

layers are composed of aragonite. Bet^\een the outer ciil-

citic prismatic and tlie inner nacreous la\ers, a middle

aragonitic prismatic \a\ex is present in some mvtilid

shells (Blackwell et al.,' 1977; Carter, 1980a). Lutz and

Rhoads (1980) and Lutz (1982) recognized 4 distinct las-

ers in the vent mussel Bothi/mocliohis thcnnophilus. in-

cluding (1) fibrous prismatic calcite, (2) nacre (arago-

nite). (3) irregular prismatic aragonite (pallial niNostrac-

um), and (4) nacre (aragonite).

The shell microstructures in Bathyinocliolii.s tlienno-

phihis are most similar to those in species of Mytihis.

Idasola, and Modiolus (Carter, 1990). According to Car-

ter (1990), the outer shell la\er of B. thcntiophihis is

calcitic with outer homogeneous and inner, predomi-

nantly fibrous prismatic sublayers. The middle and inner

shell layers are largeh' comprised of nacre, with minor

irregular simple prismatic structure. Hence, the findings

of Carter (1990) corroborate those of Lutz and Rhoads

(1980) and Lutz (1982).

Carter (1990) noted that the outer shell layer of m\-

tilids in general shov\s considerable mineralogical and

microstructural yariabilitv. In studies of m\tilids from

the Carboniferous to Recent, some species exliibit a cal-

citic fibrous outer shell la\er, whereas other species ha\e

various, different calcitic and/or aragonitic prismatic, ho-

mogeneous, and even minor crossed structures in this

layer. Waller (1978) generalized that a calcitic outer shell

layer in the M\tiloida, where present, consists of regular

fibrous prismatic structure. Carter (1990), however,

identified various combinations of aragonitic and/or cal-

citic irregular simple prismatic, irregular spherulitic pris-

matic, irregular fibrous prismatic, regular fibrous pris-

matic, and homogeneous structures in the outer shell

lavers of this order.

The shell microstructures of the 6 undescribed mxtilid

species investigated in this stud\' contain 4 calcified shell

layers composed of aragonite and calcite, although dif-

ferences in the layers can be significant owing to various

combinations of aragonitic and/or calcitic structures

present within the shell la\ers. The arrangement of shell

layers can also vary appreciably from one part of a mv-

tilid shell to another, because of alternating adductor

myostraca, dissolution, or other effects. Thus, in the as-

sessment of shell microstructure patterns, careful ex-

amination of the entire shell is necessary, as conducted

in this stutK'. These differences in shell microstructure

appear to be of great potential \alue in s\stematic stud-

ies of nn+ifids from deep-sea h\drothermal-vent and

cold-water sulfide/methane seep environments.

.ACKNOWLEDGMENTS

This is Contribution No 98-05 of the Institute of Marine

and Coastal Sciences, Rutgers Universit\', supported by

New Jersey state funds, the Deep-Sea Ecology- and Bio-

technologv' Center, and National Science Foundation

grants OCE 89-17311, OCE 92-17026, and OCE 96-

02205. We thank Richard Custafson for collecting some

of the mxtilid specimens and Alan Poole)' for help with

the illustrations.

LITERATURE CITED

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uitm.stnictiire iii h\o subspecies of tlie ribbed mussel, Geu-

kcu.sia tlciuissn (DilKwi, 1817)), Biologiciil Bulletin 1.52:1-

10.

Caiter. J. G. 19S0a. Guide to bi\al\e shell microstnictures. In:

Rlioads, D. C. and R. A. Lutz (eds.) Skeletal Groiiih of

Aquatic Organisms. Plenum Press, New York, pp. 64.5-673.

Garter, J. G. 1980b. Selected mineralogical data for tlie Bi\a]\ia.

7/1 RlioatLs. D. C. imd R. A. Lutz'(Eds.) Skeletal Gnmth of

Acjiiatic Organisms- Plenum Press, New York, pp. 627-643.

Garter, J. G. 1990. Glossaiv of skeletal biominer;ilizatiou. In: Gar-

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and Eiolutionan/ Trends. Van Nostnuid Remhold, New

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genetic significance of moUusciUi shell ultnistnictiire. In:

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cold seep emironments. Malacolo0a 37:.36.3-^373.

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comvid clams from h\dr(3tliemi;J \ent luid cold seep emi-

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19.5-200,

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species of bivalves from deep-sea Inclrotheniial vents. Anier-

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Lutz, R. A. iuid D. C., RlioacLs. 1980. Shell structure, mineralogy,

iuid niicromoq^hologv of deep-sea dieniial vent bi\;ilves

from tlie Galapagos Vtik: ecologiciil implications. Proceedings

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MacClintock, C. 1967, Shell stnicture of patelloid mk\ bellero-

phontoid gastropods (Mollusca). Bulletin of die Peabcxfv

Museum of Natural Hi.story 22:1-140.

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and mineralogv" of die Bivalvia: Introduction. Nuculacea —

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iv). Zoology Supplement .3:80—85,

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THE NAUTILUS 112(3):9()-94, 1998

Page 90

Molluscan Taxa and Bibliography of Henry van der Schalie

Mark E. Gordon

Zoolog\' Section

UniversiU- ol Colorado Museum

Campus Box 315, Hvuiter Building

Boulder, CO 80.309 USA

and

New Mexico Museum of Natural

Histor\' and Science

1801 Mountain Road NW

Albuquerque, NM 87104 USA

Henrs van der Schalie was bom in Amsterdam, The

Netherlands, on Jamiarv 8, 1907. His family immigrated

to the United States in 1909. following the death of his

father, and settled in Paterson, New Jersey. Van left New

Jersey in 1925 to enroll in Calvin College, Grand Rapids,

Michigan. During his undergraduate studies, he devel-

oped an interest in parasitology-. After receiving his A.B.

in 1929, Van began graduate studies in parasitologN' at

the University of Michigan (UM) under George R.

LaRue, Chair of the Department of Zoologx'. But before

long he accepted an assistantship in and transferred to

the Mollusk Division of the Museum of Zoologs at UM.

He received an M.S. in 19.31 and Ph.D. in 1934. Thus

began a career in malacolog>' at the Museum of Zoolog)-

that extended from his initial appointment as a museum

assistant in 1929 to his retirement as curator of mollusks

and professor of zoolog\' in 1977. A review of activities

in the Division of Mollusks during this period is pre-

sented in van der Schalie (1981).

Van became assistant curator of mollusks in 1934 and

was promoted to curator of mollusks in 1944 fcjllowing

the retirement of Calvin Goodrich. Initially appointed as

an instructor of zoology (1937-1944), he later held as-

sistant (1944-19.50). associate (1950-1956), and full pro-

fessorships (1957-1977) in the Department of Zoology,

UM, and served in visiting or adjunct capacities at other

institutions. With the onset of World War II, his research

returned to parasitologv. He studied mollusks as vectors

for various diseases, particularlv schistosomiasis. Subse-

(juently, he ser\ed on numerous national and interna-

tional committees and as a consultant to various inter-

naticmal organizations involved in medical malacology.

Van was widely known as a researcher, tnlucator, and

advocate in malacology, parasitology, environmental con-

servation, and general education. He received many re-

search grants and was member of numerous professional

societies. In recognition for his earlv contributions to

malacology, the fossil alasniidontiue (Hivalvia: Uiiiono-

idea) taxon, Vonder.schalica Modell (1943: 112; tvpe-spe-

cies: Unio kolasii Modell, 1931), was named after him.

Although retiring as curator emeritus from the Univer-

sitv' of .Michigan in 1977, he did not cease his profes-

sional interests and continued his research and advcxacy

until his death on 15 April, 1986.

In adthtion to data in van der Schalie (1981), bio-

graphic information has appeared in Kraemer and Berrv'

(1986) and Abbott (1987); however, none of these ref-

erences listed or reviewed his publications or the taxa

that he described. Van der Schalie published 142 manu-

scripts during the period of 1932-1986. Several of these

works were published initially as a summary or an ab-

stract, with the complete version of the paper pubhshed

later in a different journal, A few papers were reprinted

in other journals; in the following bibliography, the sub-

secjuent printing is noted in parentheses after the orig-

in;il reference. Between 19.34 and 1939, Van co-authored

3 new species of prosobranch (Hydrobiidae) and 4 pul-

monate (1 Ancylidae, 2 Oleacinidae. 1 Sagdidae) gastro-

pods from Guatemala with his mentor Calvin Goodrich,

and described 3 new species of unionoid bivalv es (2 from

the United States, 1 from Canada).

MOLLUSCAN TA>L\

Abbreviations: CM = Carnegie Museum. Pittsburg,

Pennsylvania; UF = Florida Mu.seum of Natural Histon',

University of Florida, Gainesville; UMMZ = Museum

of Zoology, University of Michigan, Ann Arbor

(i^iiailtic. Fcnis.sin (Laevapex) Goodrich and van der Schalie,

1937 (1.5 March), Miscellaneous Publications, Museum of

Zoologv', Universitv- of Michigan 34:23, pi. 1, fig. 7-7a ;

type localitv': Aguada de Copo, one mile south of La Lib-

ertad, Petc^n, Guatemala. Holotxpe: UM.MZ 6.5583. The

generic allocation of this species is luicertain.

aharadoi. Spiraxis Goodrich and van der Schalie, 19.37 (15

March). Miscellaneous Publications, .Museum of Zoologv,

Uui\ersitv of Michigan 34:23, pi. 1, fig. 1: hpe localitv:

limestone knoll five miles north of El Pa.so de los Caballos,

Peten, Guatemala. Holohpe: UMMZ 65165.

brouksiana, Anodonta van der Schalie, 1938 (4 June), Annals

of the Carnegie Museum 27:167, pi. 16; tvpe locality:

Spout Pond Ann, Fcrrviand District, Southern Shore,

Newfoundland. Holotvpe: CM 61.13137. The taxonomic

status of this .species is not clear. {\\. R. Hoeh, personal

conumuiication).

cicnchi. S()»w/(igi/n/.v (ioodrich and van der Schalie, 1937 (15

March), Miscellaneous Publications, Museum of Zoologv;

Univer.sitV' of Michigan 34:.37, pi. 1, fig. 6: tvpe localitv:

Ri'o de la Pasicin, S;iyaxclu'', Petc^n, Guatemala, Holotvpe:

M. E. Gordon, 1998

Page 91

UMMZ 65371. Aroapi/n^iis ciciichi {fide Hershler and

Thompson, 1992).

franccsac. Cochliopii Ciootlritli and \an der Schalie, 1937 (15

March), Miscellaneon.s Pnbhcations, Museum ot Zoolog\',

Universih' of Michigan 34:3S, pi. 1, fig. 3; Upe localiU':

Ri'o de la Pasion, at the mouth of Arrovo Chajchim'c,

northwest of Ponenir, Alta Vera Paz, Guatemala. Holo-

t\pe: UMMZ 6533S. Cochliopiiui franccsac [fide Hershler

and Thompson, 1992).

ftinibiis. Spiraxis Goodrich and \an der Schalie, 1937 (15

March), Miscellaneous Publications, Museum of Zoolog\;

Universit\' of Michigan 34:23, pi. 1, fig. 2; t\pe localit\':

limestone kiioll fi\e miles north of El Paso de los Caballos,

Peten, Guatemala. Holot\pe: UMMZ 65166.

jonesi, LampsiUs van der Schalie, 1934 (4 Mav), The Nautilus

47(4): 125, pi. 15, fig. la-b, 2, 3a-b; type localit\-: Pea River

Preston's Mill, Dale County, Alabama. Tvpe material orig-

inally was deposited in the Alabama Museum of Natural

Histor)': however the malacological collection of that nni-

seum was given to the F"lorida Museum of Natural Histon

with se\'eral lots subsequentlv transferred to the Museum

of Comparatise Zoolog\-, Harvard Uni\ersit\' (KG,

Thompson, personal communication). Lectotvpe: UF

6555S, designated by Johnson (1967; the specimen se-

lected was the male figured in van der Schahe. 1934).

Pti/chobrnnchtis jonesi (fide Atheam, 1964),

mcglaineriac. Medionidiis van der Schalie, 1939 (24 June), Oc-

casional Papers, Museum of Zoologw University of Mich-

igan 407:1, pi. 1; hpe locality-: Tombigbee River Epes,

Sniiipter Count^, Alabama. Holot\pe: UMMZ 130460.

pa-iioncnsi.s. Aiiinicold Goodrich and van der Schalie, 1937 (15

March), Miscellaneous Publications, Museum of Zoologw

Universitv of Michigan 34:36, pi. 1, fig. 4; tvpe localih':

Arroyo Subin, tributarv- of the Ri'o de la Pasion, about two

miles above Santa Teresa, Peten, Guatemala. Holotvpe:

UMMZ 65357. Aroapijraus pasionensis (fide Hershler and

Thompson, 1992).

taintori. Xenodi.scuin Goodnch and van der Schalie, 1937 (15

March), Miscellaneous Publications, Museum of Zoolog\,

University- of Michigan .34:26, pi. 1, fig. 5, 5a-b; hpe lo-

cality; woodland just east of El Paso de los Caballos, Pe-

ten, Guatemala. Holotvpe: UMMZ 65177.

BIBLIOGRAPHY OF HENRY VAN DER SCHALIE

van der Schalie, H. 1932. The station of Diisnoiniii tiicjiutra

(Raf ). The Nautilus 45(3):104-105.

Goodnch, C. and H. van der Schahe. 1932. I. On an increase

in the naiad fauna of Saginaw Bav, Michigan; II. The naiad

species of the Great Lakes. Occasional Papers, Museum

of Zoology. University of Michigan 238: 1-14.

van der Schalie, H. 1933. Notes of the brackish water bivalve,

Pohjmesoda caroUniana (Bosc). Occasional Papers, Mu-

seum of Zoologv', Universih- of Michigan 258:1-8.

van der Schalie. H. 1934. Lampsilis jonesi. a new naiad from

southeastern .i^labama. The Nautilus 47(4):12.5-127.

van der Schalie, H. 1936a. Transposed teeth of North Amer-

ican naiades. The Nautilus 49(3):79-S4.

van der Schalie, H. 1936b. Ovovi\iparitv among mollusks. The

Nautilus 50(l):lf-^19.

van der Schalie, H. 1936c. The naiad fauna of the St. Joseph

River drainage in southwestern Michigan. The American

Midland Naturalist 17(2);52.3-527.

van der Schalie, H 1936d. An unusual naiad fauna of a south-

em Michigan lake. The American Midland Naturalist

17(3):626-628.

van der Schalie, H. 1937a. A mussel taken from the stomach

o{ Rana catesbiana Shaw. The Nautilus 50(3): 104-105,

Clench, W.J. and H. van der Schalie. 1937. Lampsilis cariosa

(Say), The Nautilus 50(3):105.

van der Schalie, H. 1937b. K method lor shippmg freshwater

mollusks. Basteria 2(1):4— 6.

van der Schalie, H. 1937c. The relationslnp of the gravid pe-

riods ot certain nuissels in Michigan to the pearl button

industn-. Transactions of the American Fisheries Society'

66:406^10.

Goodrich, C, and H. van der Schalie. 1937. .Vlollusca of Peten

and north Alta Vera Paz, Guatemala. Miscellaneous Pub-

lications, Museuui of Zoologv, Universitv of Michigan 34;

1-50.

van der Schalie, H. 1938a. The naiad fauna of the Huron

River, in southeasteni ,Michigan. Miscellaneous Publica-

tions, .Museum of Zoologv, Universitv of Michigan 40:1-

83.

van der Schalie, H. 1938b. On the occurrence of Helix lactea

Miiller in North America. The Nautilus 51(4):132-1.34.

van der Schalie, H. 1938c. Remarks on some of Dr. Paul

Bartschs experiments. The Nautilus 51(4):134-135.

van der Schalie, H, 193Sd. Review of Ovster culture and ovs-

ter biology-, b\ Orton. Copeia 1938:52.

van der Schalie, H. 1938e. Anodonta brooksiana. a new naiad

from Newfoundland. .Annals of the Carnegie Museum

27(12): 167-170.

van der Schalie, H. 1938f. Hitch-hiking mussels and pearl but-

tons. Michigan Conservation 7(Jiine):-l-5.

van der Schalie, H. 1938g. Contributing factors in the deple-

tion of naiades in the eastern United States. Basteria 3(4):

51-57.

van der Schalie, H. 1938h. The naiades (fresh-water mussels)

of the Cahaba River in northern .\labama. Occasional Pa-

pers, Museum of Zoologv-, Universitv of Michigan 392:1-

29.

van der Schalie, H. 1939i. Hendersonia occidta (Sav) in Mich-

igan: its distribution, ecology-, and geographical signifi-

cance. Occasional Papers. Museum of Zoologv, Universitv

of Michigan .399:1-8.

van der Schalie, H. 1939j. Mcdionidus nicglantciine. a new

naiad from the Tombigbee River with notes on other nai-

ads of that drainage. Occasional Papers, Museum of Zo-

ology, Universitv of Michigan 407:1-6.

van der Schalie, H. 1939k. Distributional studies of the naia-

des as related to geomoiphologv. Journal of Geonioq-)hol-

o,g\- 2(3);251-257.

van der Schalie, H. 19391. Additional notes on the naiades

(fresh-water mussels) of the lower Tennessee River The

American Midland Naturalist 22(2):452-457.

van der Schalie, H. 1939in. Aquatic mollusks of the upper

peninsula of Michigan, part II; the naiades (fresh-water

mussels). Miscellaneous Publications, Museum ofZoologv.

Universih- of Michigan 43:3.5 — 15.

van der Schalie, H. 1940a. Larger land snails from pine woods

in northern Michigan. Papers of the Michigan .\cadcmv

of Science, Arts, and Letters 25:367-370.

van der Schalie, H. 1940b. Notes of Mollusca from Alta Vista

Paz, Guatemala. Occasional Papers, Museum of Zoologv-,

University^ of Michigan 413:1-11.

van der Schalie, H. 1940c. The naiad fauna of the Chipola

River, in northwestern Florida. Llovdia 3(3);191-208.

Page 92

THE NAUTILUS. \'oI. 112. No. 3

van der Schalie. H. 1940d. .^estivation of Li/innacii l/iiicctitii

(Gould). The Nautilu,s 53(4):134-135.

van der Schalie, H. 1940e. .\e.stivation of fresh-water mussels.

The Nautilus 53(4): 137-138.

van der Schalie. H. 1941a. Zoogeograph\ of naiades in the

Grand and Muskegon ri\ers of Michigan as related to gla-

cial histor\'. Papers ot the Michigan Academy ot Science,

Arts, and Letters 26;29T-,310.

van der Schalie, H. 1941b. The taxonomy of naiades inhabiting

a lake environment. Journal of Concholog>'21(8):246-253.

van der Schalie, H. and F. Locke. 1941. Hermaphroditism in

Anodonta grandis. a fresh-water mussel. Occasional Pa-

pers, Museum of Zoologv', Universih^ of Michigan 432:1-

7.

van der Schalie, H. 1941c. On collecting fresh-water mussels.

Annual Report of the .American Malacological Union tor

1940: 9-14.

Clench, W.J. and H. van der Schalie. 1944. Notes on naiades

from the Green, Salt, and Tradewater rivers in Kentucky.

Papers of the Michigan Acadenu' of Science. Arts, and

Letters 29:22.3-228. "

Goodrich, C. and H. van der Schalie. 1944. A revision ot the

Mollusca of Indiana. The American Midland Naturalist

32(2):257^326.

van der Schalie, H. 194.5a. The value ot mussel distribution in

tracing stream confluence. Papers of the Michigan Acad-

emv of Science, .\rts. and Letters .30:.3.5.5-.373.

van der Schalie, H. 1945b. What has happened to the genus

lo? Mollusca 1:59-61.

van der Schalie, H. 1947. The ecology' of molhisks for the

biology teacher American Biology Teacher 9(6): 174-176.

van der Schalie. H. 194Sa. The Michigan pearl button indus-

try. Annual Reports of the .American .Malacological Union

for 1947:8 (repnnted m 1963. Sterkiana 9:29).

van der Schalie, H. 1948b. The land and fresh-water mollusks

of Puerto Rico. Annual Report of the American Malaco-

logical Union for 1947:18 (reprinted in 1963, Sterkiana 9:

29).

van der Schalie, H. 1948c. The commerciallv \aluable mussels

of the Grand River in Michigan. Miscellaneous Publica-

tions of the Michigan Department of Consen ation Insti-

tute of Fishery Research 4:1—12.

van der Schalie, H. and E.G. Berry. 1948. A(|natic puliiionates

from Lake Tahoe. The Nautilus 62(1):.3— 1.

van der Schalie, H. 1948d. The land and fresh-water mollusks

of Puerto Rico. Mi.scellaneous Publications, Musemri of

Zoologv'. Universih of Michigan 70:1-1.34.

van der Schalie, H. 1949a. The fresh-water mussels of the

Mississippi River from St. Paul to the mouth of the Ohio

River. Annual Reports of the American Malacological

Union for 1948:16 (reprinted in 1963, Sterkiana 9:29).

van der Schalie, H. 1949b. Biographic notes on .\rnold Ed-

ward Ortmami as revealed b\ some of his letters. .Annual

Reports of the American Malacological Union lor 1948:

11-12 (reprinted in 1963, Sterkiana 9:.3()).

van der Schalie, H. 1950. Another techni(]ne for the prepa-

ration of radulae. Ainnial Reports of the American Mala-

cological Union for 1949:8.

Rogick, D. aTid H. van der Schalie. 19,50. Studies on fresh-

water Brvozoa. Ohio Journal of Scicnee 50:1.36-146.

van der Schalie. H. and A. van der Schalie 1950. The mussels

of the Mississippi River The .American Midland Naturalist

44(2):448-466.

\an der Schalie. H. 1951a. Arnold Edward Ortmaim as re-

vealed by his letters. The Nautilus 64(4):I.34-141.

van der Schalie, H. 19.51b. .Arnold Edward Ortmann as re-

vealed by his letters-II. Tlie Nautilus 65(l):23-26.

van der Schalie. H. 19.52a. .An old problem in naiad nomen-

clature. .Annual Reports of the .American Malacological

Union for 1951:4-5 (reprinted in 1963, Sterkiana 9:.30),

van der Schalie, H. 1952b. ,An old problem in naiad nomen-

clature. The Nautilus 65(3);9:3-99.

van der Schalie, H. 1953a. Ecology and distribution of Lijitt-

nacii (Bidimnca) iiicga.soina in Michigan, .Anniuil Reports

of the American Malacological Lhiion. for 1952:14 (re-

printed in 1963, Sterkiana 9:31).

van der Schalie. H. 19.53b. Interim report: bilharzi;isis control

project at Qiil\iib, Egypt. United Nations World Health

Organization EM/BILy6:l-41.

van der Schalie. H. 1953c. Mollusks from an interglacial de-

posit (Sangamon age?) in Mead Counts; Kansas, The Nau-

tilus 66(3):80-90.

van der Schalie, H. 1953d. Nembutal as a relaxing agent for

mollusks. The American Midland Naturalist 50(2):511-

512.

van der Schalie, H. 19.54a. Bilharziasis in Eg\pt The Biologist

36(l-4):3;^.34.

\'an der Schalie, H, 19.54b. Obser\ations and report on a pro-

gram for bilharziasis control in the Gezira irrigation area

of the Sudan. United Nations World Health Organization

EM/BIL/5:l-5.

van der Schalie, H. 19.54c. Eg\pt's fight against bilharziasis.

WHO Newsletter 7(1 ):4.

van der Schalie, H. 19.54d. Cakin Goodrich, 1874-1954. Re-

port to the Director of the Museum of Zoologv, Universih'

of Michigan 56:9-14 (reprinted in 1955. The Nautilus

68(4):1.35-I40).

van der Schalie, H. 1954e. The problem with snail control in

Egypt. Report of the Director of the Museum of Zoology,

Universitv of Michigan 56:35-38.

van der Schalie. H. 195.5a. Bilharziasis control project at Qal-

Mib. United Nations World Health Organization EM/BIIV

6:1-41.

\an der Schalie. H. 19.5.5b. Discussion of the paper bv L.J.

Olivier. American Journal of Tropical Medicine and Hy-

giene 4(3):424-425.

\an der Schalie, H. and D. S. Dundee. 1955. The distribution,

ecolog\', and life histor\' of Painatiupsis cinciniiaticn.sis

(Lea), an amphibious operculate snail. Transactions ot the

American Microscopical Socieh' 74(2): 119-133,

\'an der Schalie, H. 1955c. Calvin Goodrich (obituary). .Annual

Reports of the .American Malacological Union for 1955:1.

\an der Schalie. H 195.5d. Bilharziasis in Eg\pt. .Annual Re-

ports of the .Amencaii Malacological Union for 1955:23.

van der Schalie. H, 19.56a. Facilities for training malacologists

and medical zoologists in the Uni\ersit\' of Michigan.

United Nations World Health Organization Bilharziasis

Conference 2:1-5.

van der Schalie, H. and D. S. Dundee. 1956. The moiphology

of Pomatioj)si.s cincinnaiiensus (Lea), an amphibious pro-

sobranch snail. Occasional Papers. Museum ot Zoologv;

Uiii\ersit\ of Michigan .579:1-17,

\an der Schalie, H. 19.56b. Human blood fluke (schistosomi-

asis) control in Egxpt. Annual Reports ot the .American

Malacological Union for 1956:9.

van der Schalie, H. 19.56c. Studies on Ponmtiojtsis shells in

M. E. Gordon, 1998

Page 93

Michigan. Annual Reports of the American Malacological

Union for 1956:22.

1 der Sehalie, H. 1957. Concerning the t'uhire of The Bi-

ologist. The Biologist 40(1-2): lT.'"2.S-29.

1 der Sehalie, H. and H. J. Walter 1957. The egg la\ing

habits of Pomatiopsis cinciniinticnsis (.Lea). Transactions

of the American Microscopical Society- 76(4):4()-l — 122.

I der Sehalie. H. 195Sa. Problems in control of intermediate

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I der Sehalie, H, 1960b. Pearls, food, and buttons: practical

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1 der Sehalie, H. and P. W. Parniiilee. 1960. .\nimal remains

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1 der Sehalie, H. 1961a. Peter Olaus Okkelberg, 1880-1960.

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1 der Sehalie, H. 1961c. The naiad (fresh-water nnissel) fau-

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1 der Sehalie, H. and L. L. Getz. 1962c. Reproductive iso-

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van der Sehalie. 1962. Mu.ssel distribution in relation to stream

confluence in northern .Michigan. Annual Reports of the

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\an der Sehalie, H. and G. C. Robson. 1963. Bixalve. Five

pages in Enc\clopffdia Britannica

van der Sehalie, H. 1963a. Mussel distribution in relation to

former stream confluence in northern Michigan, U.S.A.

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van der Sch;Jie, H. 1963b. People and their snail-bome dis-

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van der Sehalie, H. 1963c. Comparison of temperature and

moisture responses of the snail genera Pomatiopsis and

Onconulania Ecolog\' 44(l):7.'3-83.

\an der Sehalie, H. 1963d. Laboratorv guide and notes for

medical malacolog\- bv Emile A. Malek (book review).

.\merican Journal of Tropical Medicine and Hygiene

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bution, ecolog)', and life histors' of the mussel, Arf»io/iflifl.s

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van der Sehalie, H. 1963e. Studies of the biolog\- of Poma-

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van der Sehalie, H. and G. m! Da\is. 1964. Stunting of On-

comclania fonnosa in culture. Annual Reports of the

American Malacological Union for 1964:13-14.

van der Schahe, H. 1964. Notes on die sex of Campclonm.

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seum of Zoology, University of Michigan 641:1-15.

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van der Sehalie, H. 1965b. Second European Malacological

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van der Sehalie, H, 1966a. The role of snail intermediate hosts

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van der Sehalie, H. 1966b. Hermaphroditism among North

American freshwater mussels. Malacologia 5(1): 77-78.

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van der Sehalie, H. 1969c. Man meddles with nature — Ha-

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van der Sehalie, H. 1969d. Snail control problems in Hawaii.

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THE NAUTILUS, Vol. 112. No. 3

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1969;5.5-.56.

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9(1):44 (the vohmie pubhcation date for this and tlie fol-

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van der .Schalie, H. 1970b. .American nnissel resonrces m re-

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vail der Schalie, H. 1970c. Book re\ie\v: A guide for the iden-

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in the .Americas. Pan American Health Organization Sci-

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van der Schalie. H. 1970d. Hermaphroditism among North

American freshwater nuissels. Malacologia 10(1):9.'3-112.

van der Schalie. H. 1970e. .Mussels in the Huron River above

Ann Arbor in 1969. Sterknana .39: 17-22.

van der Schalie. H. 1971. Review: Terrestrial slugs by N.W.

Riinham and PJ. Hunter. Science 17.3(4000):904. '

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Natural Historv Press, Garden Citv, pp. 116-1.36.

van der Schalie, H. 1972b. Dam(n) large rivers. . .then what?

Michigan .Alumnus (April):10-12 (reprinted in 1973, The

Biologist .55(2):29-35).

van der Schalie, H. 1972c. Problems in culturing snail inter-

mediate hosts. Malacological Review .5(1): 10.

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temperature on growth and reproduction in aijuatic snails.

Malacological Review 6(1):60.

Liang, Y.-S., H. van der Schalie, and E. G. Beny. 1973. Trans-

mission of ostracods in snails. Malacologiciil Review 6(1);66.

van der Schalie, H. and E. G. Berry. 1973b. Effects of tem-

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Environmental Protection Agency Research Series, U.S.

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<><lhi])hop.sis (ipciiti Temcharoen, a new snail host lor

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Liang, Y.-S. and H. van der Schalie, 1975b. (;ulti\'ating Fii.s-

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ciolfi luptitica. Malacological Revii-w 8(1-2): 123.

van der Schalie, H. 1975. An ecological approach to rare and

endangered species in the Great Lakes region. Michigan

Academician 8(l):7-22.

van der Schalie, H. and II. D. Blankespoor. 1976. Attachment

and penetration of miracidia obsened bv scanning elec-

tron Tiiicroscop)'. Science 191 (4224):291 -1293.

viui der Schalie, H. 1977. Malacolog)- and [)urasitolog\- as the\-

pertain to health. Bulletin of the .American Malacological

Union for 1976:.5.5-57.

van der Schalie, H. and H. D. Blankespoor. 1977. Potential use

of solar energy for snail-control. The Biologist 59(1): 16-24.

van der Schalie, H, 1978. Flukes that affect world progress.

Wards Bulletin 17(l):l-,3

van der Schalie, H. 1979. Swimmers itch. Literature. Science,

and .Arts (Uui\ersitv of Michigan) 2:-l— 7.

van der Schalie, H 1981a. Is professional training too restric-

tive, or can one earn a living working on snails? Spark

(Cal\in College), September: 12-14.

van der Schalie, H. 1981b. Fiftv years of malacologv' at the

University of Michigan (1929-1979). BulleHn' of the

American Malacological Union for 1980:1-5.

van der Schalie. H. 1981c. A.swan revisited. BioScience 31(6):

420.

van der Sch;ilie, H. 1981d. Pitst, present, ;md fiiture status of die

MoUusca of the upper Tombigbee River. Sterkiana 71:8-11.

van der Schalie, H. 1981e. Molhisks in the .Alabama River

drainage: past and present. Sterkiana 71:2-1—40.

van der Schalie, H. 1986. Mussels in the Detroit River op-

posite Cobo Hall. Michigan Academician 18(l):31-.34.

ACKNOWLEDGMENTS

John B. Burch and Renee A. Sherman (MZUM), Walter

R. Hoeh (Department of Biological Sciences, Kent State

University', Ohio), and Fred G. Thompson (Florida Mu-

seum of Natural Hi.storv, Unixersitv of Florida) provided

much appreciated assistance and information during the

preparation of this manuscript. M.G. Harase\v)'ch and

Jose H. Leal, respectiveh' past and present editors of

The Nautilus, and two anonvinous reviewers provided

direction that improved the manuscript. Finally, this

manuscript would have been considerably more difficult

to prepare if it had not lieen for the large stack of re-

prints and bibliographies given to me bv Van in 1979

during m\ first visit to the Museum of Zoologv', UM, and

subsequent coninnmications.

LITERATURE CITED

.Abbott, R. T, ed. 1987. Register of American malacologists: a

national register of professional and amateur malacologists

and private shell collectors, second edition. .American Ma-

lacologists, Inc., Melbourne, Florida. 168 pp.

Atheani, H. D. 1964. Three new uniouids from .Alabama and

Florida and a note on Lnmpsili.s /o/ics; The Nautilus

77(4):134-139.

Hershler, R. and F. G. Thompson. 1992. .A review of the ai|U.it-

ic gastropod subfamilv Cochliopinat' (.Prosobranchia: Hy-

drohiidae). Makicological Review, supplement 5:1-140.

|olinson. R. I, 1967, .Adtlitions to the nnionid fauna of the Gulf

diaiuage of .Alabama, Georgia, and Floritla (Mollnsca: Bi-

valvia). Breviora 270:1-21.

Krai'Mier, L, R. and E. G. Beri-v 1986 Things trulv excel-

lent. . .": Henn' van der Schalie. 1907-1986. The Nautilus

1()0(4):116-119.

.Modell, H. 1943. Tertiare Na|aden III Archiv Inr .\Iollusk<n-

kunde 75:107-117.

van der Schalie, II. 1981. Fiftv vears of malacologv at the Uni-

versitv of Michigan (1929-1979). Bullelm of the .Anx'ncan

Malacological Union, for 1980:1-5.

THE NAUTILUS 112(3):95-98, 1998

Page 95

A New Favaiiia (Miirexiella) from the Panamic Province

(Gastropoda: Muricidae) and Designation of a Lectotype for F.

(M.) exigua (Broderip, 1833)

Carole M. Hertz' and Barbara

W. Myers-

Santa Barbara Miist'uni nl Natural

Historv'

2559 Puesta del Sol Road

Santa Barbara. CA 93105 USA

ABSTRACT

A new species of Faiaiiia (Miircxiclla) is described (roni the

eastern Pacific Ocean and compared witli similar species F

(M.) cxigiia (Broderip, 1833) and F. (M.) lappa (Broderip.

1833). Favartia vcnustula Poorman. 1983, is tonnd to be a

junior synonym o( F (M.) cxi<i!i(i. and a lectot\pe is designated

for F. (M.) exigua.

Ki'ij words: eastern Pacific, new species, Costa Rica, Ecuador,

Mexico, Panama.

INTRODUCTION

Over ten \ears ago, Carol Skoglund of PhoenLx, Arizona,

brought to the Marine In\ertebrate Department of the

San Diego Natural Iliston Musemn se\eral specimens

of a species of Favaiiki for identification. Anthonx-

D'Attiho, then Acting Curator, and the authors examined

the specimens, noting that they were similar to F. cxif^tia

(Broderip, 183.3). But, at the time, no t\pe or compar-

ative material was available to us.

Since then, we have examined hvent\-hvo specimens

of this species from the Hertz, Kaiser, Koch, Shask\' and

Skoglund collections. We studied the type material of

the three more similar species, F.(M. ) cxit^uo, F.(M. ) rad-

icatii (Hinds, 1844), junior s\aiomnn of F. (A/.) lappa

(Broderip, 1833), and F. (M.) vemistiila Poorman. 1983,

as well as comparative material of other eastern Pacific

Favartia (Muirxiclla). As a result, we determined that

the specimens belong to an undescribed species.

The following abbreviations for institutions are used

in the te.xt: AMNH, American Museum of Natural His-

toPi'; BMNH, The Natural Histon' Museiun. London;

SBMNH. Santa Barbara Museum of Natural Histon:

' Mailing address: 3883 Mt. BKickbum A\e., San Diego, CA

92111, USA,

- Mailing address: 3761 Mt. Augustus Ave,, San Diego, CA

92111, USA.

SDNHM. San Diego Natural Histor\ Museum: USNM,

National Museum of Natural Historx, Smithsonian In-

stitution.

SYSTEMATICS

Family Muricidae Rafinesque, 1815

SubfamiK- Muiicopsinae Radwin and D'Attilio, 1971

Genus Faiaiiia Jousseaume, 1880

Subgenus Muivxiclla Clench and Perez-Farfante, 1945

Faiaiiia (Muirxiclla) panlskoghiiuli new species

(Figures 1-5)

Description: Shell small, up to 17.1 mm length X

11.2 nmi width, biconic, spire elongate. Protoconch of

three pale tan, pustulose turbinate whorls, buttressed on

last whorl. TeleocoTich of five whorls: suture indistinct;

shoulder sloping: six or seven thickened varices on liody

whorl, eight or nine on penultimate whorl. Leading edge

of varix foliate forming short, straight, open spines at

periphery spines occasionalK' slightK' recurved. Aperture

ovate, lip edge crenulate, lirate within reflecting spiral

cords. Inner lip erect along entire length, smooth within;

anal sulcus weakly defined. Siphonal canal of moderate

length, straight, narrovvK' open to right, weaklv recurved

distalK- with tAvo to four well-preser\'ed canal termina-

tions. Spiral sculpture of two strong cords on first three

teleoconch whorls: penultimate whorl with two strong

cords and a minor cord between, anterior to these are

hvo additional, minor cords; shoulder with two or three

minor cords. BoeK' whorl witli fi\e strong major cords,

interspaces each with a minor cord: gap beh\een major

cords and canal: hvo or three major cords on canal.

Shoulder of bod\- whorl with hvo to four minor cords.

Entire shell surface lamellose, forming webbing betsveen

cord terminations. All major cords divided along their

length b\' tour incised hues. Operculum muiicopsine,

imguiculate, annulate centrally witli basal nucleus. Shell

color cream. Orange-browai flush on varices, spine ter-

minations, and occasionally in intei'varical area.

Page 96

THE NAUTILUS. Vol. 112, No. 3

Figures 1-3. Faiaiiia {Murexielln) patil.skofilundi new species. Holohpe. SDNHM 7S066. 17.1x11.2 mm. Pedro Gonzales. Islas

Las Perlas, Panama, in 5.5 m. 1. Apertural view; 2. Dorsal view; 3. Paratope A, USNM S8026T. 8.5X5.1 mm. Isla la Plata, Ecuador.

Type material: Holotype: SDNHM 78066, 17.1X11.2

mm, t\pe localits-, R. Hubert coll., ex D'Attilio Collec-

tion, October 1971; Paratspes: Parat\pe A, USNM

8800267, 8.0X5.1 mm, Isla la Plata, Ecuador, 1°16'S,

81°05'10"W, D. R. Shasky coll., June 1979: Parat>pe B,

Shaskv Collection, 15.5X9.6 mm, same as Paratvpe A;

Paratope C. SBMNH 144458, 10.4X5.9 mm, off Isla Vi-

radores, Playas del Coco, Guanacaste, Costa Rica.

10°.34'N. 85°34'W, dredged 9-15 m, R. Koch coll.. 2-4

April 1986; Parat\pe D, Koch CollecHon, 10.2x6.0,

same as Paratvpe C; Parat\pe E, AMNH 290728,

11. 5X7. .3 mm, S of Isla Chitre, Islas las Perlas, Panama,

8°36'N, 79°4'W, dredged 15-18 m, C. and R Skoglund

and R. and W. Koch "coll., 15 April 1984; Parat\pe F,

Skoglund Collection. 13.4X7.9 mm, same as Paratxpe E;

Parat\pe G, Skoglund C'ollection, 12.2x7.8 mm, same

as Paratvpe E; Paratype H, Skoglund Collection.

11.6X7.8 mm, same as Paratype E; Paratvpe I, Shask-\

Collection, 16.2X10.5 mm, Isla Venado, Panama,

8°52'30"N. 79°39'3(AV. J. McDaniel coll.. 8 Septemlier

1979; Paratype J, Shaskv Collection, 12.9X7.8 mm. N

side of Isla Salango, Ecuador, 1°35'15"S, 80°52'52"W, D.

R. Shasky coll., 15 Septemi)er 1978; Paratvpe K. Koch

Collection, 11.5X7.2 nun. Isla Negritos Adentro. Golfo

de Nicoya, Costa Rica, 9°57'N, 84%2'W, dredged 12-27

m, C. and P. Skoglund and R. and W. Koch coll., 8 Ma\

1982; Paratype L, Koch Collection, 11.6X7.0 mm, Bah-

liis de Iluatulco. Oaxaca. Mexico, 15°40'N, 96°08'W.

dredged 9-30 m, R. and W. Koch , 2-4 June 1991; Para-

type M, Koch (Collection, 15.5X9.6 mm, same as Para-

tvpe E, 16 April 1984; Paratopes N-P; N, 14.6X9.0 mm;

Paratvpe O. 12.7X8.2 mm; P, 12.5X7.3 mm, data for all

same as Paratvpe E; Paratvpe Q. Skoglund Collection

10.1X6.9 mm,' off Isla Viradores Sur, Phivas del (loco.

Guanacaste, Costa Rica. 10°.34'N, 85°34'W, dredged 9-

18 m. C. and R Skoglimd coll., April 1986; Paratvpe R.

Skoglund Collection 12.1X8.2 mm, off Isla Rancheria,

Golfo de Chiriqui. Panama, 7°35'N, 8r40'W, dredged

24-36 m. C. and P Skoglund coll., March 1986; Paratvpe

S, Hertz Collection, 10.5X7.3 mm. Islas Tres Marietas,

Navarit, Me.xico, 21°41'N, 105°36'W. diving in 13.7 m,

K. L. Kaiser coll., 22 Febniarv 1995; Paratvpes T-U, K.

L. Kai.ser Collection: T, 12.6x7.9 mm; U, 1().9X7.4 mm.

Biiliia John Huston, Balii'a Banderas. Jalisco. Me.vico,

20°30.55'N, 105°21.11'W, 9 m. in sand under rocks. 2

Januar\- 1995. K. L. Kaiser coll.

Type locality: Pedro Gonzales, Islas las Perlas, Pana-

ma, 8°25'N, 79°05'W, 5.5 m depth.

Distribution: Favaiiia (Muivxiclla) ptiulskafihiiuli is

knowm to occur from Islas Tres Marietas, Na\arit, Mex-

ico, its most northern locality, with an intermittent dis-

tribution south to Isla Salango, Ecuador, in 5.5-36.0 m

depth.

Discussion: The new species was compared with the

three s\iitvpes oi Fovaifia {Miircxiclla) cxi^iui (BMNH

19841227,' Figures 6, 7). Yokes (1988) considered F. (M)

cxiiiuti to be a \alid species and referred to the illustra-

tion of the largest SMitvpe figured in Yokes (1984. pi. 2,

fig. 3), although she erroneousK' considered F (M.) rad-

icfitn as a .s\Tionym of F. (M.) cxiguo in the 1984 paper.

We have selected the largest specimen (16.2X9.4 mm)

of the .sMitvpe series of F (A/.) cxi'^ua as the lectotvpe.

Two smaller specimens (12.6X7.5 mm and 10.8X6.8

mm) are chosen as paralectotvpes. Favartia (M.) paul-

skofjltiiuli differs from F. (M.) cxig^ua in having an indis-

tinct suture and sloping shoulder with thickened varices,

whereas F. (M.) cxigtia has an impressed suture and

somewhat excavated slioukler with sliarpK' elevated var-

ices. The new species has five strong major cords with

one minor cord in each interspace, whereas F. (M. ) ex-

C. M. Hertz and B. W. Myers, 1998

Page 97

Figures 6, 7. Favaiiin (MiircxicUa) exigiia (Broderip, 1833),

BMNH 19S4122, sviihpe lot. Localin-: "PSalango". 6. Aper-

tural view. 7. Dorsal \iew. Largest specimen, 16.2X9.4 mm,

selected as lectot)pe. Two smaller specimens, 12.6x7.5 mm

and 10.8x6.8 mm, selected as paralecto^y-pes. Specimens pho-

tographed with kind permission of the trustees of The Natural

Historv Museum, London.

Figure 4, 5. Favaiiia (MiircxicUn) pniilskonliindi new spe-

cies. 4. Paratvpe A, camera lucida drawing of protoconch (1.1.

mm diameter). 5. Paratope A, camera lucida ilrawing of oper-

culum (3.5 mm length).

iiiua has fi\e strong cords with no minor cords in the

deeply-cut interspaces, the edges of tlie cords somewhat

overlapping the interspaces.

Since Yokes (1970) stated that 'the tspe of A/. Itipjxi

is no longer to be foimd," the new species was compared

with the holot)pe of F. (M.) radicata (Hinds, 1844)

(BMNH 1907.10,28.136, Figure 8), a junior synonym of

F. (M.) lappa (Broderip, 1833) (Radwin and D'Attilio,

1976; Fair, 1976) and comparative material of K {M.)

lappa in private collections. Favartia (M.) paulskoglundi

differs from F. ( M. ) lappa in the number of spiral cords

on the liod\- whorl, five major cords with strong minor

cords between on the new species, and five major cords

with two strong minor cords on the shoulder and no

minor cords on the bodv whorl on F. (M.) lappa. In F.

(M.) paulskog,hiiuli there are si.x to seven varices which

do not obscure the indistinct suture, whereas in F. (M.)

lappa the five varices project above and obscure the su-

ture. The type species of F. (M. ) radicata had no re-

maining protoconch.

C'oniparisons between the new .species and F. (M.)

vcnusfula (holotvpe, SDNHM 81610), show that F (M.)

vcnii.stnla and F (A/.) cxigiia are conspecific. Both spe-

cies are of comparable size with similar protoconchs ( F

(M.) cxigua with slightly more than 2.5 smooth, convex

whorls and F (M.) vcnusiula with 3 smooth, conve.x