THE NAUTILUS

Volume 111, Number 1

Januanj 29, 1998

ISSN 0028-1344

A quarterly devoted

to malacology.

Marine Bicloc

Woods Hole Or. '-

FEB 0 5 I9d8

■oods Hole, MA 025'i'

EDITOR-IN-CHIEF

Dr. M. G. Harasevvych

Division of Mollusks

National Museum of

Natural Histor)'

Smithsonian Institution

Washington, DC 20560

MANAGING EDITOR

Dr. Jose H. Leal

Bailey- Matthews Shell Museum

3075 Sanibel-Captiva Road

Sanibel, FL 33957

CONSULTING EDITORS

Dr. Rudiger Bieler

Department of Invertebrates

Field Museum of

Natural History

Chicago, IL 60605

Dr Arthur E. Bogan

Freshwater Molluscan Research

36 Venus Way

Sewell, NJ 08080

Dr. Robert T Dillon, Jn

Department of Biology

College of Charleston

Charleston, SC 29424

Dn William K. Emerson

Department of Living Invertebrates

The American Museum of Natural

History

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 Zoologj'

Harvard University

Cambridge, MA 02138

Dr James H. McLean

Department of Malacolog)-

Los Angeles County 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

Dn Paula M. Mikkelsen

Department of Living Invertebrates

The American Museum of Natural

History

New York, NY 10024

Dr Donald R. Moore

Di\asion of Marine Geology

and Geophysics

Rosenstiel School of Marine and

Atmospheric Science

Universit)' of Miami

4600 Rickenbacker Causeway

Miami, FL 33149

Dr. Gustav Paulay

Marine Laboratory

University of Guam

Mangilao, Guam 96923

Mr Richard E. Petit

PO. Box 30

North Myrtle Beach, SC 29582

Dr. Edward J. Petuch

Department of Geology

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

Harvard University

Cambridge, MA 02138

Dr. Geerat J. Vermeij

Department of Geology

Universit)' of Caliibrnia at Davis

Davis, CA 95616

Dr G. Thomas Watters

Aquatic Ecology Laboratory

1314 Kinnear Road

Columbus, OH 43212-1194

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T H E€7N AUT I L U S

CONTENTS

; —— Volume 111, Number 1

Woods Ho;e Oc(.-:-ir.cQr Januanj 29, nMS

L -^0 ISSN 0028-1344

FEB 0 5 1998

Riidiger Bieler

Paula M. Mikkeisen

Wo^rl- lio'^'. I

Ainmoniccra in Florida: Notes on the Smallest Living

Gastropod in the United States and Comments on Other

Species of OinalogxTidae (Heterobranchia)

John B. Wise

Moqihologv and Systematic Position of Rissoello caribaca

Rehder, 1943 (Gastropoda: Heterobranchia: Rissoelhdae)

13

Edward J. Petuch

The Molluscan Fauna ol the Wawa River Region, Miskito

Coast, Nicaragua: Ecolog\\ Biogeographical Imphcations,

and Descriptions of New Taxa

99

Jesiis Ortea

Jose Espinosa

The Protoconch of FastigicIIa caiiiwta Reeve, 1848

(Mollusca: Cerithiidae)

45

Notices 46

This is the first number of THE NAUTILUS pubhshed by The Bailey- Matthews Shell Museum. All correspondence

pertaining to subscriptions, purchase of back issues, and missing issue claims should be addressed to:

Dr. Jose H. Leal

THE NAUTILUS

P. O. Box 1580

Sanibel, FL 33957 USA

Phone: (941) 395-2233

FAX: (941) 395-6706

e-mail: [email protected]

Beginning with volume 112, Dr. Leal will become Editor-in-Chief of THE NAUTILUS. All new manuscript submis-

sions should be addressed to Dr. Leal at the above address.

THE NAUTILUS 111(1):1-12, 1998

Page 1

Ammonicera in Florida: Notes on the Smallest Living Gastropod

in the United States and Comments on Other Species of

Omalogyridae ( Heterobranchia)

Riidiger Bieler

Center for Evolutionar\' and

Environmental Biology

Field Museum of Natural History

Roose\elt Road at Lake Shore Dri\'e

Chicago, Illinois 60605, U.S.A.

[email protected]

Paula M. Mikkelsen

Department of Invertebrates

American Museum of Natural History

Central Park West at 79th Street

New York, New York 10024, U.S.A.

[email protected]

ABSTRACT

The first record of a species of Ammonicera in Florida, with

an additional record from Yucatan, Mexico, is presented and

the gross morphologN' of the living animal is described for

the first time. This smallest living snail in the United States

is identified as Ammonicera minoiinli.s Rolan, 1992, origi-

nally described from Cuba. Comparisons are made with

closely similar species, especially A. japonica Habe 1972, a

possibly conspecific form with known wide distribution in

the Pacific Ocean. Various taxonomic problems in the genera

Ammonicera and Omalog,ijra are addressed, and current

composition of the family Omalogyridae is discussed. Lists

of currently recognized omalogvrid species in the Atlantic

Ocean (including the Mediterranean Sea) are presented.

Key words: Florida Keys, Gastropoda, lower Heterobranchia,

Ammonicera. Omalogyra. Omalogyroidea, Atlantic Ocean. Sys-

tematics.

INTRODUCTION

The family Omalogyridae is a poorly known group of

extremely small marine snails. Placed in their own su-

perfamily Omalogyroidea, they are currently classified as

members of the unresolved "lower heterobranch" gas-

tropods (e.g., Haszpnmar, 1988; Bieler, 1992; Healy,

1993). Even the most basic taxonomic and distributional

information is sketchy for this group, with most faunistic

studies missing or dehberatelv omitting the usuallv less-

than-one-miilimeter size range of the adult shells. The

few studies that have concentrated on this group have

brought many new species to our attention, such as the

recent series of works with excellent scanning electron

micrographs by Sleurs of omalogyrids in Papua New

Guinea (198.3) and in the Republic' of Malcbves (1985b),

by Palazzi (1988) in the Mediterranean and Madeira,

and those by Rolan in the Cape Verdes (1991) and Cuba

(1992). Whether these areas are exceptional in their high

species diversitv- of omalogvrids is doubtful, although it

is surprising that no true omalogyrids were reported in

some Caribbean studies that otherwise dealt with min-

ute species (e.g., De Jong & Coomans, 1988; Rios,

1994).

In addition to their small size, omalogyrids have

anatomical features that set them apart from caeno-

gastropods with which they were usually grouped.

This led to early speculations about their systematic

position. Because of their unusual radular characters,

G. O. Sars (1878) placed the at the time monotvpic

family as the only member of his new higher taxon

"Prionoglossa," giving it equal rank with other groups

such as Taenioglossa and Ptenoglossa. Jeffreys (1859a)

thought these animals the only surviving members of

the otherwise extinct genus Euomphalus Sowerby,

1814. Fretter (194S) showed in a detailed anatomical

study that Omalogt/ra atomii.s (Philippi, 1841) differs

greatly from the "prosobranchs" with which it was tra-

ditionally placed. Omalogyrids have regained interest

in recent years because of their presumed basal po-

sition within the heterobranchs (Haszprunar, 1988;

Ponder, 1990, 1991). Their exact relationships remain

uncertain; recent suggestions (Bandel, 1996; Pacaud

& Le Renard, 1996) to combine the Omalogyridae

with several other families in a superfamily Architec-

tonicoidea are not supported by anatomical data (Hea-

ly, 1993; Huber, 1993).

At the nomenclatural level, certain confusion exists

in the literature about the usage of genus-group

names such as Ammonicera versus Ammonicerina, and

about the identity- and authorship of Omalogi/ra's type

species.

The discovery of an Ammonicera species in the

Florida Keys, representing the smallest gastropod

known in the United States, is here used to summarize

existing data on Omalogyridae in the Atlantic Ocean

and to address additional taxonomic problems.

Page

THE NAUTILUS, Vol. Ill, No, 1

ABBREVIATIONS USED

AMNH American Mu.seum of Natural History, New

York, U.S.A.

BMNH The Natural Hi.ston Museum, Ujiidon, Unit-

ed Kingdom

FMNH Field Museum of Natural Historv, Chicago,

U.S.A.

MLP Museo de La Plata, Argentina

MNCN Museo Nacional de Ciencias Naturales, Ma-

drid, Spain

MNHN Museo Nacional de Historia Natural, Santiago,

Chile

ZMB Zoologisches Museum, Humboldt Universitat,

Berhn, Germany

SEM Scanning Electron Micrograph

RESULTS

FamiK Omalogyridae G. O. Sars, 1S7S: 215 [as Hom-

alog>Tidae]

(often erroneously credited to Fischer, 1885; e.g., Ab-

bott, 1974)

Genus Ammonicera Vayssiere, 1893

Ammonicera ininoiioUs Rolan, 1992

(Figures 1-8)

Omalogijra species. — V'okes and Vokes, 1984: 168, ligs. 7, 7a

(SEM).

Ammonicera minortalis Rolan, 1992: 40, 42, figs. 10, 11 (teleo-

conch), 1.3, 15 (protoconch) (all SEM).

Holotype: (MNCN 15.05/6794): shell diameter 0.35

nun: t\pe locahty: north of Cuba, Baracoa; holotype

from 4 m depth.

Type material studied: 3 paratypes, AMNH 226450,

from Kpe localit\.

Florida material studied: 2 Florida Keys specimens

collected and observed ahye, one each from station FK-

045 [Indian Key Fill, Mile Marker 79, Monroe County,

24°53'25"N. 80°40'28"W, Gulf side, rocks in 0.5-1 m

among Thalassia/Halodulc seagrasses, 20 September

1996[ and station FK-062 [Missouri Key, Mile Marker

39.5, Monroe County, 24°40'29"N, 81°14'21"W, Gulf

side of Missouri-Ohio Key bridge, snbtidal rocks, 14

April 1997]. Also empt\- shells from sta. FK-040 [Mis-

souri Key site as above, 12 March 1996], FMNH 279010

(1 shell); FK-057 [Missouri Key site as above, 26 Sep-

tember 1996], FMNH 279011 (10 .shells incl. SEM ma-

terial), AMNH 288137 (5 shells). All localities were fully

marine and were sampled by the authors by "rock wash-

ing" (bnishing and rinsing of rock surfaces).

Distribution: Now known from north and south

coasts of Cuba (Rolan, 1992), the Florida Keys (this pa-

per), and the Yucatan Peninsula (Vokes & Vokes, 1984).

Description: Shell (Figs. 1-2) extremely minute, di-

ameter 0..34-0.46 mm (0.2-0.4 mm, fide Rolan. 1992),

planispiral, tightly coiled, glossy, uiiii'orinly dark brown,

resembling a miniature annnonite in shape and sculp-

ture. Protoconch (Figs. 3^) of 1.3 whorls' (identical in

SEM but described as ".3/4 whorl," by Rolan, 1992), di-

ameter 120-135 |jLm, distinctively sculptured v\'ith one

major spiral cord at mid-whorl, reticulate sculpture pe-

ripheral to major cord, and .3—4 smaller spiral ridges cen-

tral to major cord. No distinction of a separate larval

shell ("protoconch 11"), indicating the absence of a free

swimming larval stage. Coihng near-planispiral, with

slight initial hyper.strophy (compare Figs. 3 and 4). Te-

leoconch of about 1.3 rounded whorls (1-1.5 whorls fide

Rolan, 1992), sculptured with prominent elongated a.xial

tubercles, regularly spaced, equally sized, beginning im-

mechately after protoconch, numbering 15-19 (1.3-17

fide Rolan, 1992) on body whorl, fading to no axial sculp-

ture at the periphei"v. Tubercles and spaces between al.so

with fine growth Unes. Periphery (Fig. 6) uniformly

rounded, smooth except for fine growth lines. No dis-

tinct spiral sculpture (but occasionally with extremely

fine hues, see specimen in Fig. 1). Outer lip (Figs. 1-2)

thin, sharp, ending in a single plane perpendicular to the

plane of coiUng; aperture circular; columella without

folds or grooves. Head-foot (Figs. 7-8) translucent to

nearly transparent. Animal ghding rapidh- on short foot,

with blunt, very active propochum. Shell held nearly ver-

tically as the snail crawls. Transparent operculum on

hindfoot serving as a support for the coil of the shell.

Head with two finger-shaped tentacles, each held in an

erect arch curxang toward the midline; eyes black, near

base of tentacles. Radula and internal anatomy not stud-

ied.

Habitat: Anim;ds not observed in situ, but collected

from shallow snbtidal rocks covered on one or several

surfaces with various polvchaete wonn tubes, marine ;d-

gae, sponges, tunicates, and numerous other attached or

free-living mollusks. Diet unknowm (but see below).

Remarks: Although the diet ot Ammonicera minor-

talis has not been confirmed, it is hkely to feed on the

variety of algal species growing in its subddal rock hab-

itat, based on literature records on the habitat/tbet of

other Omalogyridae: on Codiuin and Zxstera (Omalo-

gijra atomiis [as Eiiomphahis nitidissimiis] — Jeffreys,

1859a, 1859d); on Ulva (O. atomus—Fretter, 1948); on

Fiictis (A. rota — Nordsieck. 1972); on Ulva and Enter-

omoiyJia (O. atomtis — Fretter & Graham, 1978; Gra-

ham, 1988); on Fucits, Laminaria, Cladophora. Coralli-

na. Ulva {Ammonicera rota — Fretter & Gr;iliam, 1978);

on Padina ( Hawaiian A. japonica — Kay, 1979); piercing

algal cells and sucking out the contents, and depositing

egg stranils on bases of Cladophora (A. rota /A. fischer-

iana — Franc, 1948; Graham, 1988); on Halimeda {A. ja-

ponica and others — Sleurs, 1985a, 1985c); on Zostera,

Ulva, Cifstoseira, and epiphytic diatoms (O. atomiis, A.

fisc1wriana—C,:vj}m\. 1993). Bullock tf «/., 1990, pnnid-

ed the most det;iiled ;iccount, reporting Azorean O. ato-

iints and ,\ fischeriana from a varieh' of algae, incluihng

' Ascertained using the method of Taylor as summarized by

[alilonski & Liitz (1980: ;5;}0, fig. 4)

R. Bieler and P. M. Mikkelseii, 199.S

Page 3

Figures 1-6. Amnionicera minortalis. shells by scanning electron nncroscopy (tonr different shells, F'MNH 279011). 1. Apical view.

2. Umbilical \ie\v. 3. Protoconch (detail of fig. 1). 4. Protoconch (detail of fig. 2). 5. Apertural \iew. 6. Dorsal view (from "above,"

as seen in crawling animal). Scale bars: Fig. 1 = 100 (j.m (Figs. 2, 5, 6 at same scale); Fig. 3 = 20 (j.m (Fig. 4 at same scale).

Page 4

THE NAUTILUS, Vol. Ill, No. 1

8

Figures 7-8. Ammonicera 77iinortalis. sketch of liNing animal.

Maximum shell diameter = 0.42 mm, 7. Right lateral view. 8.

Dorsal view.

Enteromorpha, Cijstoscira, Viva, Ptcrocladia, Petjson-

nelia, Halopteris, Asparagppsis, as well as Codium. Om-

alogijra atomus, which they also found on Gclidium and

Sarga.ssum, was the dominant species on Chondria and

the only moUusk found on Funis in that study.

DISCUSSION

Species-level identification: Roliin (1992) described

this species based on empty shells collected from north

and south coasts of Cuba (3-20 m). A comparison with

the excellent original illustrations and with paratypic ma-

terial at AMNH proved the identity of the Florida Keys

specimens. No other known Atlantic species combines

such axial teleoconch sculpture with reticulated sculp-

ture of its protoconch. Rolan apparently was unaware of

an earlier record of this iorm, as "Ottialogyra species,"

by Yokes and Yokes (1984) who collected it in Arrecife

Alacran, about 140 km north of Progreso, Yucatan, in

the Gulf of Mexico.

According to Rolan (1992: 42), only Ammonicera ja-

ponka Habe, 1972, described as "Japan's smallest gas-

tropod" from Honshu, is "superficially similar but it has

very constant spiral striae." Habe's species (1972:11.'>-

116, figs. 1^) was described as 0.42-0.68 mm in di-

ameter, dark brown in color, with 'about Ifi annulations

in the body whorl" (Habe, 1972:116). Habe did not

mention or illustrate the spiral sculpture noted by Rolan.

Additional specimens were described and illustrated as

Ammonicera japonica from Hawaii by Kay (1979:92, figs.

32A-C [SEM], as Omalogijra; earlier reported bv Kay &

Switzer, 1974:278, table 1, from Fanning Island). Kay

mentioned sculpture "from 16 to 18 axial ribs on the last

whorl, the ribs becoming obsolete at the periphen,. " Spi-

ral striae were not described but faint spiral sculpture is

visible in one illustrated shell (Kay, 1979:fig. 32B).

Sleurs' (1985a:4-5, pi. 1, figs. 1, 6, 9 [SEM]; as Omal-

ogtjra) description of A. japonica from Papua New Guin-

ea was very similar. He described the protoconch in de-

tail "with reticulated sculpture at the abapical side" (his

fig. 9); spiral sculpture of the 0.3 to 0.45 mm large te-

leoconch was not discussed, but shows very faintly in one

SEM illustration (his fig. 6). Fuk-uda's illustration of this

species from the Ogasawara (Bonin) Islands (1994:pl. 35,

fig. 697a-c; as "Omalogijla" japonica), shows no spiral

sculpture. The large specimen illustrated (0.4 mm) has

about 19 axial ribs.

The protoconch and teleoconch sculpture of the shells

oi Ammonicera minortaJis and A. japonica are extremely

similar according to the SEMs provided bv Rolan (1992),

Kay (1979), Fukuda (1994) and Sleurs (i985a), respec-

tively, and suggest synonymy of the Caribbean and Indo-

Pacific species. Faint spiral teleoconch sculpture appears

to occur in some indi\'iduals of both nominal species.

However, the di.sjunct distributional pattern makes fur-

ther study necessary. No siniOar fomi has been de-

scribed or recorded from the eastern Pacific (Shasky,

1989).

As also noted by Rolan (1992), Ammonicera iniuortalis

is similar to the European A. rota (Forbes & Hanley,

1850) in its teleoconch characters (but the latter has a

greater number of whorls and axial tubercles continuing

over the periphery). Ammonicera rota has, however, a

very different protoconch without reticulated sculpture

(see, e.g., Rodriguez Babio & Thiriot-Quie\Teux, 1974:

pi. 2, F-H; as A. fischeriana). Also similar is A. plicata

Sleurs, 1985, from the Maldives (19S5b:20 ff , figs. 2, 7,

10, 13, 14), which has a larger teleoconch (0.45^ to 0.65

mm) with weaker axial ribs and a protoconch lacking the

reticulate sculpture present in A. minor-falis and A. ja-

ponica.

Genus-level identification: Omalogyridae currently

comprises three recognized extant genera: Ammonicera

Yayssiere, 1893, Omalogtjra Jeffreys, 1859, and Retro-

tortina Chaster, 1896. The last (with type species by

monot\pv: R. ftiscata Chaster, 1896) has a sinistral te-

leoconch that distinguishes it from Ammonicera and

Omalogtpa.

Bandel (1988:9) also placed Orbitestella Iredale. 1917,

in this family, but Ponder (1990) showed that this genus

belongs in the Yalvatoidea, not OmiJogv roidea. Bandel

(1988), who viewed om;ilog\Tids as small-bodied mem-

bers of Architectonicidae or Architectonicoidea (pp. 10,

17), attempted to introduce a new fossil genus "Neam-

phitomaria." but did not designate a type species. Ban-

del (in Dockeiy, 1993:92) subsequently provided such a

R. Bieler and P. M. Mikkelsen, 1998

Pas^e 5

designation (the Upper Cretaceous Psctidomalaxis stan-

toni Sohl, 1960) and thus \ahdated Ncainphitomaria of

that date [not as of 1988 as is frequently cited; see ICZN

Art. 13(b)]. Amphitomaria Koken, 1897, and Neamphi-

tomaria Bandel, 199.3, were then placed in a new family,

Amphitoniariidae, by Bandel (1996), thus removing the

genus agiiin from the Omalogyridae.

The placement of the present species in the genus

Ammonicera, rather than Omalogijra, is here accepted

because of (1) its protoconch sculpture with strong spi-

ral ribs and grooves (in contrast to small tubercles in

Omalogijra; e.g., Rolan, 1992); (2) the presence of dis-

tinct cephiilic tentacles (absent in Omalogijra); and (3)

its strong teleoconch sculpture (absent or weak in Om-

alogyra). The kniovvn radulae of Omalogijra and Am-

monicera (not yet studied for A. mhwrtalis) are so dif-

ferent between members of the two nominal genera that

Sleurs (1985c: 181) suggested that they might belong to

different families. However, pubhshed radular data dif-

fer even within the two genera: Omalogijra radulae have

been described as either uniserial (Jeffreys, 1859, 1867;

Thiele, 1929; Sleurs, 1985c) or with a formula of 1-1-1

(G. O. Sars, 1878; Thiele, 1929; Egorova, 1991). Those

of Ammonicera have been described with a fomiula of

1-1-1 (Vayssiere, 1893) or 1-1-0-1-1 (Sleurs, 1985b, c),

and so definitive conclusions must await a detailed com-

parative study.

Unfortunatelv, the taxonomic history o{ Ammonicera.

Omalogijra. and their included species is exceedingly

complex and confused (see discussion below).

REMARKS ON AMMOTSllCERA AND ITS TYPE

SPECIES

Ammonicera was introduced by Vayssiere (1893:16 ff.)

for Homalogi/ra fischeriana Monterosato, 1869. He pro-

vided a full anatomical description based on histology

and studies of the radula. Franc (1948:142 ff ) and Sleurs

(1985a:9) questioned the identity of Vavssiere's material,

assuming that his work was based on misidentified "Om-

alogijra rota" Forbes and Hanlev, 1850. Compared to

Gaglini's descriptions and illustrations (1993:933-04,

934-03-04), Vayssiere s line drawings of the shell (1893:

figs. 8-9) seem to represent typical A. fi.scheriana in col-

or pattern and relatively fine crenulations of the periph-

ery, although the sketched pronounced axial ribbing is

more representative of the nominal species A. rota.

Monterosato himself considered the two nominal species

as varieties of one (e.g., Monterosato 1872:38; 1875:29),

and manv recent authors (e.g., Fretter & Graham, 1978;

Backeljau ct ah, 1984; Knudsen, 1995) have deemed

them svnonymous (see also Hoisaeter, 1968; van Aartsen

et ai. i984). Gaglini (1993), on the other hand, argued

convdncinglv for the presence of two sympatric species.

Whether or not thev will prove to be sviionvanous, they

are without doubt so closely related and mor{)hologic;illy

similar that it will not impact interpretation of the nom-

inal genus Ammonicera (in contrast to Sleurs, 1985a).

In addition to Vayssiere's extensive description, pub-

lished biological information about this/these species in-

cludes description of gross anatomy (Franc, 1948), ner-

vous system (Hulier, 1993), egg capsules (Franc, 1948

[summarized by Knudsen, 1995]; Graham, 1988), and

feeding (Gniham, 1988).

Ammonicera should not be confused with Ammoni-

cerina — see svnonvmy of Omalogijra (below).

REMARKS ON OMALOGYRA AND ITS TYPE

SPECIES

Omalogijra was introduced by Jeffreys (1859b) in the

midst of an engaged discussion (with Clark, 1859) ulti-

mately involving the identities of Helix niticlissima Ad-

ams, 1800, "Skenea" nitidi.ssima .sensii Forbes and Han-

ley, 1850, and "Tntncatella" atomu.s PhiUppi, 1841. The

current understanding of Omalogijra is based on Fret-

ter's (1948) excellent anatomic;il study on British animals

identified as O. atonuts. Other pubhshed information on

this species includes gross anatomy and radula (G. O.

Sars, 1878), nervous system (Huber, 1993), spermatozoa

(Healy, 1993), and egg capsules (Graham, 1988; Knud-

sen, 1995 [However, it should be noted that the accom-

panying SEM shell photographs, Knudsens fig. 5, seem

to be of a skeneopsid, not of O. atomit.i]). The "eggs" of

0. atomtis as described by Jeffreys (1867) and Lebour

(1937) were subsequently recognized as misidentified

glandular stiiictures (Fretter, 1948).

No type material for any of these nominal taxa could

be located; our following discussion thus has to concen-

trate on hterature review: Helix nitidissima ]. Adams,

1800, was introduced with a short description and three

illustrations (here reproduced in Fig. 9). The species was

accepted and cited, in various generic combinations, by

subsequent authors {e.g., Weinkauff, 1868:266, as "Spira

nitidissima Adams"). Many authors have considered H.

nitidis.sima J. Adams, 1800, as svnonymous with Tntn-

catella atoinus Philippi, 1841 {e.g., Fischer, 1857; Wein-

kauff, 1868; Fretter & Graham, 1978; Rolan, 1983; Gra-

ham, 1988; Poppe & Goto, 1991; Rosenberg/Malacolog,

1997). The original description by Adams (1800: 4, pi.

1, figs. 22-24) was based on the shell alone: ■H.[elLx]

testa duobus anfractibus, subtihssime transverse striata.

Obs. Corneous, pellucid, umbilicated; easily distin-

guished by the uncommon briUiancy of its glossiness."

Original figure 23, said to be of "natural size" (1800:6)

measures nearlv 3 mm. The shell, much too large to be

a European omalogyrid species, was subsequently rec-

ognized as "evidently the fry of Zonitcs radiatuliis [J.

Alder, 1830]," a land snail, by Jeffreys (1867:71). The

holotype of H. nitidissima was not located (K. Way,

BMNH, pers. comm., 1997). The interpretation as a

young stage of a British land snail is here accepted; H.

nitidi.ssima Adams is not a senior svnonvm of T atomiis.

Much of the interpretation of "nitidissima " bv sub-

sequent authors was based on "Skenea" nitidissima sensu

Forbes and Hanley, 1850, who used diis name for a dif-

ferent species. Several authors erroneously credited

Forbes and Hanley with the description of a new species

Page 6

THE NAUTILUS, Vol. Ill, No. 1

/C-^ /CtJ

tB

Jug^

10

Figure 9. Reproduction of original illustrations of Helix niti-

dissima ]. Adams, 1800 (from Adams, 1800: pi. 1, figs. 22-24).

Figure 10. Reproduction of original illustrations of Tnincatella

atomiis Philippi, 1841 (from Philippi, 1841: pi. 5, figs. 4a-d).

"Skcnca nitidissima " {e.g., Jeffreys, 1860; Nordsieck,

1972; Nordsieck & Garcia-Talavera, 1979; Gaglini,

1993). However, Forbes and Hanley themselves (1850:

158) cited the species as "S.lkcnca] nitidissima, Adams"

with full page and figure reference to Adams' original

work. It is this misidentified "nitidissima scn.sti Forbes

& Hanley " that enters into the various lengthy published

chscussions comparing "nitidissima" and TntncatcHa ato-

mus.

Phihppi (1841:54, pi. 5, figs. 4a-d) described and il-

lustrated Tnincatella atomtis, collected in Sorrento

(Campania, southern It;ilv). He emphasized that he was

able to studv the animal in detail at a magnification of

60 times and placed it in Tnincatella because of the an-

imal's similarity to members of that genus. In 1844, he

re-described the species (p. 134, pi. 24, fig. 5; again as

"n. sp.") and reproduced his 1841 illustrations of T ato-

miis. Philippi's (1841) illustrations, here reproduced in

Fig. 10, show a living specimen with planispiral shell

(with logarithmic growth), tapering tentacles, an oper-

culum, and a representation of actual size of about 0.5

mm. The t\pe material has not been located in Berlin

or Santiago (von Rintelen, ZMB, pers. comm., 1997;

MNHN, pers. obs., 1997).

Forbes and Hanley (18.50:158-160, pi. 73, figs. 7, 8)

described and illustrated a British shell under the name

"S.[hettea] nitidissima, Adams "; tlic\ did not mention the

living animal. They placed Phifippis Tnincatella atomtis.

with questicm mark, in .synonymy. Jeffreys ( 1859a: 109-

111, pi. 3, figs. 15a, b, 16a-c) discussed the species, as

Euomphalus nitidi.ssimtis. with a sketch of the animal

(showing ciliated head lobes, no tentacles, and a unise-

riate radula). He reported its range as 'from the Shet-

lands to Sicily, and probably far bevond these limits" (p.

Ill) based in part on the s)aionymy of Tnincatella ato-

mus of Pfiilippi, and expressed his astonishment over

Philippi's "mistake" of describing the animal so differ-

ently (i.e., with tapering tentacles).

Much of the ensuing confusion was based on (1) the

treatment of Philippi's Italian "Tnincatella atomtis" spec-

imens as members of the British "Omalogt/ra nitidissi-

ma" .sensu Forbes and Hanlev, and (2) the tiiscrepancy

between gross anatomical descriptions of these two spe-

cies, i.e., with or without tapering head tentacles, re-

spectively.

Clark (1859:410^13, text-figure), after reexamining

British animals reconstituted from dried specimens, dis-

agreed with Jeffreys and corroborated the correctness of

Philippi's figure of an animal with triangular tentacles,

the large eves embedded at the center of their bases. "It

appears quite clear that Mr. Jeffreys has delineated his

animal with rounded lobes, or, in other words, v\ith the

tentacles retracted. . ." (p. 411). Jeffreys (1859b:498) re-

butted: "What Mr. Clark supposed to be tentacula must

have been the shrivelled lobes of the veil. . . " Fischer

(18.59:364-367) joined Clark in criticizing Jeffreys

(1859c), assuming that the latter had described a lar\'al

stage with vela instead of tentacles. Jeffreys then (1860:

108-111), in rebuttal of Fischer, affirmed that his ob-

servations were based on adult specimens without ten-

tacles. Finally, in British Concholog)', Jeffreys (1867:67-

71, pi. 1, fig.' 5; 1869:209, pi. 70, fig. 2) again described

the shell and animal in detail, reaffirming his opinion of

Philippis error, but recognized the prioritv of "Homal-

ogt/ra atomu.s" (Philippi) over "Shenea nitidissima" of

Forbes and Hanley. This "anatomically corrected " Hom-

alogi/ra atomtis, with "Skenea" nitidissima .sensu Forbes

and Hanley in sMionvinv, is the Omalogt/ra atomtis de-

scribed in det;iil by Fretter (1948) and that currently

forms our concept of the genus.

Unfortunately the original figures of Tnincatella ato-

mtis Philippi, 1841, are in conflict with the descriptions

of Fretter Philippi's illustrated gross moqihological de-

tails (i.e., tapering tentacles) are indicative of Ammoni-

cera. Meanwhile, the sketched smooth sheU appears in

line with the current concept of Oinalogt/ra. In the ab-

sence of t\pe material, it is impossible to explain this

discrepanc\'. It is possible that Philippi's material con-

tiiined members of both genera and his illustration is a

composite based on more than one species.

In the interest of nomenclatural stabilitv, we base

our interpretation of Philippis Tnincatella atomtis

on hi.s illustration/description of the shell alone (ex-

cluding the anatomy in original fig. 4c), thus pre-

serving this name for ^'Skenea" nitidissima sensu

Forbes and Hanley, 1850 (non Adams, 1800), and

Omalogtjra atomus sensu Jeffreys, 1859, as well as

Fretter. 1948.

The taxonomic confusion has been compounded by

uncertaint\' about the tvpe species designation and the

date of introduction of Omalogtjra. Some authors [e.g..

R. Bieler and P. M. Mikkelsen, 1998

Page 7

Waren, 1980:12) cited it as having been introduced by

Jeffreys (1860), with t\pe species TnincatcUa atoinus

Philippi, 1841, by monot>py. Others (e.g., Wenz, 1939:

647-648) gave "O. niticlissiina (Forbes & Hanley)" as

tvpe species. The date of description is often erroneously

cited as •■1867" (e.g., Abbott, 1974; Castellanos, 1989a;

Vaught, 1989; Rios, 1994).

The generic name "for the reception ol these anom-

alous nioUusks" was in fact proposed by Jeffreys (1859b:

498). In that paper, he referred bv name to "Eitoin-

phalus niti(li.ssiiini.s" (with reference to his earlier, 1859a,

article), to "E. Rota" and its "varietv tricariiuita of Web-

ster." In the referenced article, he additionally stated a

synonym for '£."' nitidi.'isimus: "I have no doubt that it

is the Tnincatclla atoiuti.s of Phihppi" (1859a:lll). Jef-

freys did not indicate a t\pe species. Following ICZN

(1985: Art. 69(i)), there are four "originally included

nominal species":

Helix nitkli.'i.'iimti J. Adams, ISOO. Now considered a land snail

[Jeffreys recognized the misidentification only in 1867; his

(1859a, b) usage thus cannot be construed as "deliberately

used in the meaning of a previous misuse" (ICZN, 1985;

Art. ll(i)].

Tnincatellti atomus Philippi, 1S41 [in sviioimiiy],

Skcnea rota Forbes and Hanlev, 1850. Now considered a mem-

ber of Amnumicern.

Skenea tricarinata Webster, 1856. Described as a potential new

species; subsequently (beginning with Jeffreys in Webster,

1857) considered a yariety/svnonvni of S. rota-

Jeffreys (1867:69 ff ) synonvmized "Skcnca nitidissima"

.icnsii Forbes and Hanley under TnincatcUa atomus, af-

ter recognizing the tnie Helix nitidissima Adams as a

land snail. He iilso svnonvmized Skenea tricarinata Web-

ster under Skenea rota. No tvpe species was designated.

Jeffreys therein changed the generic name to Homalo-

giji'a, an unjustified emendation. The first authors to se-

lect a tvpe species appear to have been Buciiuov et al.

(1884:78) who stated "TyjDe: Homalogiira atomus Philip-

pi sp. (TnincatcUa)."

We therefore offer the following sviionymies:

Otnalogiira Jeffreys, 1859b:498; tvpe species by subsequent

designation of Bucquoy et al. (1884:78), TnincatcUa nto-

;ni(.s' Philippi. 1841.

Ammonicerina O. G. Costa, 1861: 71; tspe species by .sub-

sequent designation of Dall (1927b: 1.34, as "Ammon-

ocerina"). Ammonicerina simplex O. G. Costa, 1861.

Preoccupied by Ammonicerina O. G. Costa, 1856

[Protista], This ta.xon is usually placed in synonymy

of Ammonicera (e.g., Palazzi & Gaglini, 1979); how-

ever, its tvpe species by subsequent designation be-

longs to Omalogtjra.

Homalogyra Jeffreys, 1867:67 dm unjustified emendation).

Note: In the description of their new genus Transo-

malogi/ra, Palazzi and Gaghni (1979:3.3) made Ammon-

icerina simplex O. G. Costa, 1861. the tvpe species by

original designation. This woidd make Tran.somalogijra

an objective synonym oi Ammonicerina and a subjective

svnonvin of Omalogt/ra. However, as pointed out by

Waren (1991:74), the tvpe species was misidentified,

with Palazzi & Ciaghni's illustration actually showing a

shell of Adcuomphalus ammoniformis Seguenza, 1876.

Waren (1991) thus placed Transomalogijra in the syn-

onymy oi Adcuomphalus Seguenza, 1876, as a genus in-

ceriae scdis in the "Archaeogastropoda."

Omalogt/ia atomus (Philippi, 1841)

Tnincatella atonuis Philippi, 1841:.54, pi. 5, fig. 4a-d [excluding

the sketched animal in fig. 4c].

Skenea nitidissima (Adams) sensii Forbes and Hanlev. 18.50, et

anct. [non Helix nitidi.ssimn J. Adams, 1800].

Homalogijra atomus van vitrea Jeffreys, 1867:69.

Homalogijra atomus \a.r. fasciata Monterosato, 1877:418.

Notes on other named "varieties":

Homalogipa atonnis var maculata Dautzenberg and Du-

rouchoiLX, 1914:27. The authorship of this name is usu-

ally crethted to Monterosato, 1875 (e.g., Gaghni, 1993:

928-02). However, Monterosato's applications and some

subsequent citations of the name are not available for

nomenclatural purposes because they represent nomina

nuda (Monterosat(j, 1875:29; 1878:88; Buctjuoy et al.,

1884:324). The first available introduction appears to be

that of Dautzenberg and Durouchoux (1914).

Ammonicerina atomus "var. pallida Monterosato 1884"

as cited bv Gaghni (1993:928-02) is hkewise not avail-

able as of that date. Monterosato's usage ( 1884:22) of

"var. pallida" is a nomen nudum, as is his Homalogi/ra

atomus var. zonata Monterosato (1878:88), subsequently

cited as "var, ex colore 2, zonata Monts." by Bucquoy et

al. (1884:324; hkewise a nomen nudum).

Honudogt/ra atonuis var nautilijonnis De Gregorio,

1889, was recognized bv Monterosato (1890:141) as a

juvenile of Capuhis ungaricus (Linnaeus, 1758). Nev-

ertheless, the name nautilifonnis De Gregorio, 1889,

was retained bv some authors to describe an Omalogi/ra

morph with a much widened body whorl (e.g., Nord-

sieck, 1972:148; Gaghni & Curini Galletti, 1978:210, fig.

2c), GagUni (1993:928-02-3) introduced a new infra-

subspecific name for this moqih, Omalogijra atomus var.

"inflata."

Homalogi/ra atomus var poh/zona "Brusina mss. (fide

Monterosato)" in Bucquoy et al., 1884:324, pi. 37, fig.

32. Earher references to a varietv' "poh/zona Brusina" by

Monterosato (1872; 1875; 1878) are unavailable because

they were stated in sviiommv or as nomina nuda. Gag-

hni'(1993:931-01, 9.31-02-3) showed that this is a po-

tential sviionvm of O. simplex, not O atomus.

CURRENT COMPOSITION OF OMALOGYRIDAE

Recognized western Atlantic Species: [regions of

type locahties in brackets]

Ammonicera alhospeciosa Rolan, 1992:44, figs. 17, 19, 21

[Cuba]

Page 8

THE NAUTILUS, Vol. Ill, No. 1

Amnwnicera rirrnmrirm Rolan, 1992:45, figs. 23. 26, 28

[Cuba]

Amriumicera familiaris Roldn, 1992:42, 44. figs. 16. 18, 20

[Cuba]

Aintnonicera lineofusciita Rolan. 1992:44-45, figs. 22, 24—25,

27 [Cuba]

Ammonicera minorfalis Rolan. 1992:40. 42. figs. 10-11. 1.3, 15

[Cuba]

Amnu>nicera sculpturata Rolan, 1992:40, figs. 9, 12, 14 [Cuba]

Omalogijra atomiis (Philippi. 1841:54. pi 5, figs. 4a-d) [Med-

iterranean]

Omalogtjra burdwoocliann (Strebel, 190S:.52. pi. 6. fig. 85a-c

[Burdvvood Bank, south ot Falkland Islands]

Omaloayra fiiscopardahs Rolan, 1992:.36. .38, figs, 1, 3, 5, 7

[Cuba]

Onmlogtjra taludana Castellanos, 1989a:88-89, figs. 1, 2 (plus

sketch of apertural aspect in 19S9b:pl. 1, fig. 10) [off San

Jorge Gulf Argentina]

Omaloaijra zebrina B-olin. 1992:38. figs. 2. 4. 6. 8 [Cuba]

For the western Atlantic, eleven omalogyrid species

are currently recognized. Of these, eight are to date only

known from Cuba (all described by Rolan, 1992). Two

others, Omaloaijra bunlwoodiana (Streliel, 1908) and O.

taludana Castellanos, 19S9, are faiown from subantarctic

waters off South America. Two omalogyrid species are

now recognized from the east coast of the United States;

Ammonicera minorialis and O. afomiis.

Several other nominal omalogyrid species have been

reported for the western Atlantic Ocean, but need to be

excluded from that fauna:

"Omalogyra piano rbis ■. A nominal species in the west-

em Atlantic frequently cited as an om;dogyricl is Lipp-

istcs? planorbis Dall, 1927a: 131, originally described

from "off Femandina," Florida. This deep-water species

was re-described in detail by Moore (1971: 114-116, fig.

1) as Omalogifra planorbis, and subsequently called Om-

alogijra (Ammonicera) planorbis {e.g., Abbott, 1974:81;

Rios, 1994:60). This taxon was placed in Palazzia Waren,

1991, as an "archaeogastropod" group of uncertain affil-

iations, tentatively assigned to Skeneidae (Waren, 1991:

74, 76).

"Ammonicera jischctiana": Nortlsieck (1972:149) re-

ferred to "Ammonicera fischeiiana (Monterosato. 1869)

= denscco.'itata [sic] (Jeffreys, 1884)" in "Westindien,"

without further explanation. This synonymy is erroneous.

The West Indian record for this Mediterranean species

is based on Watsons (1886) "Challenger" materiiil of

"densicostata" as explained in the following.

"Omalogifra (Ammonicera) densicostata": Homalogt/ra

densicostata Jeffreys, 1884:129, pi. 10, fig. 1. was de-

scribed from "Porcupine" stations (1098-2002 m) off the

coast of Portugal. Additional material from a "Bulldog"

cruise off Labrador (2967 m) was also included in the

original description. Abbott (1974:81) reported this spe-

cies as Omalogyra (Ammonicera) densicostata Iroui deep

water off Portugal, the Azores, and Labrador Moore

(1971:114) showed that the Labrador ("Bulldog") ma-

terial in fact belongs to "Lipj)i.ste.s" planorbis Dall, 1927,

thus removing the Labrador record for "O." densico.s-

tata. Watson (1886:677) added a "Homalogyra densicos-

tata (?)" record from north of the island of Culebra,

between Puerto Rico and the Virgin Islands ("Challeng-

er" staHon 24, 71.5 m). Moore (1971:11.5-116) doubted

both the syiionviiiv of the Challenger material and that

of the shallow-water material reported by Dautzenlierg

(1889:46) for the Azores, thus restricting densicostata

again to the eastern Atlantic. The species was considered

a member of the eastern Atlantic omalog\Tid fauna until

recently (e.g., Sabelli et al., 1990; Gaglini. 199.3). Hom-

alogifra densicostata was syiionymized imtler Adeiiom-

phaliis ammonifonnis Seguenza, 1876, and placed as an

"archaeogastropod" of uncertain affiliations, tentatively

assigned to Skeneidae (Waren, 1991:74 ff.).

Recognized eastern AtlanticAlediterranean Spe-

cies: [regions of type localities in brackets]

Ammonicera biimayi Rolan. 1991:112, figs. 1.3-14 [Cape Verde

Archipelago]

Ammonicera ftscheriana (Monterosato, 1869:274—275. pi. 13,

fig. 1) [Mediterranean]

Ammonicern tignrn (Palazzi. 1988:105, figs. 8. 18) [Madeira]

Ammonicera midlistriata Rolan, 1991:112, 114, figs. 1.5-16

[Cape Verde Archipelago]

Ammonicera nolai Rolan. 1991:110. figs. 8-9 [Cape Verde Ar-

chipelago]

Ammonicera oteroi Rolan, 1991:110. 112, figs. 10-12 [Cape

Verde Archipelago]

Ammonicern robusta Rolan, 1991:114-115. figs. 17-18 [Cape

Verde Archipelago]

Ammonicera rota (Forbes & Hanley, 18.50:160, pi. 73, fig. 10;

pi. 88, figs. 1, 2) [Ireland]

Ammonicera rotundata (Palazzi, 1988:105, figs. 10, 21, 27)

[Madeira]

Annnonicera verdensis Rolan, 1991:109, figs. 6-7 [Cape \'erde

Archipelago]

Omalogt/ra afomus (Philippi, 1S41:.54, pi. 5. figs. 4a-d) [Med-

iterranean]

Onuilogi/ra di.scutus Palazzi, 1988:104, figs. 1, 20 [Madeira]

Omalogtpa simplex (O.G. Costa. 1861:72, pi. 11 figs. 3 a, b)

[Mediterranean]

Omaloayra undosa Palazzi, 1988:104, figs. 5, 15 [Madeira]

Retrototiina fuscata Chaster. 1896:2 [Strait of Gibraltar]

In the eastern Atlantic, fifteen omalogyrid species are

currently recognized, comprising ten species of Ammon-

icera, four of Omalogifra. as well as Retrototiina fuscata

(for Mediterranean records see also Sabelli et al., 1990;

Le Renard et a/./CLEMAM, 1997). Omalogifra atcunus

is the only species known from both sides of the Atlantic;

it is widely distributed, ranging from the Mediterranean,

Madeira, and the Azores to Norway, Iceland. Greenland,

and in New England (Abbott, 1974; Bullock, 1969, 1995;

Fretter & Gniliam, 1978; Thorson, 1944) from Maine to

Rhode Island. Egorova (1991) recognized material from

Antarctic waters, previously identified and cited as O.

atomits, as members of a moiphologicallv extremely sim-

ilar species, O. antarctica Egorova, 1991.

Several other nominal omalogyrid species ha\e been

described for the eastern Atlantic. Of the.se, Omalogyra

apeiia Sykes, 1925:192, Ironi off Portugal, was recog-

nized as a member of the "archaeogastropod" genus Eii-

R. Bieler and P. M. Mikkelsen. 1998

Paee 9

daronia Cotton, 1945, Iiy Waren (1991:80). Homalogijra

granulosa Svke.s, 1925, also from off Portugal, was

placed in the ■"archaeogastropod" genus Rctigyra Waren,

1989 (see Waren, 1992:168). Homalogijra paiadoxa

"Monterosato (? MS.)" of Svkes (1925:192) is a nomcii

niuliim. Two other noiinn;il species introduced bv Sykes

{H. sititiosa Sykes, 1925, and H. (?) marshalli Sykes,

1925) are in need of further study (see Palazzi, 1992).

An additional Mediterranean species, O. ausonia P;ilazzi,

1988, was recently made the t\pe of Palazzia Waren,

1991, and transferred to the "archaeogastropods," with

tentative placement in the Skeneidae (Waren, 1991).

Nominal species Homalogi/ra oniata Dautzenberg, 1889

(p. 46, pi. 4, fig. 9a-d), described from the Azores, is still

in need of rein\'estigation.

It should be noted that Palazzi (1988) used "ausonia"

(Italy) and "disculus" (little chsk) as nouns in apposition

in the original descriptions; recent usage as "Palazzia

ausoniac" or "Oiiwlogi/ra disciila" (e.g.. Sabelli et al.,

1990; Giannuzzi-Saveili ct al. 1994; Arduini ct al.. 1995)

are incorrect subsecjuent spellings.

ACKNOWLEDGMENTS

This project, as part of a broader study of lower heter-

obranch gastropods, was supported under National Sci-

ence Foimdation grant DEB-9318231 to RB. Field and

laboratory' work in Florida was made possible through

Visiting Scientists Awards by the Smithsonian Marine

Station at Unk Port (SMSLP) to RB; Dr Mary E. Rice

and the station staff are gratefulK acknowledged for

their support. Field collecting in the Florida Keys was

supported through supplementary funding from the

Bertha LeBus Charitable Trust and Field Museum's

Marshall Field Fund. Dr. Kenneth J. Boss (Museum of

Comparative Zoologw Harvard Uni\'ersit\) and Richard

E. Petit (North Myrtle Beach, South CaroUna) kindly

provided hterature, and Roberto Cipriani (FMNH) as-

sisted with translations. We thank Dr. Sergio Letelier

(MNHN) for hospitality- extended during RB's \isit to

Santiago, Kathie Way (BMNH) and Thomas von Rin-

telen (ZMB) for information on tvpe holdings of their

respective collections. Dr. Cristian F. Ituarte (MLP) for

a specimen loan, and Richard E. Petit (North Myrtle

Beach) as well as tvvo anonymous reviewers for their

comments on the manuscript. The excellent facilities

and helpful staff of the AMNH and FMNH libraries are

also ackniowledged. This is SMSLP Contribution no.

434.

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THE NAUTILUS 111(1):13-21, 199S

Page 13

Moiphology and Systematic Position of Rissoella carihaea

Rehder, 1943 (Gastropoda: Heterobranchia: Rissoellidae)

John B. Wise

Departnieiit of Malacolog)-

Houston Museum of Natural Science

One Hermann Circle Drive

Houston. Texas 77030-1799 USA

E-mail; j\[email protected]

ABSTRACT

Rissoella caribaea is one of onlv three known western Atlantic

species of rissoellids. It was first described by Rehder ( 1943a,b)

hving amongst algae on the mangrove prop-roots at Bonefish

Key, Lower Florida Keys, Florida. Rissoella caribaea, like other

rissoellids, has a pair of cephalic tentacles and a verv distinctive

(diagnostic) pair of oral lobes on either side of its bifid snout.

It has a rhomboid-shaped jaw, a s\inmetrical radula. each row

composed of rachidian and one pair each of lateral and mar-

ginal teeth. Rissoella caribaea is a simultaneous hermaphrodite,

with well de\eloped penis, \'as deferens and a large portion of

the female reproductive tract within the mantle floor. This

study supports the allocation of this species to the genus Ris-

soella. However, this assignment is based upon our current and

very- fimited understanding of rissoellid morphologv and phy-

logenetics.

Keij words: Heterobranchia, Rissoellidae, Rissoella. anatoniN'.

INTRODUCTION

Rissoella caribaea Rehder, 1943a, one of only three

knovvTi western Atlantic rissoeUid species, occurs from

the Florida Keys to Brazil and Puerto Rico (Robertson,

1961a; Abbott,' 1974; Rios, 1994). Although the genus

has a worldwide distribution, and the family is consid-

ered important for developing an understanding of gas-

tropod phvlogenedc relationships, the anatomies and life

histories of these minute gastropods are poorly under-

stood (Bartsch, 1920; Thiele, 1912, 1925, 1929; Ankel,

1936; Lebour, 1936; Fretter 1948; Robertson, 1961a,

1985; Fretter & Graham, 1962, 1994; Haszprunar, 1988;

Ponder, 1966, 1983; Huber, 1987, 1993; Ponder 6f Wors-

fold, 1994; Simone, 1995).

Members of the Rissoellidae are thought to be basal

Heterobranchs because they possess the following pu-

tative synapomorphies : 1) ciliary tracts located on the

right side of the mantle ca\atv'; 2) gonoduct on mantle

floor; 3) eve position, posterior to the base of the ce-

phalic tentacles, although Ponder and Yoo, 1977, re-

ported its position to be variable in the family; and 4)

sperm morphology (Haszpnmar, 1988; Healy, 1993;

Waren & Ponder, 1994). Moreover, molecular studies

support the hypothesis that the Rissoelloidea are sister

taxa to the Pyramidelloidea and other "lower" Hetero-

branchs (Harasewvch ct al.. 1997).

This study was undertaken to proxdde further insight

into rissoellid anatomy and biology, and more specifical-

ly, the assignment of R. caribaea to the genus Rissoella.

MATERIAL AND METHODS

Specimens of Rissoella caribaea were collected by wash-

ing the tops and often embedded undersides of rocks

and coral rubble. Debris was placed in a 0.5 mm sieve,

rinsed with seawater to eUminate mud and sand, and

sorted under a dissecting microscope. Snails were kept

alive in small bowls of aerated seawater

Living snails were observed and photographed with a

Pentax 35mm camera mounted on a Zeiss Tessavar dis-

secting microscope. Snails were dissected whole, once

their shells were cracked with forceps and shell frag-

ments removed. Whole animals and/or their alimentary

and reproduction systems (once excised) were stained

with toluidine blue. Line drawings were produced using

a dissecting microscope with a camera lucida.

Shells were decalcified using a commercial decalcifier

(Decalcifving solution. Krajian, J.T Baker). Specimens

were rinsed in 3 changes of seawater and fixed in 10%

formalin buffered with filtered seawater. Snails were em-

bedded in paraffin, sectioned at 3-.5 |xm, and stained

with hematox-yhn and eosin-Y (Sheehan & Hrapchak,

1980).

Opercula and shells were cleaned by sonication, air

dried, coated with gold-palladium and examined with a

Cambridge S-650 scanning electron microscope set to

5-10 KEV.

Several specimens were fixed in 2.0% glutaraldehvde

buffered in 0.025M sodium cacodylate in filtered sea-

water Once fixed, tissues were rinsed with sodium cac-

odylate buffered/seawater These specimens were de-

Page 14

THE NAUTILUS, Vol. Ill, No. 1

J. B. Wise, 1998

Page 15

hydrateil in a graded series of ethanol, critical point

dried, coated and examined v\ith a scanning electron mi-

croscope, as ahove.

Specimens used in this study were collected in the

Florida Kevs, Florida Bay at Craig Key, Florida

(24°50'N, 46°45'W), December 1994; and Fiesta Key,

Florida (24°49'N, 8()°47'\V), October 199fi and Febru-

ary- 1997. Identification was confirmed by comparing this

material against the holotype of Rissoella caribaea

(USNM No. 536046). Voucher specimens on deposit at

Houston Museum of Natural Science, Houston, Texas.

(HMNS Nos. 43688-43690).

SYSTEM ATICS

Class Heterobranchia

Family Rissoellidae Gray, 1850

Genus Rissoella Gray, 1847

Type species: Rissoella diaphana Alder, 1848

Diagnosis: Shell transparent, smooth, broadly ovate to

more slender, 1-2 mm in length. Umbilicus broad to

narrowK' keeled or absent. Aperture ovate to semicir-

cular, with thin outer lip. Protoconch of 1-1.5 dextral

whorls. Operculum translucent, corneous, ovate, with

ventral peg on columellar side. Head-foot white, yellow,

gray or black. Head with pair of cephaHc tentacles and

pair of oral lobes (= triangular lobes, Fretter, 1948: oral

tentacles. Ponder & Yoo, 1977) on either side of bifid

snout. Eyes black, often surrounded by clear mound.

Nervous svstem highly concentrated, with fused gangfia

and untvNasted visceral loop. Mentum absent. Propodium

bifurcate. Foot broad anteriorly, tapered posteriorly. Jaw

present. Radula \ariable, with 3, 5, or 7 teeth/row. Ra-

chidian cusps numerous, few or absent. Feeds on dia-

toms, micro-algae, and detritus. Simultaneous hermaph-

rodite, with opafjue or orange ovotestis iilong columellar

side of visceral mass. Egg capsules hemispherical to

ovoid, with 1-2, and rarely 3 eggs. Development direct.

Remarks: For a discussion of the genus and type spe-

cies see Robertson, 1961b, 1962, and Ponder, 1966, and

Ponder & Yoo, 1977.

Rissoella caribaea Rehder, 1943a

Rissoella caribaea Rehder, 1943a: 194, pi. 20, fig. 7.

(Holot\pe:USNM 536046: t\pe localitv: Bonefish Key,

Florida Keys); Abbott, 1974:723, ;ifter Robertson, 1961a.

Description: Shell (Figs. 1^): Globular, transpar-

ent, thin, fragile, 1 to 1.8 mm in length, composed of 2

to 3 adult whorls (Figs. 1, 2). Whorls convex, with mod-

eratelv shallow sutures. Body whorl 75 to 80% of shell

length. Teleoconch smooth, with \'erv fine axial growth

lines. Umbibcus narrow, with sharp keel. Aperture elon-

gate-ovate (Fig. 3). Protoconch smooth, dextral, with

1.25 whorls (Fig. 4). Operculum ovate, translucent, with

peg extending from ridge parallel to columellar oper-

cular edge (Figs. 5, 6).

Head-foot (Figs. 7-9, 11, 12): Head colored with dark

brown, redchsh browTi, or black pigmentation (Figs. 7,

11). Dark brown, gray, or black tentacles oval in cross-

section. Oral lobes equal to, or slightly longer than ce-

phalic tentacles. Both tentacles and oral lobes cihated

(Figs. 7, 9, 11). Eyes black, with well developed lenses,

posterior to tentacles on unpigmented, shght protuber-

ance (Fig. 11). Snout bifid and bilobed. Mouth vertical

slit, medial and ventral to head. Foot broad anteriorly,

mediallv bifurcate, wide, tapering posteriorly. Foot un-

pigmented, translucent, gray to white. Opaque crescent-

shaped ridge on dorsal surface of propodium (Fig. 11).

Mentum absent. Ventral surface of foot unpigmented

and highly cihated (Fig. 8), Pedal gland opens along mid-

dle of sole, producing a verv fine, transparent attach-

ment thread. Opaque visceral mass gray-white or brown,

with scattered and/or aggregates of black and/or dark

red-browTi cells.

Alinientan/ tract (Figs. 7, 11, 13-19): Gut extends pos-

teriorly from bulbous snout, under mantle floor to stom-

ach, located within visceral mass. CylindriciJ stomach

wide mediallv, tapering at junctions of esophagus and

intestine. Jaw, consisting of two rhomboidal sections

joined together at one of two long sides to form a tent-

hke stmcture, just posterior to buccal opening. Jaw cov-

ered with numerous, tightly interlocking scales. Radular

ribbon approximately 150|jLm long, 50(xm wide, with

about 30 rows of teeth (Fig. 13). Radula symmetrical,

with large rachidian teeth, each consisting of wide basiil

portion supporting a laterally rounded upper portion

(Fig. 14). Upper portion with medial indentation and

numerous fine serrations (Figs. 14, 15). Basal portion of

lateral teeth broad, with numerous serrations. Anterior

portion hook-shaped, serriform (Figs. 13, 16, 17). Mar-

ginal teeth similar to lateral teeth, with broad base and

hooked serrated anterior portion, with cuspidate crest

(Figs. 18, 19).

Pallial eaviti/ (Figs. 11, 12): Short, spacious palhal cav-

ity. Anterior mantle edge folded, finely crenulate, with

black pigmentation arranged as stripes. Pigmented hy-

pobranchial gland (=pigmented mantle organ? sensu

Robertson, 1985), within mantle roof, exudes copious,

thick blue-white exudate that mainlv' e.xits right side of

Figures 1-6. Scanning electron niicrophotographs of the shell and operculum of Rissoella caribaea. 1. Apertural view of shell

(scale bar = 200 (xm). 2. Dorsal view of shell (scale bar = 150 (xni). 3. Enlargement of aperture (scale bar = 100 jxm). 4. Apical

view of protoconch ( scale bar = 50 |xm). 5. Attachment surface of operculum (scale bar = 100 jjim). 6. Outer surface of operculum

(scale bar =100 \Lm).

Page 16

THE NAUTILUS, Vol. Ill, No. 1

Figure 7. Living Rissoelhi carihticn (.scale bar = 250 |xni). Figures 8-10. Scanning electron niicrophotographs of critical-point

dried R. caribaea. 8. Propodium/sole (.scale bar = 25 |xm). 9. Penis on neck posterior to cephalic and labial tentacles (scale bar =

50 (jLni). 10. Penis (scale bar = .30 |xni) (c = ciliated strip, ct = cephalic tentacle, f = foot, hbgl == hypobranchial gland, niae =

mantle edge, ol = oral lobe, p = propodinm pe = penis, sn = snout).

mantle, however, occa.sionally emitted from the left side,

a result of cilia that covers left mantle edge laterally and

extend one fourth of the way across anterior mantle edge

(Fig. 12). Majority' of exudate is moved out of the right

side of mantle cavity by ciliated strips. Pigmented hx-

pohranchiiil gland shape is \arial)le, comprised of large

cells filled with thick yellow-white contents (released

when sn;iil is disturbed). Pigmented hypobranchial gland

divitied into partitions, with cells iorming an oblong or-

gan having one or several salient projections (Figs. 7,

11). Projectiems arranged generally peipendicular to the

long axis of the gland. Perimeter of hyjiobranchial gland

may be open (not continuous) or cntirclv closed. Ventral

and dorsal ciliated strips (right side of mantle cavit\) join

posteriorlv but extend only a short distance into mantle

cavitv (Fig. 12). Water flow is from left to right. Large

pallial heart, yvith contractions at about 80/min. No pal-

lial kidney yvas found. Rectum-anus on right side, near

posterior end of mantle cayits'.

Rcprodudivc st/.stcm (Figs. 9, 10, 12): Simultaneous

hermaphrodite. Pallial portion of reproductive tract con-

sists of both male and female counteqiarts. Penis on

right side of mantle floor on snail's neck (Figs. 9, 12).

Penis attenuated anteriorly, with bulbous posterior and

scattered cilia (Fig. 10). Vas deferens joins bullions por-

J. B. Wise, 1998

Page Vi

11

12

Figures 11-12. Diagrams of the anatoniv of RissocUa caribaea. 11. External anatomy of fi. carihaea (scale bar = 250 jji,m). 12.

Mantle cavit\' and reproductive tract of fi, caribaea (scale bar = 250 fjim). (abg = albumen, eg = capsule gland, cs = ciliated strip,

ct = cephalic tentacle, fc = fertilization chamber, hbgl = hvpobranchial gland, hd = hennaphroditic duct, mae = mantle edge, ol

= oral lobe, o\d = o\iduct, ovt = o\'otestis, pe = penis, sn = snout, vd = vas deferens, vm = \isceral mass).

tion, extends posteriorly through mantle floor to fertil-

ization chamber. Fertilization chamber joins hermaph-

roditic duct that extends posteriorly to join ovotestis

within visceral mass (Fig. 12). Hermaphroditic duct

serves as a conduit for eggs and spemi and possibly as

seminal vesicle, as endogenous sperm are often present.

Ovotestis coils within visceral mass on columellar side.

Oviduct travels posteriorly from fertihzation chamber

turns and extends anteriorlv, becoTning part of capsule

gland. Female duct terminates as a short papilla on the

mantle floor, immediately posterior and on the right side

of penis (Fig. 12). Double-lavered egg capsule typically

contains two embryos. Larval development unknown.

DISCUSSION

Rissoella caribaea was first described by Rehder (1943a),

who stated that "It lives in sparselv populated colonies

on clean rocks, actively moving about, its white tentacles

contrasting stronglv with the black body, which is visible

through the idmost transparent shell." Later Robertson

(1961a) described tlie operculum, radula, and external

morphology of R. caribaea. He determined that the shell

and radula of R. caribaea (type species of the subgenus

Phijcodrosus Rehder, 1943) closely resembles that of R,

zebra (t\pe species of the subgenus Jcjfrei/silla Thiele,

1925). The difference he noted were that the rachidian

of R. caribaea is more narrow, with "untoothed cusps '.

On the basis of his comparison of these two species, he

determined that the subgenus Phycodrosus is a junior

svnonvnn of Jeffrei/silla.

When the anatomy of R. caribaea (this study) is com-

pared to the other rissoelUd species for which the inter-

nal anatomy is k-nown, there are a number of similarities

and differences (Fretter, 1948; Simone, 1995). In 1948,

Page 18

THE NAUTILUS, Vol. Ill, No. 1

Figures 13-19. Scanning electron niicrophotograplis ot the radiila of Ri.ssdclhi caribaca. 13. Radiilar nlibon, each row consisting

of a rachidian, and a single lateral and marginal toolli on each side (scale bar = 10 (xni). 14. Racliidian tooth (.scale bar = 10 (xni).

J. B. Wise, 199S

Page 19

Fretter descrihed the liiologv and anatoniN- of hvo ris-

soellids {Rissoclla diapluina and R opaliua ) inhabiting

the coralhne tidepcjols of the British Isles. She deter-

mined that although their internal anatomy is similar, the

two species are easily distinguished by shell shape, ten-

tacles, and snout moq^hologv'. As in the rissoellids de-

scribed by Fretter (1948), Rissoella caribaca exliibits in-

traspecific variation in the amount of pigmentation and

color of the exposed body parts ( = headfoot) and vis-

ceral mass, which is \isible through the transparent shell.

All rissoellids have a pair ot cephalic tentacles antl a p;iir

of oral or triangular lobes that Fretter (1948) and Si-

mone (1995) described as bifiucations of the snout. The

oral lobes (Figs. 7, 11) oi RissocUa caribaca are separate

from the bifid snout, as illustrated b\' Ponder and Yoo

(1977) for R. (Rissoclla) atriiiiaciila from S.W. Austraha.

In most species, the oral lobes are shorter and slightly

wider than the tentacles. Rissoellids have well developed

black eves, containing large lenses. The eves are poste-

rior to the tentacles and high on the head, atop a shght

protuberance, that in R. caribaca is surrounded by an

area of unpigmented tissue (Fig. 11). Simone described

the eves as being on short stalks, however his illustra-

tions of Rissoclla oniaia show the eyes to be on similar,

slight protuberances, and not stalks.

The pedal gland of R. caribaca produces an attach-

ment thread, which it uses to secure itself to the sur-

rounding substratum. No attachment thread was ob-

served in R. (liaphana or R. opalina. However, the pedal

gland produces a secretion, probably mucus, which is

"essential to the locomotor mechanism of the snail"

(Fretter, 1948; Fretter & Graham, 1962). In R. oniafa,

as in other taxa, the pedal mucous gland opening is lo-

cated in a medial, longitudinal sht in the posterior half

of the foot's ventral surface. No attachment thread or

mucus production was reported for R. omata. because

no li\ing snails were a\;iilable to Simone (1995).

The mantle cavitv of rissoellids is reduced in size by

the large capsule gland and duct that projects dorsally

from the body wall. As in the hvo species described by

Fretter (1948) neither a giU nor an osphradium is pres-

ent in R. caribaca. Simone (1995), howe\er, reported the

presence of "gill vestiges," with functional ciha and vari-

able leaflet number and shape in R. omata. These giU

vestiges are located on the left side of the mantle roof,

just posterior of anterior mantle edge. As in R. opalina

and R. diphana the rectum-anus of R. caribaca is located

on right posterior portion of the mantle floor. However,

unhke R. caribaca, the two British taxa have two rows

of ciliated strips extenchng anteriorly from each side of

rectum; one on the roof and one on the floor of the

mantle caxdty. In R. caribaca. both ventral and dorsal

ciliated strips extend only a short distance into the man-

tle cavits' where they join (Fig. 12). Furthermore, in R.

opalina and R diapJiana underKing each cihated strip

is a broad band comprised of secretoiy cells. In partic-

ular, the dorsal band is broader and composed entirely

of mucous cells, which Fretter (1948) suggested may

function as the hvpobranchial gland. No bands of mucus

producing cells were observed within the cihated strips

oi R. caribaca. Dissimilarly, the rectum/anus of R. ornata

apparently extends along the right side of the mantle

caNatv to terminate in what Simone ( 1995) described as

a special fecal chamber. This chamber is located at the

right comer of the mantle edge. From his illustrations

it appears that Simone (1995:figs. 9, 10) confused the

ventral and dorsal ciliated strips (similar to those in R.

caribaca). with what he respectivelv defined as head flap,

and folds, both peq^endicular and parallel to the mantle

border. The hvpobranclual gland of R. omata, located

posterior to these structures within the mantle roof is

nf)t present in R. caribaca. Although no Iddnev was

found in R, caribaca, it is present within the mantle cav-

it)' in both R. opalina and R. diaphana, opening near the

anus (Fretter, 1948; Fretter & Graham, 1962)" In R. or-

nata, the kidne\' is between stomach and albumen gland

within the \isceral mass (Simone, 1995).

The alimentarv tract of R. caribaca extends posteriorly

from its bifid snout to the jaw and then the radula (Fig.

13). The jaw of R. caribaca is ver\' similar to the jaw of

R, omata (Simone, 1995). The radulae of R. opalina and

R. diaphana (not figured) are simflar to diat of R. cari-

baca. However, Fretter (1948) stated that the lateral and

marginal teeth are absent and replaced by intermediate

teeth. The radula of R. ornata is short (10-12 rows),

each row containing a single rachidian, with approxi-

mately 10 equal sized cusps and one pair each of cus-

pidate lateral and marginal teeth. Esophagus, stomach,

digestive gland and intestine-anus are quite similar for

these taxa, the only cUfference being the position of the

rectum-anus (see abo\'e).

Rissoelhds are hermaphrodites, simultaneously pro-

ducing both eggs and sperm, within the same tubules of

the ovotestis, although Fretter (1948) reported that

sperm mav mature first. The gonadal duct ( = hermaph-

rodite duct, Fretter, 1948; Fretter & Graham, 1962; Si-

mone, 1995) or hermaphroditic duct (this study) extends

anteriorlv from the gonad to join the fertilization cham-

ber in R. caribaca (Fig. 12). In contrast, the hermaph-

rodite duct in R. onmta, R. opalina, and R diaphana

splits into a vas deferens and an albumen gland. The vas

deferens passes anteriorly beneath the palfial oviduct, to

the left of the midtUe of the mantle floor That portion

of the vas deferens just prior to the duct's terminus is

the glandular prostrate. The hermaphroditic duct ap-

pears to function as the seminal vesicle in R. caribaca

15. Enlargement of rachidian cutting edge, top portion (scale bar = l(xni) 16. Lateral and marginal teeth (scale bar = 10 (xm).

17. Fine serrations on sides ot lateral and marginal teeth (scale bar = '2 \x.m). 18. Crested marginal teeth (scale bar = 2 jjim). 19.

Enlargement of single marginal crest (scale bar = I jjim).

Page 20

THE NAUTILUS, Vol. Ill, No. 1

and in the species described by Fretter (1948). Fretter

(1948) described a deep gutter, on the median side of

the ventral longitudinal strip, into which the feniiile duct

opens "at summit of long papilla." At the anterior part

of the female duct terminus is the folded penis sitting

in a grooN'e, with its tip immediately anterior to the fe-

male aperture. Fretter and Gndiani (1962) also de-

scribed a third conduit that comprises a muscular sac

connected to the posterior end of the capsule gland by

a short duct and opens to the mantle cavity via a longer

duct. They suggested that it removes excess "secretions

and sperm" from the genital duct and is homologous

with a receptaculum seminis. In R. carihaca, the penis

is on the right side of the mantle floor on snail's neck

(Figs. 9,10) and not within a groove. The reproductive

tract in R. onmta is similar to that in fi. opalina, however

its vas deferens extends anteriorly on the right side of

the head and ends as a short tubular penis. The albumen

and capsule glands of R. onmta are very similar to those

present in R. carihaca, R. diaphana, and R. opalina.

Although this study provides new insights into ris-

soelhd biology, and more specifically the anatomy of Ris-

soella carihaca, it is not possible (due to limited available

data) to assess its systematic position within the family.

As determined bv Robertson (1961a), RissocUa carihaca

possesses the following characters typical of the family:

1) oral lobes, 2) perpendicular opercular peg, 3) large

hypobranchial gland (=PMO? sensii Robertson, 1985;

Wise, 1996), and 4) symmetrical radula consisting ot a

single central tooth and one pair each of comb-shaped

lateral and marginal teeth. However, these features are

present in all known rissoeUids and cannot be used to

resolve relationships within the rissoellids. Moreover,

studies utihzing radular moq^hology to separate rissoel-

lid species into subgenera have seldom examined

enough specimens or species to reach well-supported

c(3nclusions. Geographical variation within a species, on-

togenettc changes, and whether or not tooth morj^hology

is a result of diet or phylogeny remain areas that are

underexplored for rissoelBds. Until a better understand-

ing of known rissoelhd taxa exists, relationships within

the genus or family cannot be robustly resolved.

ACKNOWLEDGMENTS

I thank R. Germon (USNM) for the loan of the holotype

of Rwsoclla carihaca, and Dr. M. G. Harasewych for

field assistance and logistical support. I am particularly

grateful to the staff at the Texas Children's Hospital,

Houston, Texas: J. Barrish for all his time at the SEM

microscope and in the darkroom; C. Langston, tbrector

of the histology laboratory and her staff for graciously

blocking, sectioning, and staining of snails; and Dr. J.

Hicks for allowing me to work in the pathology lab. I

also thank Dr P. Mikkelsen for the provitling the Thiele

papers. The critical comments of R. Bieler, R. CoUin, R.

Robertson, and an anonymous reviewer greatly im-

proved this manuscript. Lastly, this work would have

been possible without the support ;ind assistance ot Dr

M. Rice and the dedicated staff at the Smithsonian Ma-

rine Station at Link Port, Florida. This is paper is Smith-

sonian Marine Station Contribution no. 480.

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Thiele, J. 1925. Gastropoda der Deut.sciien Tief see-Expedition.

n. Verlag von G\istav Fischer in Jena.

Thiele, ]. 1929. Handhuch der Systematischen WeichtierkTui-

de. Teil. Verlag von Gusta\ Fischer in Jena.

Wise, J.B. 1996. Morphology ant! phylogenetic relationships of

certain pyTamidellid taxa (Heterobranchia). Malacologia

.37(2):44.'3-511.

THE NAUTILUS lll(l):22-44, 199S

Page 22

The MoUuscan Fauna of the Wawa River Region, Miskito

Coast, Nicaragua: Ecology, Biogeographical ImpUcations, and

Descriptions of New Taxa

Edward J. Petuch

Department of Geology

Florida Atlantic Uni\ersitv

Boca Raton, Florida 33431 USA

and

Department of Paleontology

The Graves Museum of Archaeology

and Natural Histor\-

Dania, Florida 33004 USA

ABSTRACT

Faunal surveys (1991-1992) of the biologically-unex|3lored

\\'awa Ri\er region of the Miskito Coast of Nicaragua ha\-e

shown that the intertidal and estuarine areas contain tour dis-

tinct molluscan assemblages; the Nvritina — Polijnu'sodn As-

semblage, the Littorina — Petricola Assemblage, the Agaronia-

Micromactra Assemblage, and the Hastula — Donax Assem-

blage, and that the shallow subtidal area contains a single as-

semblage, the Ficus-Pacipecten Assemblage. A list of macrom-

ollusks found in these fi\e assemblages is giyen, and eight new

endemic mollusks are described. These include: (Gastropods)

Ficus villai n.sp. (Ficidae), Caiicellnrin mediaiucricnna n.sp.

(Cancellariidae), Conns pnschalli n.sp. (Conidae), and (Bi-

valves) Noetia (s.s.) lindae n.sp. (Arcidae), Plicatula miskito

n.sp, (Plicatulidae), Mactra inceri n,sp., Micromactra miskito

n.sp. (both Mactridae), and Petricola (Petricolaria) donnae

n.sp. (Petricolidae). This new faunal data, in conjunction with

previous Central American biogeographical studies, has dem-

on.strated the existence of a new molluscan faunal area within

the western Caribbean, here referred to as "Miskitia". The

Miskitian area, which may represent a new subprovince of the

Caribbean Molluscan Province, is distinctly Panamic-appear-

ing, containing molluscan assemblages that are dominated by

nomially-Eastem Pacific genera such as Agaronia, Noetin (s.s.),

Pnrijx-ftrn. and Micromactra

INTRODUCTION

Within the pa,st decade, faunal .survey.s in prexioiisly-

une.xplored areas of the Caribbean Sea region have un-

covered large number,s of new and unusual marine mol-

hisks (Petuch. 1987, 1988, 1990). Besides tniderscoring

our lack oi kTiowledge of such geographically-proximate

malacofaunas, these discoveries have shown that the Ca-

ribbean is not faunally homogeneous as was previously

thought (Warmke & Abbott, 1961), but is composed of

at least seven discrete, mutuallv-exclusive subregions

(Petuch, 1981,1982,1987,1988, 1997). Each of these

subregions has been found to contain a high degree of

endemism at both the generic and specific level and,

altogether, have demonstrated that the geographically-

small Caribbean Sea region contains the highest level of

molluscan diversitv found in the Americas.

While the coast;il faunas of the Yucatan Peninsula,

Honduras (Petuch, 1981,1987,1988; Yokes & Yokes,

1983), Costa Rica (Houbrick, 1968), and Panama (Olsson

& McCinty. 1958; Petuch, 1990; Radwin, 1969) have

been studied, the mollusks of the Caribbean coast of Nic-

aragua had never been adequately suneyed and the re-

gion has remained enigmatic. Over the past ninety years,

onh' a few scattered reports on Caribbean Nicaraguan

mollusks have appeared in the literature (reviewed in next

section), and these have only offered tantiilizing bits of

information that hinted at a very unusual and highly en-

demic fauna. In December, 1991 and July, 1992, while

working with the Caribbean Conservation Coi-poration of

Gainesville, Florida (under the auspices of IRENA (In-

stituto de Recursos Natuniles), the environmentiil agency

of the Nicaragtian government), I had the opportunity to

visit and collect ;ilong a part of the Miskito Coast near

the mouth of the Wawa Ri\er. This study, coincident;illy

conducted in the only area of die entire coast that had

ever been sampled (Fluck, 1905a, 1905b, 1905c, 1905d,

1905e, 1906; see next section), is part of a major biotic

survey that is being undertaken hy the Nicaraguan gov-

ernment in preparation of the establishment ot a wildbfe

and cultunJ protected zone along the Miskito ('oast.

The Miskito Coast was found to contain a highly un-

usual molluscan fauna, one that was both ahpical of the

Caribbean Molluscan Pnnince and that exliibited a high

degree of endemism. In many ways, the Nicaraguan Ca-

ribbean coastal m;ilacofauna more closely resembles the

faunas of the Eastern Pacific Panamic Province and the

eastern South American Brazilian Province than it does

E. J. Petuch, 199S

Paee 23

the fauna oi the surrounding Caribbean Province. This

Pananiic- Brazilian appearance was further emphasized

h\ the discinerv ol new endemic species, several of

which belong to genera that were thought to have been

restricted to the Pananiic and Briizilian Provinces.

In this paper, descriptions of the various coastal en-

vironments of the Wawa River area are given, along with

lists of the dominant macromollusks foimd in each hab-

itat. Eight new Misldto Coast species are described, in-

cluchng; Gastropods — Ficus villai n.sp. (Ficidae), Can-

ccHaria mcdiamericana n.sp. (Cancellariidae), Conus

paschaUi n..sp. (Conidae): Bivalves — Noetia liiulac n.sp.

(Arcidae), Plicatula iniskito n.sp. (Plicatulidae), Mactra

inceri n.sp., Micromactra miskito n.sp. (both Mactridae).

and Pctricflla (Pctricolaha) donuac n.sp. (Petricohdae).

A discussion of the biogeographical relationships of the

Misldto Coast fauna is given in the final section.

PiiKMors Mal.acolocuc.al Rese.-mk.h Ai.()N(. Tiik

MisKrro Co.AST Of Nic.-\r.\c.u.\

Prior to the present research, onlv one other attempt at

a comprehensive survev of the macromollusks of the

Miskito Coast had ever been undertaken. This pioneer

work was done bv the Rev. W.H. Fliick, a Moravian mis-

sionarv who ministered to the Miskito and Sumo Indians

at the tuni of the centurv. In a series of six papers pub-

lished in 1905 and 1906, Fluck gave interesting anec-

dotal insights into the environments and natural history-

of the area, molluscan food utilization bv the local In-

dians, and detailed descriptions of some of his tech-

niques for collecting mollusks. Most importantlv, he pro-

vided sever;il annotated species hsts, arranged phvloge-

neticallv, of the gastropods and bivalves that had been

collected near the mouth of the \\'awa River, W'ounta

and Bluefields Lagoons, and the offshore Man-of-War

and King's Kevs.

In four of his papers, Fluck lists fiftv-two species of

marine and estuarine mollusks from the Wawa River

area, one of which he described as new ( Strouibiis pug-

ilLs nicaragiicnsis Fluck, 190.5). Of these, five were not

encountered in the present study; the gastropods Tonna

maculosa, Poliniccs lacteus. Ci/matiiim pilearc. and Col-

iimhclla mcrcatoiia. and the bivalve Arcopa^ia fatista.

Here, arranged by publication date, are hsts of Fluck's

Wawa species, ;ilong with their updated ta.xonomic des-

ignations (where apphcable):

Fluck, 1905b. (SlicU Collecting on the Moscpiito Coast

of Nicaragua— 11. The Nautilus 19(2):16-19).

Fluck's Ta.\on

Murex brevijrons Lamarck

Murex bellus Reeve

Cijmatimn pilearc Lamarck

Piii-fnira floiidana Conrad

Fasciolaiia tulipa Linnaeus

Mdontiena incloni^cna Lin-

naeus

Nassa vibex Say

Updated Nomenclature

Chicoreiis incr^us E. \'okes

Vokcsimurex garcmi (Petuch)

Tiiai.s floridana (Conrad)

Na.ssnrins vibex (Sav)

Valuta virescens Solander

Marffnella apicina Menke

Mar^mclla ffittata Dillwvii

Oliia( Aoflron ia ) Irstacea

Lam.

Columbella im-rratoria Lin-

naeus

Columbella [Anachis) lyr/ila

Sow.

Cancrllaria reticulata Linnae-

Terebra i

■rea GmeJi;

Valuta lacertina Petuch

rniimin pniino.suin (Hinds)

Pntnuin ffittatuin (Dillwvai)

Anarania hilli Petncli

Costanachi.'i leledti (Ducios)

Cancellaria mcdiamericana

n.sp.

Hastula cinerea (Boni)

Fluck. 1905d (Shell Collecting on the Mosquito Coa.st

of Nicaragua— IV. The Nautilus 19(5):55-57).

Fluck's Taxon

Cypraea exanthema Linneans

Dalium perdix Linnaeus

Ftjrula papijratia Sav

Paliniees bninnea Luik

Palinices lacteus Gnilding

Crepidula conrexa Sa\'

Venniculaiia spirata Philippi

Litorina lineata Pliilippi

Litorina calumcllam D'Orb.

Litarina an^ulifera Lam.

Ncritina lineata \ar reticulata

Updated Nomenclature

Ci/praea zebra Linnaeus

Tanna mncula.sii Linnaeus

Fiais villai n.sp.

Palinices hepaticus (Roding)

Littorina lineata d'Orbignv

Littorina nebulosa (Lamarck)

Littarina an^ulifera (La-

marck)

yeritina piratica Russell,

1940

Fluck. 1905e. (Shell Collecting on the Mosquito Coa.st

of Nicaragua-V The Nautilus 19(7):7S-80).

Fluck's Taxon

Phalas campechiensi.^ Gmehn

Tagelus poeijii Dall

Periplanm inequivalvis

Schum.

Mactrat Mactrella I alata

Spengler

Mulinia ffiadelaupensis Re-

cluz

Updated Nomenclature

Tagelu.s plebeius (Lightfoot)

Periploma margaritaceum

(Lamarck)

Mactrellana alata (Spengler)

Mulinia clenjana (Orbigny)

Fluck. 1906. (Sliell Collecting on the Mosquito Coa.st of

Nicaragua-VI. The Nautilus 20(1): 1-4).

Fluck's Taxon

Tellina {Eun/tcllina) ans.ulosa

Gmelin

Tellina iCijclatellina) fausta

Pultene\

Sanouinalaiia i Psammatella)

operculatai Gmehn)

Strigilla canmria Linnaeus

Iphigenia brasiliana (La-

marck)

Danax denticulatu.s Linnaeus

Danax cat/ennensi.s Lamarck

Pitar (Hysteraconcha) diane

(Linnaeus)

Chione cancellata (Luuiaeus)

Ctjrena {Pohjmesoda) pana-

mensi.s Prime

Updated Nomenclature

Arcapagia fausta (Pultenev)

Sanquinolaria cruenta (Light-

foot)

Danax stiiatus Linnaeus

Pahpnesoda placans iHanleij)

Page 24

THE NAUTILUS, Vol. Ill, No. 1

14-1

-14°

Figure 1. Map of tlie Miskito Coast of Nicaragua, in the vi-

cinity of the Wawa River mouth, showing collection stations.

Distance between Stations 1 and 4 is appro.xiinately 11 km. 1

= shallow embayment on southern side of Bragman's Bluff

(containing Agaronia — Micromactra Assemblage); 2 = clay

cliffs and clay talus "reefs" innnediately north and south of

Puerto Cabezas (containing Littorina — Petricola Assemblage);

3 = beach on outer side ot Barra de Wawa, northern side of

Wawa River mouth (containing HasUila — Donax and Ficus —

Pacipecten Assemblages); 4 = beach on southern side of Wawa

River mouth, adjacent to Wawa Village (containing Hastiila —

Donax and Fiais — Pacipecten Assemblages); 5 = southern

shore of Laguna Karata, at Karata Village (containing Neriti-

na — Pohjmesoda Assemblage); 6 = mangro\e jungle at Sal-

onque de Kauropura (containing Neritiiut — PoUjnwsoda As-

semblage).

Tivela rniuiroiiles (Bom)

Chama CDHi^rt-fiata f^oiirad

Ami tiinbonatu Lamarck

Sca])harca incongnta var

braxiliana Lam.

Area occidentalis Philippi

Pinna muricata Linnaeus

Pecten dislocatus Sav

Ostrea virninirn Giiielin

Area iinbricata Bniguiere

Aniidina hriisiUaiui (Lamarck)

Area zebra (Swainson)

Atriiia semimida (Lamarck)

Paeipecten leueophaeiis

(Reeve)

Crassostrea virgitiiea (Gnie-

lin)

Since Fluck's .serie.s of papers, only a few scattered

references to the Mi.skito Coast fauna have appeared in

the Caribbean molluscan hterature. Most of these have

been in the Johnsotiia series of monographs published

at Har\ard University (1941-1974)(ie. Clench & Abbott,

1941; Bequaert, 1942, etc.), and were based upon the

Fluck Collection in the Museum of Comparative Zool-

ogy. No references to newlv-collected Miskito material

have been mentif^ned prior to the present study.

In the 1980's, I had received several suites of unusual

western Caribbean gastropods from local commercial

shell dealers, particularly Mr Leonard C. Hill of Miami,

Florida. Within these specimen lots were large numbers

of new species that had been dredged from shallow

depths (10-.35m) off Roatan Island and the northern

Honduras coast by commercial shrimp boats. Many of

these were subsequently described bv myself (Petuch,

1987) and were later discussed within a geochronological

and biogeographical framework (Petuch, 1988:149-156).

Although the bulk ot these new taxa were from the area

immediately north ol the Miskito Coast, a few have now

been collected in Nicaragua and have been found to be

ecologically-dominant Miskito organisms. Two of these

recently described gastropods, which range from the

Roatan Island-Punta Patuca area of Honduras to the

Nicaraguan Miskito Coast, include Agaronia hilli Pe-

tuch, 1987 and Vokcsimurex garciai (Petuch, 1987).

Following the legacy of the Rev. Fluck and his pioneer

research, I here report on the most recent malacological

survey of the Miskito Coast. In total, one hundred and

four species, from five distinct molluscan assemblages,

were collected, doubUng Fluck's total of fift)'-two spe-

cies. Of these, eight are new to science. The habitats

and ecological preferences of the molluscan assemblages

of the Wawa River area of the Miskito Coast are given

in the following section.

Coi.i.ECTiNc Stations And Cciast.\l En\'ironments

In order to giiin a general insight into the molluscan

faunas of the varied ct)astal habitats, sLx vvidelv-spaced

collecting sites were selected in the Puerto Cabezas-

Wawa River area (Figure 1). Four of these were along

the open coast, ranging from approximatelv 2.5 km north

of the m;un pier of Puerto Cabezas southward to the

southern shore of the Wawa River mouth. Two others

were from within the estuary of the Wawa River From

these six sites, altogether, five distinct molluscan assem-

blages could be discerned. The assemblages are named

after dominant or characteristic gastropods and bivalves,

one of each, and these include: 1. The Nciitina — Poly-

mcsoda Assemblage (brackish water estuarine), 2. the

Littorina — Petricola Assemblage (coastal clay chffs), 3.

the Agaronia — Micromactra Assemblage (protected

coastal, (|uiet intertidal), 4. the Hastula — Donax Assem-

blage (open coastal, high energy intertidal), and 5. the

Ficus — Pacipecten Assemblage (offshore, shallow subH-

dal). The physical en\ironment and molluscan assem-

blages of each collecting site are discussed herein.

Station 1. (Agaronia — Micromactra Assemblage)

An unnamed, widely open embayinent containing shal-

low, reiativelv quiet water conditions, was located at ap-

E. J. Petuch, 1998

Page 25

proximately 2.5 km north of Puerto Caliezas, between

the mouth of La Bocana Creek and Bragman's Bluff,

The southward curvature ot Bragman's Bluff ;Jong with

its accompanving sandstone and cla\' block talus reef,

both offer protection from strong easterly and north-

easterly winds and waves, allowing the creation of almost

lagoonal comlitioiis within the open embavment. These

quiet conditions, which are especiallv pre\alent imme-

diately south of the bluff, are further enhanced by the

presence of a large sand bar that rims par;dlel to the

coast just offshore.

This shallow (0-lm depth) environment, comprising

a Hat muddy-sand substrate and nuuky water contlitions

with only minimal wave action, contiiined a highly un-

usual, highly atypical Caribbean molluscan assemblage.

The dominant mollusks, found nowhere else in the Ca-

ribbean pnnince, give this assemblage a chstinctive East-

em Pacific appearance, and the entire comnmnity close-

ly resembles assemblages that are found on similar bio-

topes along western Central America. Characteristic

mollusks from the Bragman's Bluff open embavment lo-

cahty include:

Gastropods

Stminhit.s piigilis nicaraguensis Fluck, 1905 (Figure 6)

Agaronia hilli Petuch, 1987 (Figures 4, 5)

Conus poschalli n.sp. (Figures 2,3)

Bivalves

Noctio lindac n.sp. (Figures 9, 10)

Plicattila ini.skito n.sp. (Figures 16-19)

Diplodonta notata (Dall & Simpson, 1901)

Trachycardiiim muiicatum (Linnaeus, 1758)

Mactra inceri n.sp. (Figures 7, 8, 11)

Micromactrci miskito n. sp. (Figures 12, 13)

Mulinia clcnjaiia (Orbigny, 1846) (Figure 20)

Macoma brevifrons (Say, 1834)

Macoma const ricta (Bruguiere, 1792)

Tcllina punicea Bom, 1778

Sangninohiiia sangitinolcnta (Gmehn, 1791)

Periploina inargaiifacciim (Lamarck, 1801) (Figure 21)

Similar biotopes from around the Caribbean Basin all

contiiin assemblages that are typically dominated by oh-

vid gastropods of the genera Oliva (particularly mem-

bers of the Oliva reticularis Lamarck, 1811 species com-

plex; see Petuch & Sargent, 1986:120-122) and OlivcUa.

and bivalves of the genera Glycyineris, Divaricella, Co-

dakia, Anomalocardia, and the telhnids Tcllina radiata

Linnaeus, 1758 and Tcllina listen Boding, 1798. All of

these ta.xa were missing from the Bragman's Bluff col-

lecting station and were not found at any other Miskito

coast locahtv' (both this stud)- and that of the Rev. Fluck

in 1905-1906). Instead, the gastropod component at

Bragman's Bluff was dominated bv the oli\id genus Aga-

roiiia and the bivalve component bv mactrids {Mactra,

Micromactra, and Mulinia). The primarilv-Panamic ge-

nus Micromactra (represented by M miskito n..sp.), pre-

viously unknown in the Caribbean Province, was es-

peciallv common at Station 1.

Behind the sand bar, tlirectly off the beach south oi

Bragman's Bluff, large beds of dead bivalve shells have

accumulated. These beds, composed mostly of dead

Noctia valves, produce the proper substrate for the at-

tachment of the dwarf oyster, Plicafula miskito n.sp..

This new species, which closely resembles the Eastem

Pacific Plicatula pcnicillata Carpenter, 1857, further

demonstrates the Panamic appearance of the Bragman's

Bluff assemblage.

Station 2. {Littorina — Pctricola Assemblage)

From the main pier of Puerto Cabezas northward to the

mouth of Lagima Kruldra (approximately 8 km north),

the coast is characterized by high (20-50m) red and

white clay cfiffs (Figure 14). Lenses of rounded quartzite

pebbles are plentiful within the cfiffs and, upon erosion,

often form thick pebble beds at the cfiff bases. At several

localities immetliately north of the pier, and particularK'

adjacent to the mouth of La Bocana Creek, large talus

slopes composed of massive blocks of clay and soft sand-

stone have extended seaward and have, in effect, pro-

duced "rockv " intertidal en\ironments. These clay

"reefs" often extend into the subtidal region and offer a

wide variety of habitats within the vertical range. The

clay cfiffs biotope at, and immediately northward of

Puerto Cabezas, was chosen as the second collection sta-

tion.

The intertidal-shallovv subtidal clay cfiff and clay

"reef environment contained a very impoverished mol-

luscan fauna, with onlv a few species dominating the

area. Characteristic mollusks from the Puerto Cabezas

clav chffs locahtv include:

Gastropods

Diodora cai/cncnsis (Lamarck, 1822)

Littorina lincata Orbignv, 1842

Littorina uchulosa (Lamarck, 1822)

Littorina riisei Morch, 1876

Thais floridana (Conrad, 1837)

Bivalves

Pholas campcchicnsis Gmefin, 1791

Pctricola (Pctricolaria) donnac n.sp. (Figures 24, 25)

What was particularly impressive about die clay cfiffs

fauna was the noticeable absence of typical common Ca-

ribbean hard substrate taxa, manv of which were ex-

pected to have been encountered along this coast. Ab-

sent from the supratidid zone were the classicallv Carib-

bean fittorinid genera Tcctarius (Cenchritis), Nodilitto-

rina (Echinolittorina) and Echininus. Within the

intertidal zone, the absence of the neritid genera Lin-

ncrita, Theliosti/la, and Pupcrita, the potamicfid genus

Batillaria, the cerithiid genus Ccrithium (Thericium),

and the pulmonate genus Siphonaria was particularly

Page 26

THE NAUTILUS, Vol. 111. No. 1

Figures 2-13. M..llt.sks charactenstic of the A^nronin-Micromartra Asse.nhlac^e. 2, 3. Conns (Lcptoconus) paM "^-,^P'^"«^

58 mm- 7 8. A/«rtr« inceri new species, holot>pe, length 35.5 m>u; 9, 10, Noctin (s.s.) h.uhw new species, hoiotNpe, lei <Sh 25.5

^r^n Macira inceri new speces, parat>pe,- length 34 nun ipal!>al hne outUned u.th „>k); 12, 13. M.cromaCra nusk.to new

species, hol()t>pe length .30 niiu.

E. J. Petuch, 1998

Page 27

^

^

Figure 14. Aerial \ie\v of clav cliff coastline in the vicinitv' of Puerto Cabezas. Arrows point to clay talus "reefs" near the month

of La Bocana Creek, on the northern side of the citv. Station 2 included these cliffs and "reefs" and others inimediatelv to the

south (to the left). The holot\pe of Pctricola (Petricolaria) doimac n.sp. was collected on the beach between the first two arrows

on tlie left.

noteworth). Instead, the entire intertidal zone was dom-

inated by only three species of Littorina, of which Lit-

foriiia nchiilo.sa (Lamarck, LS22) was the most abundant.

The sliallow subtidal zone of the clav "reefs ' exposed

at extreme low tides, was relatively barren externally, but

was riddled yvith immense numbers of holes. The holes

represented the burrows of the pholadid and petricohd

bivalves Pholas campcchicnsis Gmelin, 1791 and Pctri-

cola (Pctricolaria) donnac n.sp., and these two burrow-

ers were the dominant organisms in this area. The large

carnivorous muricoidean gastropod Thais floridana

(Conrad, 1S.37) yvas also frequently encountered on the

subtidal clay "reefs", and several specimens were found

to be teethng on the hmpet Diodora caijcncnsis (La-

marck, 1822).

Stations 3 and 4. (Hastida — Donax Assemblage and

Ficiis — Pacipecten Assemblage).

South of the Puerto Cabezas pier, the high clav chffs

disappear, and the coastline becomes flattened with

wide, jungle-lined beaches and high-energv surf zones.

Along these beaches two distinct molluscan assemblages

accumulate; a Hflst(//fl-Do/)a.v-dominated assemblage

that li\'es in the sandv intertidal zone amid the heavy

surf, and a Pacipecten dominated assemblage that hves

in the subtidal area directh- offshore (approximately 0.5

to 1 km otf the coast, in depths of 10-25m). Most of the

components of the former assemblage can be coUected

within the surf at low tide and their dead, beached shells

are unusually well-preser\'ed. The components of the

latter assemblage, on the other hand, only wash up onto

the beach after storms or heavy blows, and their shells

are most always in a smashed, fragmentary condition.

Preliminary information on the composition of Carib-

bean beach communities (ie. Warmke & Abbott, 1961:

17,19; Houbrick, 1968) and western Caribbean offshore

communities (Raduin, 1969:229-236: Petuch, 1987:62-

63; Petuch, 1990) made it possible to determine yvhich

of the two assemblages represented the home commu-

nities of all beach specimens collected.

The high-energ\' beach emironment, the biotope of

the Hastula-Donax Assemblage, houses die most typical

Caribbean fauna found along the Miskito Coast. For

study, hvo beach areas that particularly characterized

this biotope were selected on either side of the W'awa

River mouth; with Station 3 located on the northern side

along the outer coast of the Barra de Wawa, and with

Page 28

THE NAUTILUS, Vol. Ill, No. 1

Figure 15. View of the beach along the southern side of the Wawa River nioutli. Piles of driftwood, brought down bv the strong

river currents, line the beach for several Idlonieters. The Fig Shell, Finis villai n.sp.. and the scallop, Pacipecien leucophneus (Reeve,

1852), were especially common along this area. Station 4 included this localitv and the open beach emironment in the distance.

This is also the t^-pe localit}' of Fluck's Stroiuhus pui^ilis nicnragucnsis (whicli is plentiful as fresh beach specimens).

Station 4 located on the southern side adjacent to Wawa

Village (Figure 15). Characteristic mollusks from the

Wawa surf locahty include:

Gastropods

Hastula cincrca (Bom, 177(S)

Bivalves

Solen ohliquus Spengler, 1794

Stripilla canmria (Linnaeus, 1758)

Striiiillii <^al>hi Olsson and McGinty, 195S

Sthjiilla mirahili.s (Philippi, 1841)

Strii^^itla pi.sifonnis (Linnaeus. 1758)

Stri^illa pwditcta (Tryon, 1870)

Strifiillfi pscudocnnidiia Boss, 19fi9

Tellina sijharitica D;J1, 1881

Donax dcnticulatus Linnaeus, 1758

Donax mediamericamis Pilsbry, 1919 (Figure 43)

Donax .striatum Linnaeus, 1767 (Figure 42)

Iphi'^cnia hra.siliaiia (Lamarck, 1818)

Tagchts plebeius (Lightfoot, 1786)

Pitar dionc (Linnaeus, 1758)

Tivcia mactroidc.s (Bom, 1778)

With the exception oi Donax mediamericanus Pilsbry,

1919, which is endemic to the Caribbean coast of Cen-

tra] America (type locality, Li\ingston, Guatemala), all of

the other members oi this assemblage are wide-ranging

tropical western Atlantic suri-loving species. Although

the fauna! composition of this assemblage is, b\' itself,

unremarkable, the abundant presence of six sympatric

species of the tellinid genus StriojUa and three species

of large Donax is noteworthy. Other Caribbean surf eco-

Figure 16-29. .Vlcjllusks characteristic of the Agaronia — Micronmctra. Llttorina — Petricola, and Neritina — Poliptiesoda Assem-

blages. 16, 17. Plicattila miskito new species, upper valve of holotype, length 13 mm; 18. Plicatula miskito new species, interior

view of bottom valve of holotvpe attached to interior of Noetia liiulnc valve. Arrow points to spond\l()id hinge; 19. Plicatula miskito

new species, upper valve of holotvpe attached to bottom \alve in life position, within Woititi lindac \aKe; 20. Miiliiiiti rlen/ann

E. J. Petuch, 199S

Page 29

Is-

20

19

23

22

24

25

27

28

29

(d'Orbignv, 1846). length 23 mm; 21. Periploma margnritaceum (Lamarck, 1801), interior of left \-aI\-e, length 31 mm; 22. Cras-

sostrea rhizophorae (Guilding, 1828) upper specimen length .35 mm; 23. Petricola (Petricolaria) pholadifonnis (Lamarck, 1818),

length 35 mm. From beach at Lake \\'orth, Floricia, (for comparison with Petricola donnne). 24, 25. Petricola (Petricolaria) donnae

new species, holot>pe, length 28 mm; 26, 27. Crassostrea virginica (Gmelin, 1791), length 97 mm. Elongated \'ariant that resembles

Crassostrea labelleensis Olsson & Harbison, 1953 from the Floridian Pleistocene; 28. Neritina piratica Russel, 1940, length 13 mm;

29. Polijmesoda placans (Hanley, 1845), length 28 mm.

Page 30

THE NAUTILUS, Vol. Ill, No. 1

Figure 30. \'iew of a section of the reil mangrove (Rliizophora nian^lc) forest at Salonque de Kauropura, near Lagiina Karata.

The arrow (extreme lower left center) points to a small clnnip of the mangrove oyster, Crassostrea rhizophomc (Gnikling, 1828).

The nerites Ncritiiw piratica Russell, 1940 and Neritina recUvata (Sav, 1822) were common here on submerged roots and o\ster

clumps. On the higher dn,- branches and roots, the periwinkles Litforinn ndndosa (Lamarck, 1S22) and Liftorina anffilifcra (La-

marck, 1822) and the melampid Melampus coffeus (Linnaeus, 1758) could be found in abundance. Open muddy-sand bottom areas

between mangrove forests housed large, dense colonies of the corbiculid bi\aKe Polymesoda placans (Hanley, 1845). This area

encompassed Station 6.

Figures 31-45. Molliisks characteristic ot the Finis — Pacipecien and lliLstula — Donax Assemblages. 31, 32. Cancdlaria iiu-ili-

anwricana new species, holotyjie, length 34 nmi; 33, 34. Fiais villai new species, holotype, length 67 mm; 35. Det;ul of shell

sculpture of holotvpe of Ficiis villiii: .36. CiinrrUaria n-tirulatn (Linnaeus. 1767), length 27 mm. From beach at Sanibel Is,, Florida,

E. J. Petuch, 1998

PasJe 31

35

41

45

for comparison with Canccllaria uu-diamcrieana: 37. CnnccUaria pctuclii Harasewvch. Petit & Verhecken, 1992, length 35 mm.

Trawled from 25 ni depth off \'ictoria, Espinto Santo, Brazil, for comparison with Canccllaria mcdiamericana: 38. Pacipcctcii

Icucopliaetis (Ree\e, 1S52), length 38,5 mm; 39. Ccrithioclava aarciai Houbrick, 1985, jnvenile specimen, length .34 mm; 40. Fictis

lindac Petuch, 1988, length 99 mm, Trawled from 20 m depth off the Monges Islands, Gulf of N'enezuela, for comparison with Ficus

villai; 41. Detail of shell sculpture of Ficus lindac; 42. Donax strintiis Linnaeus, 1767, lengtli .34 mm; 43. Donax mediamcricanus

Pilsbry, 1919, length 30 mm; 44. Ficus communis Roding, 1798, length 67 mm. From beach at Sanibel Is., Florida, for comparison

with Fin/5 lillai; 45. Detail of shell sculpture of Fictis communis.

Page 32

THE NAUTILUS, Vol. 111. No. 1

systems usually coutaiii oue or two Doiiax species and

only two or three StiiiiiUa species.

The other moUuscan assemblage found along the

Wawa beaches, the Ficits-Pacipccten Assemblage, is

probably only partially represented, with its community

structure data being biased toward those species whose

shells most readily accumulate along the shore after

stonns. Neyertheless, sufficient iniormation exists about

this intriguing and unstudied ecosystem, mostly gleaned

from beach deposits, to show that this offshore com-

munity is, in many ways, as atypical of the Caribbean

region as is the Agaronia — Mkromactra Assemblage of

shallower water areas. Characteristic moUusks of the

Miskito Coast offshore, subtidal area include:

Gastropods

Turbo castanca Gmelin, 1791

Vermiculaiia spirata (Phihppi, 18.36)

Ccrithioclava garciai Houbrick, 1985 (Figure 39)

Crepidula coiwexa Say, 1822

Xcnophora conchyUophora (Bom, 1780)

Cypraea (Macrocypraea) zebra Linnaeus, 1758

Pseudocyphoma intermedia (Sowerby, 1828)

Naticarius canrena (Linnaeus, 1758)

PoJinices hepaticus (Roding, 1798)

Phaliitin granulattim (Bom, 1778)

DiMorsio robinsoni Petuch, 1987

Tonna galea (Linnaeus, 1758)

Ficus villai n.sp. (Figures 33, 34, .35)

Chicorcus hrcvifrons (Lamarck, 1822)

Chicoreus mergiis (E.Vokes, 1974)

Vokesimurex cahtittii (Bernardi, 1859)

Vokesimurex garciai (Petuch, 1987)

Talityphis expansus (Sowerby, 1874)

Costanachis vclcda (Duclos, 1846)

Nassaritis vibex (Say, 1822)

Fasciolaria ttdipa (Linnaeus, 1758)

Fusinus dowianus (Olsson, 1954)

Vasmn miiricatiim (Bom, 1778)

Valuta laceriina Petuch, 1990

Cancellaria mediamericana n.sp. (Figures 31, 32)

Prunum guttatum (Dillwyn, 1817)

Pntiiuin pniiiiosum (Hinds, 1844)

Conufi aureopunctatus Petuch, 1987

Conus daucus Hwass, 1792

Conns garciai daMotta, 1982

Conus rosemaryae Petuch, 1990

Conus spurius Gmehn, 1791

Terchra tauriiia Lightfoot, 1786

Bivalves

Area imhricata Bruguiere, 1789

Area zebra (Svvainson, 1833)

Anadara hrasilianu (Lamarck, 1819)

Anadara ovalis (Bruguiere, 1789)

Barbatia doniingensvi Lamarck, 1819

Caloosarea notahilis (Roding, 1798)

Atrina seiniiiuda (Lamarck, 1819)

Argopecten gibbtis (Linnaeus, 1758)

Paeipectcn Icucophaeus (Reeve, 1852) (Figure 38)

Lopha ecjuestris Say, 1834

Diplodonta seniiaspcra (Phihppi, 1836)

Chaina congregata Conrad, 1833

Laevicardium lacvigatum (Linnaeus, 1758)

Maetrellona alata (Spengler, 1802)

Tellina augultisa Gniefin, 1791

Tellina similis Sowerby, 1806

Sanguinolaria cntenta (Lightfoot, 1786)

Chione eaucellata (Linnaeus, 1767)

Chione (Lirophora) latilirata (Conrad, 1841)

Chione (Lirophora) paphia (Linnaeus, 1767)

Pitar circinatiis (Bom, 1778)

Corbula confracta Say, 1822

Because of their thin and Hghtweight but yet struc-

turally-strong shells, Ficus villai n.sp. and Pacipecten leu-

cophaeus (Reeve, 1852) (single valves) are collected

abundantly along the beach, most often in perfect con-

dition. Judging from its abundance as beach shells, the

western Caribbean endemic Pacipecten Icucophaeus

must form dense beds just offshore, much as the closely-

related analogue species Pacipecten tumbezensis

(d'Orbigny, 1846) does along the subtidid areas off west-

em Central America. The Pananiic appearance of the

Miskito Coast subtidal assemblage is further enhanced

by the presence of Ficus villai n.sp. (possibly a close

relative of the Panamic Ficus vcntricosa (Sowerby,

1825)) and by Cancellaria mediamericana n.sp. (resem-

bling members of the Panamic Cancellaria ohcsa Sow-

erby, 1832 species complex). The subtidal assemblage

also shows a close relationship to the faunas of Carib-

bean Honduras to the north (Petuch, 1987; Petuch,

1988:15.5-156) and to Caribbean Panama in the south

(Petuch, 1990), especially in containing such distinctive

western Caribbean endemics as Ccrithioclava garciai

Houbrick, 1985, Vokesimurex garciai (Petuch, 1987),

Fusinus doivianus Olsson, 1954, Voluta laceriina Petuch,

1990, Conus garciai da Motta, 1982, Conus rosemaryae

Petuch, 1990, and Conus aureopunciatus Petuch, 1987.

Stations 5 and 6. (Neritina — Polymesoda Assemblage)

The Wawa River and its numerous small tributaries ex-

pand into three large lagoons before coalescing and nar-

rowing into a single large final channel. These lagoons,

the Lagima Yulu-Laguna Karata complex (to the west of

the m;un Wawa channel) and the smaller Lagima Kau-

ropura (to the east of the main channel) come together

near the Salonque de Kauropura, producing a rechan-

nehzed Wawa River mouth diat empties into the Carib-

bean Sea appro.ximately 11 km south of Puerto Cabezas.

The headwaters of these lagoons are essentially fresh-

water, iiecoming brackish in the southern part of Laguna

Karata and giadually becoming more saline at the Sal-

on(jue de Kauropura. Open oceanic safinity does not ap-

pear until well past the Barra de Wawa at the river

mouth.

E, J. Petuch, 1998

Page 33

Since these brackish coastal lagoons are virtually unex-

plored biologicallv, t\vo different localities were selected

within the lagoonal svsteni; one along the shghtly brack-

ish southern Laguna Karata near Karata Village, and one

along the more sahne Salontjue de Kauropura. The

shoreUnes of the lagoons and fin;il river channel are hned

with dense thickets of the red mangrove [Rliizophora

inau<ilc) (Figure 30) and most of the molluscan fauna is

mangrove-associated. Open channel and lagoon bottoms

are covered with nniddv-sand, and ovster bars occur in

deeper areas. Characteristic moUusks from the Karata

and Kauropura estuarine locahties include:

Gastropods

Neritina piratica Russell, 1940 (Figure 28)

Neritina recliiata (Say, 1822)

Littorina aniiithfcra (Lamarck, 1822)

Littorina nchulosa (Lamarck, 1822)

Cerithkica plictilosa vcracnizcnsis Bequaert, 1942

Melongena melongena (Linnaeus, 1758)

Mdampus coffcus (Linnaeus, 1758)

Bivalves

Crassostrca rluzophorac (Guilchng, 1828) (Figure 22)

Crassostrea virginica (Gmehn, 1791) (Figures 26, 27)

Poh/mcsflda placans (Hanlev, 1845) (Figure 29)

Although taunisticallv imposerished, the Wawa estu-

ary is probabl)' tvpical of the Miskito coastal lagoonal

system, and shows a degree of endemism. The presence

of Neritina piratica Russell, 1940, Cerithidca plictilo.sa

veracnizensis Bequaert, 1942, and Poh/mcsoda placans

(Hanley, 1845), all of which are restricted to the western

Caribbean, demonstrates the close relationship of the

Wawa assemblage with drose to the north (Honduras)

and to the south (Costa Rica).

Within the final channel of Barra de Wawa, a large,

slender variant of Crassostrea virginica (Gmehn, 1791)

(resembhng the Floridian Pleistocene C.labcUccnsis Ols-

son & Harbison, 1953) forms massive reeflike bars in

deeper, more sahne water. The large carnivorous gastro-

pods Thais jioridana (Conrad, 1837) and Melongena me-

longena (Linnaeus, 1758) are common on these bars and

are the primary predators of the oysters.

SYSTEMATIC SECTION

The holotypes and paratopes of the following new spe-

cies are deposited in the tvpe collection of the Division

of MoUusks, Section of Invertebrate Zoology', the Car-

negie Museum of Natural History, Pittsburgh, Pennsyl-

vania. A voucher collection of at least one specimen of

each species Usted has also been deposited in the m;un

collection of the Carnegie Division of MoUusks.

SYSTEM ATICS

Gastropoda

Caenogastropoda

Tonnoidea

Ficidae

Ficus Roding, 1798

Ficus villai new species

(Figures 33, 34, 35)

Description: Ficus of average length for genus (adults

av. 62 mm), elongated, pyriform, belonging to Ficus

communis Roding, 1798 species complex. Shell orna-

mented with thick spiral cords that are better developed

than longitudinal cords. Spiral cords colored with irreg-

ular, small, pale brown and white dots. Some specimens

with broad, dark browii longitiuhnal flammules, produc-

ing "zebra" pattern. Shell thin, hghtAveight, but struc-

turally strong. Shoulder rounded, gently sloping. Suture

depressed, subcanaficulate. Spire elevated above shoul-

der line, shghtly sc;ilariform. Shell surface covered with

evenly-spaced, intersecting longituchnal and spiral cords,

producing fenestrate patterns. Spiral cords much stron-

ger and better-developed than longituchnal cords, ap-

proximately four times as thick (Figure 35). Thinner, fine

secondarv cord present between each pair of thick pri-

mary cords, with secondary cord being appro.ximately as

thick as thin longitudinal cords. One or two extremely

fine, hairhke, tertiary spiral cords present on either side

of secondary spiral cord, giving areas between thick pri-

mary cords a silk^v appearance. Average specimen (based

on tvpe lot) with 26 large primary cords on combined

body whorl and siphonal canal. General apertural shape

elongatelv ovate, tapering gradually into open siphonal

canal. Aperture proportionally wide, approximately equal

in size to ventral half of body whorl. Edge of labrum in

adult specimens (such as holotype) thickened, reinforced

with thin, white, callus-like deposit. Well-preserved

specimens show slight undulation along labral edge, with

raised areas corresponding to edges of large, primary

spiral cords. As typical of all known members of Ficus

communis species complex, protoconch bulbous, mam-

millate in appearance, smooth, consisting of iy2 whorls.

In fresh specimens, color somewhat variable, generally

tan or pale brown with large, wide brown, longitudinal

flammules of varving intensities (such as on holotvpe.

Figures 33, 34). On many specimens (such as holotvpe),

primary cords exliibit fiiint pattern of alternating pale

wliitish-tan and fight brown elongated spots. Depressed

subsutural area whitish-tan, paler than rest of body

whorl. Anterior half of siphonal canal fight brown, darker

than body whorl. Protoconch pale whitish-tan. Interior

of aperture brown, becoming fighter in color toward la-

bral edge, and with prominent, widely-spaced dark

brown stripes that correspond to external primarv spiral

cords. Columellar side of siphonal canal white. Interior

of siphonal canal characteristically dark brown, several

shades of intensity darker than aperture interior or body

whorl exterior.

Page 34

THE NAUTILUS. Vol. Ill, No. 1

Type material: Holotype (CMNH 47339) length 67

mm, width 36 nini; ParatApe lot (7 specimens) (CMNH

37340) ranging from length 44 mm, width 27 mm, to

length 77 mm, width 44 mm. All t^pe material from the

t\-pe locality-.

Type locality: On beach, high tide hne, Barra de

Wawa, northern side of Wawa River mouth (Station 3),

Misldto Coast, Nicaragua.

Additional localities: Fragmentary- specimens were

also seen on beach, high tide line, southern side of Wawa

River mouth (Station 4), near Wawa Village, Misldto

Coast, Nicaragua.

Distribution: At present, collected only from the area

of the Wawa River mouth, Misldto Coast, Nicaragua, but

may extend from the Honduran Misldto Coast, near

Cabo Gracias a Dios, southward to Bluefields, Nicara-

gua. Discussions with local shrimpers have indicated that

Ficii.s lillai n.sp. is frequently taken in shrimp nets dur-

ing shallow (app. 15-20 m depth) trawls off the Cabo

Gracias a Dios coast, and that the local fishermen are

famiUar with this locally-common species. Broken, frag-

mentary specimens of Fictis villai were also collected at

Puerto Cabezas (Station 2), but the condition of these

was too poor to be part of the tspe lot.

Etymology: Named for Dr Jiiime Villa, Professor of

Biological Sciences, Florida Atlantic University, who as-

sisted me in collecting along the Wawa River area and

who was instrumental in securing my involvement in the

C.C.C. Misldto Coast research project.

Remarks: As presentK' imderstood, the Finis com-

itiuuis complex consists of three shallow water species

with disjunct distributions within the tropical and warm-

temperate western Atlantic; Ficiis communis Roding,

1798, (Figures 44, 4.5) a Carohnian Pro\ince, warm-tem-

perate species that ranges from Cape Hatteras, North

Carohna to Miami, Florida, and from Naples, Florida

around the Gulf of Mexico to Isla Contoy, Yucatan, Mex-

ico, and two tropical, Caribbean Prtnance species, Ficiis

villai n.sp., restricted to Miskitia, and Fiats lincloe Pe-

tuch, 1988 (Figures 40, 41), restricted to northern Co-

lombia and the Gulf of Venezuela area. These chstribu-

tions are shown in Figure 46.

Of the two pre\ioiisly-described species. Finis villai

is most similar, in l)oth shell size and thickness, to the

Carohnian Fcommunis. The new species differs from its

common and well-known congener in the following

ways; 1. shell shape — F villai is a more slender shell with

a higher, stepped spire; 2. shell sculpture — the new spe-

cies has a very- distinct sculpture pattern composed of

thick, prominent, high-rehef, primary spiral cords that

dominate the sculpture (Figure 35), while F. communis

has a much lower-relief pattern composed of weaker pri-

mary cords that are nearly equal in size to the secondary-

cords (Figure 45); 3. shell color — F. villai is a much

darker-colored shell, generally with dark longitudinal

flaminnlcs (Figure 33) and always with a prominent.

Figure 46. Distribvitions of the shallow water Ficus Roding,

1798 species of the western Atlantic. C = Fiais communis

Roding, 1798; V = Ficus villai new species; L = Fiats lindae

Petuch, 1988.

dark brown end to the siphonal canal, while F. communis

is a Ughter-colored shell, lacking both the longituchnal,

"zebra'"-t\pe flammules and the darklv-colored anterior

end.

The remiiining member of the complex, Ficus lindae

is the least sin-iilar to the new species, differing in having

a larger and much thinner, almost paperlike, shell. As in

the comparison with F. communis, F villai also chffers

from F. lindae in three distinct ways; L shell shape — the

new species has a proportionally more slender, less in-

flated shell with a higher, more elevated spire; 2. shell

sntlpture — F. villai exliibits a sculpture pattern of dom-

inant, high-relief primar\ spir;d cords with interstitial

smaller secondary and tertiary spiral cords, while F. lin-

dae (Figure 40) has much reduced, much lower-relief

priman spiral cords, a single reduced, secondar\- .spiral

cord, and no tertian- cords; 3. shell color — although both

species have similar browni shells \\-ith longitudinal

darker brown flammules, F. villai has a pale, whitish-tan

sulisutural area and a dark browii siphon;il end, both of

which are lacking in F lindae.

The new species also shows some moq-)hol()gical sini-

ilaritv to Ficus carolae Clench, 1945, a deep water (200

m depth) species from off the Florida Keys and Cam-

peche Banks of Yucatan, Mexico. This bath\-;il ficid

E. J. Petuch, 199S

Page 35

somewhat resembles F. villai. partitiilarlv in shell size,

shape, and proportions, hut clifters in having less-devel-

oped primarv spirit! cords and in haxing a paler-colored

shell that is conspicuously and characteristically marked

with scattered large, elongated, dark hrowii spots. Ficiis

carohw mav represent a deep water. continent;il slope-

dwelling offshoot of the shallow water Ficus communis

complex The t(.\'o other knovvm deep water western At-

lantic ficids. F atlautica Clench & Aguayo. 1940 (9(M) m

depth off central Brazil) and F htnielli Clench &

Aguayo, 1940 (2O(M00m depth from Cuba to Trinidad),

both have extremelv inflated, paper-thin shells, short,

stubb\' siphonal canals, and regiilarlv-arranged, darkly-

spotted color patterns. These tvvo species, together, dif-

fer greath' from the F. commiini'i complex, and appear

to represent a separate, distinct, unrelated group.

With the discovery of the Misldtian Ficiis villai, it is

now known that the western Atlantic houses sLx species

of Ficidae. Of these, three h\e in deep water (200—400

m), continental slope areas and three prefer shallow wa-

ter, neritic environments. The three shallow water spe-

cies can be separated as follows (+ = present, - =

absent):

Cancellarioidea

Cancellariidae

Cancellaria Lamarck, 1799

Canccllaria mediamciicana new species

(Figures 31, 32)

Description: Tvpical western Atlantic Cancellaria s.s..

of average length for genus (adults a\'. 35 mm), crudely

biconic, with inflated bocK' whorl. Shoulder angled, with

flattened subsutural area. Anterior canal short, open. In-

ner hp along columella with 2 colmnellar and 1 siphonal

folds. Posterionnost columeUar told largest, overlying si-

phonal fascicle. Body whorl and spire whorls with heav\',

coarse cancellate sculpture. Shell thick, structurally

strong, reinforced by heavy cancellate sculpture. Outline

of body whorl and spire whorls rounded. Edge ot shoul-

der acute, marked with carina-like spir;il cord. Flattenetl

subsutural areas give scalariform appearance to spire.

Shell surface coxered with evenK-spaced, intersecting

longitudinal and spiral cords, producing even cancellate

sculpture. Cords ol equal size and shape. Intersection of

cords producing low, rounded bead, gi\ing shell surface

pebbly appearance. Holotype with 33 longitudinal cords

and 14 spiral cords on bod\ whorl. Aperture proportion-

ally large, broad, semielliptical in shape. Inner edge of

labnun with 13 large, thin cords that extend into aperture.

Figure 47. Distributions ot the geographically-restricted spe-

cies i>( Cancellaria Lamarck, 1799 (aensu stricto) in the western

Ariantic. R= Cancellaria richardpetiti Petnch, 1987; A = Can-

cellaria adelae Pilsbr)-, 1940: M = Cancellaria medianKricana

n.sp.; P = Cancellaria pctnchi Harasevwch, Petit & Verhecken.

1992. The distnbution of the widespread Cancellaria reticulata

(Linnaeus, 1767) is not shown.

Labral cords cUsappear in interior of aperture. Entire col-

umeUar region overglazed with thin callus. Callus thicker

along anterior end, thinning toward posterior end. Both

columeUar folds and single siphonal fold with single large,

flattened keel. Posteriomiost and largest lold with %'ery

faint, poorly-developed secondary ridge along anterior

side. Protoconch paucispiral in fonn, composed of 2V2

low, inflated, glassy whorls. Protoconch of holotxpe devi-

ated from main shell axis by approximateh' 15° (Figure

48). Base shell color pale cream-white. Bod\- whorl with

3 wide, evenly-spaced, continuous dark reddish-brown

bands, one around anterior end, one just anterior to mid-

body and one aroimd shoulder Spire whorls with single

wide, redcUsh-browai band. Anteri(jr tip ol shell, anterior

can;il, and siphonal fasciole pale yellow-tan. Protoconch

pale cream-white. ColumeUar and siphonal folds white.

Type material: Holotype, (CMNH 47341) length 34

mm, width 20.5 mm; Paratypes, (CMNH 47342) (2

specimens), length 34 mm, width 20 mm and length 36

mm, width 22 mm. All t\pe material trom the tvpe lo-

cality.

Type locality: On beach, high tide hue, Barra de

Page 36

THE NAUTILUS, Vol. Ill, No. 1

Figure 48. Detail ot protoconchs of CannUaiia species. (Left)

Cnnrrllaria reticulata (Linnaeus, 1767), drawii from specimen

from Sanibel Is., Florida (Figure 44). (Right) Cancellaria me-

diamericana n.sp., drawn from holotvpe (Figures 31, .32). Note

proportionally larger size and flexure away from shell axis of

C. mediamericana.

Wawa, northern side of Wawa River mouth (Station 3),

Misldto Coast, Nicaragua.

Distribution: At present, known only from the area

of the Wawa River mouth, Misldto Coast, Nicaragua, but

may extend from the Honduran Misldto Coast, near

Cabo Gracias a Dios, southward to Bluefields, Nicara-

gua. A specimen in tlie private collection of Mr. Leonard

Hill, Miami, Florida, reportedlv taken by commercial

shrimpers from off Cabo Gracias a Dios, supports a wid-

er range for the new species.

Etymology: "mediamericana" , "Central American," in

reference to the confined range of the new species along

the eastern coast of Central America.

Remarks: Cancellaria mediamericana n.sp. is the

newest member of a characteristic western Atlantic spe-

cies comple.x centered around the widespread Cancel-

laria (s.s.) reticulata (Linnaeus, 1767) (Figures 36). That

well-known and common species ranges from North

Carohna, throughout the Gulf of Mexico, along the An-

tilles Arc, and southward to northern Brazil. Besides C.

reticulata and C. mediamericana. the complex also in-

cludes three other species with restricted geographical

ranges (Figure 47). These are: Cancellaria richardpctiti

Petuch, 1987 (100-200m depth off western Florida),

Cancellaria adelac Pilsbi'v, 1940 (carbonate sand areas,

1-10 m depth, Florida Keys), and Cancellaria petuchi

Harasewych, Petit & Verheeken, 1992 (central and

southern Brazil) (Figure 37). Of the members of the

complex, C. mediamericana is most similar to the wide-

spread C. reticulata, but differs in the following ways; 1.

shell shape — the new species has a more inflated shell

with a wider aperture; 2. columella — C. mediamericana

has only two columellar folds, with the large posterior-

most fold being flattened and keel-like, while C. reti-

culata has three folds with the large centr;il fold being

characteristically bifid in form; 3. protoconch — the pro-

toconch of the new species is proportionallv large, in-

flated, and bulbous with a slight flexure awav from the

m;un shell axis, while that of C'. reticulata is proportion-

ally much smaller (Figure 48), tightly cylindrical in form,

composed of three whorls, and is aligned with the m;iin

shell axis; 4. color — in the new species, the shell is

marked with three solid color bands, while in C reti-

culata, the color bands are discontinuous, broken into a

series of separate, large, rectangular maculations.

Cancellaria mediamericana is also simflar to the Bra-

zilian C. petuchi Harasewych, Petit & Verheeken, 1992

(Figure 37) but differs in the following ways; 1. shell

shape — ;ilthough both species have widely inflated body

whorls, C mediamericana has a shaq^ly-angletl shoulder

and a flattened, planar subsutural area, whfle C. petuchi

has a distinctly rounded shoulder; 2. shell sculpture —

while C. mediamericana has a sharji, strong reticulate

sculpture pattern that persists onto the bod\' whorl, C.

petuchi has a reduced sculpture pattern of low, rounded

ribs that become obsolete on the body whorl. Both spe-

cies, although widely separated geographically, have sim-

ilar inflated, bulbous protoconchs, triple-banded contin-

uous color pattern, and have only two, flattened, keel-

hke columellar folds.

The other two species of the complex, C.adelae Pils-

bry, 1940 and C. richardpctiti Petuch, 1987, are both

more similar to C. reticulata than to C. mediamericana.

The deep water, bathval C. richardpctiti is a distinctly

elongated shell with a high, protracted spire. The Flor-

ida Kevs endemic C. adelac. although generally similar

to C reticidata. has reduced shell sculpture, producing

a smooth l)od\' whorl like that ot C. petuchi. Both spe-

cies, C. richardpctiti and C. adelae, however, have three

columellar folds and have bifid, large central folds. In

the last character, particularly, these two species much

more closely resemble reticulata than C. mediameri-

cana. For a comparison of the sheU morphologies of C.

reticulata, C. adelae, and C. petuchi, see Harasewych,

Petit & Verheeken (1992). The widespread Caribbean,

Miskitian, and Brazihan Cancellaria species can be sep-

arated as follows ( + = present. — = absent):

1. inflateil protoconch

2. solid color bands

3. columellar callus

4. 3 cohunellar folds

5. 2 columellar folds

6. bifid central told

7. angled shoulder

+

+

+

-I-

Conoidea

Conidae

Conus Linnaeus, 1758

Lcptoconus Swiiinson, 1840

Conus (Lcptoconus) pa.'ichalli new species

(Figures 2, 3)

Description: Conus of smaller-than-average length

for subgenus Lcptoconus (adults av. 24 mm), with stock"s

body whorl, wide shoulder, and proportionally low spire.

Shoulder shaiply-angled, subcarinated. Spire whorls flat-

tened, noncanaliculate. Operculum unknown. General

shell form subpvriform, with con\ex sides on body

E. J. Petuch, 1998

Page 37

whorl. Body whorl tapers ahniptly anteriorly, pinching-

off noticeablv to produce narrow siphonal area. Spire

whorls slightly raised above suture hne, giving stepped

appearance to overall spire. Body whorl smooth and pol-

ished, marked with numerous very faint longitudin;il

gro\\'th lines. Spire whorls marked with ver\ nimierous,

closelv-spaced crescent-shapeil threads. Anterior one-

third ot body whorl encircled with 9-11 evenly-spaced,

deeply-impressed spiral sulci. Aperture proportionally

narrow, widening shghtly on anterior one-hali of shell.

Protoconch unknowii, missing on holot\pe and t\pe lot.

Based upon proportions ot broken early whorls, proto-

conch probably large and bulbous. Base color white or

pale yellowish-white, overlaid with 8-10 evenly-spaced

spiral bands of large orange dots and blotches. In some

cases, such as in holotvjie (Figures 2, 3), dots and rect-

angular blotches coalesce to produce large, widely-

spaced orange longitudinal flammules arranged in "ze-

bra" pattern. Clear white band, devoid of markings, ex-

cept for few small spots, present just anteriorward of

midbody hue. Clear band separates dots and longituchnal

flammules into two wide sections, one e.xtending from

shoulder to below midbody Une, and a narrower one

extending around anterior end of shell. Anterior tip of

shell very pale pinkish-lavender. Spire whorls white,

marked with widely spaced, orange, crescent-shaped

flammules. Early whorls (and presumed protoconch)

pale orange. Interior of aperture white. Based on small

patch still atUiering to outer hp of holotvpe (later re-

moved by cleaning in sodium hxpochlorite), periostra-

cum is thick, heaw, brown in color, marked with \'eiT

fine, longitudinal striae.

Type material: Holotype, ( CMNH 47344) length 26

mm, width 14 mm; Paratopes (7 specimens), (CMNH

47345) ranging from length 20 mm, width II mm, to

length 27 mm, width 14.5 mm. All tvpe material from

the type locality.

Type locality: On beach flat, low tide, southern side

of Bragman's Bluff (Station 1), appro.ximately 2.5 km

north of Puerto Cabezas, Miskito Coast, Nicaragua.

Distribution: At present, knowai onK' from the beach

areas near Bragman's Bluff and Puerto Cabezas, Nica-

ragua, but mav e.xtend southward to Bluefields, Nicara-

gua.

Etymology: Named for Dr Norman Paschall of Lar-

go, Florida, who assisted me in coUecting in the Brag-

man's Bluff area.

Remarks: Of the faiowai Caribbean small Lcptocouus

species. Conns pascholli is most similar to Conns por-

tobelocnsis Petuch, 1990 from the San Bias Islands re-

gion of Panama. The new species differs from its south-

em congener in the following ways; 1. shell shape — C.

paschalli is a smaller (av. 24 mm) species than C. por-

toheloensis, with a stockier, proportionally shorter, and

more pyriform body whorl; the shoulder of C. pascholli

is sharply-angled, almost carinated, while C. poiiohel-

ocnsis has a more rounded shoulder edge with only the

faintest hint of a carina; 2. spire — the spire of of C. pas-

challi. although low like that of C. porfobelocnsi.i, is dis-

tinctly stepped, with the spire whorls being shghtly

raised above each subsequent suture line; the spire of

the Panamanian species has depressed sutures, produc-

ing a graduallv-sloping spire outhne instead of a scalar-

iform one; the spire whorls are planar on the new spe-

cies, whfle those of C. poiiobeloemis are sloping and

faintly canaliculate; 3. shell sculpt n re — C. paschalli has

a smooth shell with faint longitutlinal striae, while the

shell of C poiiobeloensis is chstinctly sculptured with

fine spiral threads and striae; the anterior third to die

bodv whorl of C. paschalli is encircled with 9-11 large,

deeplv-incised sulci, while the anterior third of C por-

tobcloensis is encircled by at least 20 thin, faint raised

cords and an equal number of shallow, thin sulci; 4, col-

or— while both species have a color pattern composed

of spiral hues of dots and large coalescing longitudinal

flammules, C. paschalli has fewer rows of dots and the

indi\idual dots are proportionally larger; the anterior tip

of C. portobcloensis is pale yellow-orange while that of

C paschalli is pale pinkish-lavender 5. habitat — judging

from the tvpe localitv, the new species prefers muddy

sand locahties in shallow muddv water areas along the

river effluent zone of the Miskito Coast of Nicaragua;

Conns portobeloensis, on the other hand, lives in an off-

shore (30 m depth) carbonate substrate — clean water

emironment (see Petuch, 1990:68) off the Colon and

San Bias coasts of Panama. Neither species, nor anything

resembhng them, has been collected along the interven-

ing clean sandy coast of Costa Rica (Houbrick, 1968),

demonstrating geographical and ecological separation.

Bivalvia

Pteriomorpha

Arcoida

Arcidae

l^oetia Gray, 1857

Noctia (s.s.) lindae new species

(Figures 9, 10)

Description: Shell small for genus (adults av. 25 mm),

equivalve elongated, subtrigonal in shape. Posterior end

onh' slightly expanded, truncated, sharplv angled, taper-

ing to sharp point. Angled posterior truncation bordered

by large, prominent posterior ridge that runs from um-

bones to posterior tip. Shallow medial depression pres-

ent on posterior end above posterior ridge. Umbones

placed extremely forward, in anterior one-third of sheU.

Anterior end not expanded, rounded, without angled

ridge. Shell surface with 30-32 evenly-spaced, low, wide,

rounded primarv radial ribs. Thin, rounded secondary

radial rib present between each p;ur of v\ide primaiy

ribs. SecondaiT rib bisected bv fine incised line. Primary

and secondary radial ribs covered by and intersecting

with very fine, closely-packed concentric growi:h fines,

giving ribs sfightly roughened appearance. Intermittent

Page 38

THE NAUTILUS, Vol. Ill, No. 1

large concentric growth Une.s present, corre.sponcling to

periodic ce.ssation.s in shell growth. Shell margin and in-

ner margin marked with 30-32 shaq), well-developed

crenulations that correspond to thin secondary ribs.

Beaks large, prominent, angular, projecting above hinge

Une. Umbones well-de\eloped, recuned. Beaks and um-

bones strongK' opisthogvrate. Hinge hne straight, v\ith

coarse taxodont dentition. Posterior teeth larger than an-

terior teeth. Hinge Ugament elongated, broad, with

trans\'erse hgamentarv grooves at right angles to hinge

hne. LigamentaA' grooves 7-8 in number, placed ante-

riorly to umbones. Anterior muscle scar proportionally

small, oval, at extreme anterior end. Posterior muscle

scar proportionally large, oval, formed at base ot long,

narrow, low flange. Posterior flange extends upward into

umbonal interior Palhal line weakly developed, straight,

following edge of shell margin just above marginal cren-

ulations. SheU pale-to-bright lemon-yellow or cream-or-

ange, with posterior end and area above posteroventral

angle being darker vellow-orange. Umbones and beaks

brightlv colored, generally dark reddish-orange or bright

cherrv red. Shell interior pale yellow or yellowish-white,

with some specimens (holohpe and 6 paratvpes) having

darker veUow-orange stain deeper within. Occasional

specimens (such as one left valve paratype) with dark

red-orange stain that extends from umbonal interior to

posterior margin. Periostracum (Partially preserved on

two left valve paratypes; removed from holotype pair by

soaking in sochum hypochlorite solution) thick, black,

coarsely lameUose, marked with thin concentric striae.

Periostracum eroded off of beaks of all specimens ex-

amined.

Type material: Holotype, (CMNH 47367) length 25.5

mm, width 15 mm; Paratypes, (12 single valves, CMNH

47343), ranging from length 20 mm, width 13 mm to

length 29 mm, width 18 mm. All type material from the

type locahty.

Type locality: On lower beach flat at low tide, south-

ern side of Bragman's Bluff (Station 1), approximately

2.5 km north of Puerto Cabezas, Miskito Coast, Nica-

ragua.

Distribution: At present, known only from the beach

area near Bragman's Bluff, Puerto Cabezas, Nicaragua,

but mav possiblv be found southward along the Miskito

Coast.

Etymology: Named for mv wife, Linda Joyce Petuch.

Remarks: Noctia linilac is the first member of its ge-

nus (s.s.) to be recorded from the Caribbean region, and

only the second species of Noetia s.s. to be found in the

western Atlantic. The other species, Noetia bisulcata

(Lamarck, 1819) occurs along eastern South America,

from Surinam to Uruguay (range and illustration in Bios,

1975: fig. 949). No other Noetia s.s. is known to occur

in the area behveen N. lindae and N. bisulcata. The larg-

er Noetia ponclerosa (Say, 1822) from the Carolinian

Province belongs to (and is the tvpe of) the subgenus

Eontia MacNeil, 1938, and represents a dilierent and

separate Noetia lineage. The Panamic Noetia reversa

(Sowerby, 1833) and Noetia olssoni Sheldon and Maury,

1922 are verv' similar to N. lindae, and the new Carib-

bean species appears to be intermediate in sheU form

between its two Pacific relatives; having the more elon-

gated shape of .V. olssoni and the strong, prominent pos-

terior ridge and high beaks of .V. reversa.

In general sheU shape and size, Noetia lindae is most

simflar to the South American N. bisulcata, but differs

in the following wavs; 1. .shell .shape — N. lindae is a much

more elongated species with a projecting, distinctly

pointed posterior end, while N. bisulcata has a more

ovate shell shape with a wide, blunt posterior end; the

beaks of the new species are located much more ante-

riorly, within the anterior one-third of the shell, whereas

the beaks of N. bisulcata are more medially located, al-

most at the midbody line; the dorsid posterior area im-

mediatelv above the posterior ridge is flattened in N.

lindae. while in N. bisulcata the posterior dorsal area is

riiised and laterallv flattened; 2. color — N. lindae is a

much more colorful shell than its southern relative, often

exhibiting a bright vellow or yeUow-orange sheU with red

lunbones, while N. bisulcata has a more subdued sheU

c(jlor, being white or pale vellow- white with light orange

umbones.

Pterioida

Pectinoidea

Plicatuhdae

Plicatula Lamarck. 1801

Plicatula miskito new species

(Figures 16, 17, 18, 9)

Description: Shell small for genus (holotype 13 mm),

o\atelv subtrigonal in shape, flattened, with right valve

attached to substrate. Auricles present but poorly-de-

veloped, with posterior auricle larger and wider than an-

terior auricle. Interior porcellaneous. Hinge line straight.

Exterior of free (left) valve with ver\' fine, raised con-

centric lines that become stronger and more lamellose

toward shell margin. Area near beaks decorated with 18

very fine beaded radial ribs. Badi;il ribs flatten and dis-

appear halfwav between beak and shell margin. Shell

margin hiintlv crenulate. Beak poorly-developed, almost

planar, with small, faint shar|-)ly-pointed umbo projecting

only shghtlv abo\e hinge Une. Entire hinge stmcture

proporrionallv large, typicalh' spondyloid, with narrow,

small, elongated chrondrophore. Chondrophore pit of

upper (left) valve flanked hv two cniral ridges, each with

single, smooth ball-Upe teeth. Chondrophore pit of bot-

tom (right) \al\e flanked by hvo elongated sockets and

hvo small elongated teeth. Single adductor scar posterior

to center of shell, almost circular in shape. Pallial line

deeply incised, located subcentralK, following shell mar-

gin. Exterior of free (left) valve white with numerous

fine, radiating, p;ile reddish-brown fines and large, scat-

tered, dark retldish-brown spots. Shell margin marked

with large, prominent, equallv-spaced, dark redilish-

E. J. Petuch, 1998

Page 39

brouii streaks. Suhiiiiilional area stiunetl pale tan. Unilio

reddish-tan. Attached (right) \alve pure white. Interior

of left val\e white with huge piile tan stain, extending

from adductor scar to unibonal area. Margin of shell

interior of left \al\e marked with prominent, equallv-

spaced dark reddish-hrowii streaks, corresponding to

those seen on shell exterior. Ball-and-socket teeth of

both \'al\es pale tan.

Type material: Holotype (pair) (CMNH 47366)

length 13 mm, width 5 mm.

Type localifrv: On lower beach flat at low tide, in shell

mbble bed, southern side of Bragman's Bluff (Station 1),

appro.ximatelv 2.5 km north of Puerto Cabezas, Miskito

Coast, Nicaragua. Holotype found attached to interior of

Noctia liiulac n.sp. vahe (Figures 18, 19).

Distribution: At present known only from shell rub-

ble beds at Bragman's Bluff, Puerto Cabezas, Nicaragua.

Etjinology: Named for the Miskito Indians, as a noun

in apposition.

Remarks: Pikritula miskito n.sp. is onlv the second-

knowai representati\'e of its genus in the vs'esteni Atlan-

tic, and is the first Plicatula to be collected on the Mis-

kito Coast. The other Atlantic member, Plicatula gibbosa

Lamarck, 1801, which ranges from North Carohna to

Brazil, is an abundant but verv tlifferent-appearing spe-

cies and does not seem to be closeh' related. PUcatula

miskito differs from its widespread and well-knowoi con-

gener in the following ways; 1. size — P. gibbosa is a much

larger species, averaging 25 mm in length (Abbott, 1974:

450), while P. miskito is a much smaller animal, with the

holotvpe being onl\' 13 mm in length; 2. shape ami

sculpture — the common P.gibbosa is a \erv rugosely-

sculptured species with 5-7 prominent, high, raised ph-

cations diat gi\e the shell a strongk' ribbed appearance

(hence the common name, the "kitten's paw "), while the

shell of P. miskito lacks the raised pUcae having, instead,

fine beaded radial riblets near the umbo; the shell mar-

gin and commissure of P. miskito are relativelv straight

and flattened while those of P. gibbosa are strongK' un-

dulate.

Morphologicidlv, the new species is most similar to

Plicatula penicillata Carpenter, 1857 from the Panamic

Pro\ince, and its presence in the Miskitian malacofauna

further underscores the Pacific faunal affinities of the

Nicaraguan Caribbean coastal region. The two species

are strildnglv similar, parhcularlv in size, shape, and color

pattern. Both species exhibit the chstincti\e and char-

acteristic large dark streaks along the shell margin and

both have the large tan stain within the shell interior.

Plicatula miskito differs from P. penicillata primarily in

the surface sculpture of the tree val\-e; while the new-

species is relativelv smooth, with onl\' fiiint concentric

laminae and rachal riblets, its Panamic congener has a

more crenulate, rougher, surface, often decorated with

small ribs and spines. Otherwise, the two species are

extremely close and they form an impressixe example of

a pre\iouslv-unkno\\ii Panamic-Caribbean sibhng spe-

cies pair.

Heterodonta

Veneroida

Mactroidea

Mactridae

Mactra Linnaeus, 1767

Mactra inceii new species

(Figures 7, 8, 11)

Description: Shell topically mactrid in form, thin,

fragile, with convex, mflated valves, of average size for

genus. Shell shape generally oval-subtrigonal, with high,

pointed umbones and shghtly concave posterior-dorsal

area. Faint, low, rounded keel borders posterior-dorsal

area and posterior margin of umbones. Posterior end

shghtly pointed, anterior end rounded. Beaks placed

shghtly anterior to sheU midhne (based on total sheU

length). Lunular region wide, deeply impressed. Entire

shell exterior covered with \ery fine, closely-packed con-

centric grov\i:h fines, giving shell silk-y texture. Growth

lines become coarser toward shell margin. Beaks large,

prominent, trigonal in shape, projecting above hinge

hue. Umbones highly recurved, strongly prosogvTate.

Resihfer proportionally large, trigonal in shape, with

apex of triangle bent anteriorally Resihfer set in large,

cup-shaped chondrophone that overhangs umbonal re-

gion, producing deep subinnbon;il cavitv,-. Edges of re-

silifer-chondrophore complex bounded by thin, raised,

shghtlv' concave shelly lamina. Carchnal teeth prominent,

with posterior flange overhanging resihfer. Lateral teeth

thin, elongated, bladelike, with anterior lateral tooth hav-

ing projecting, trigonal flange. Anterior adductor scar

near extreme anterior margin of shell, thin, elongated in

shape. Posterior scar nearly circular in shape, located

near extreme posterior shell margin. Palhal hue promi-

nent, sinusoidal in form, with long, deep, sublinguiform

piilhal sinus. P;illial sinus extends horizontallv well past

shell midhne (see Figs. 8, 11). SheU exterior and interior

uniform pale cream-white. Periostracum (Present only

as fragmentary remnants along margin of holotype valve)

pale veUow-tan, thin, translucent, with silkv' texture.

T)pe material: Holotype, (single right valve) (CMNH

47.338). length 35.5 m, width 41 mm; Figured paratype,

(single right valve, CMNH 47346), length 24 mm, width

27 mm; Parat\pes, (3 single valves, CMNH 47.350),

ranging from length 25 mm, width 28 mm to length 33

mm, width 36 mm. All type material from the tvpe lo-

c;dit\-.

Type locality: On lower beach flat at low tide, south-

em side of Bragman's Bluff (Station 1), approximately

2.5 km north of Puerto Cabezas, Miskito Coast, Nica-

ragua.

Distribution: At present, kiiowir only from the beach

area near Bragman's Bluff, Puerto Cabezas, Nicaragua,

but may possibly be found southward along the entire

Miskito Coast.

Page 40

THE NAUTILUS, Vol. Ill, No. 1

Etymology: Named tor Dr. Jaime Incer, director of

IRENA, Managua, Nicaragua, in thanks for his assi.s-

tance and support of my research while I worked along

the Misldto Coast.

Remarks: Mactra inccri is the first species of Mactra

(s.s.) to be reported from the Caribbean Basin. The only

other known western Atlantic member of the genus is

Mactra ihciingi (Dall, 1897), which ranges from Vene-

zuela (Weisbord, 1964:381-382) southward to Santa Ca-

tarina, Brazil (Rios, 1975:236) and, prior to the discovery

of the Miskitian species, the genus Mactra (s.s.) was con-

spicuously absent from northern South America, eastern

Central America, and the West Indian Arc. Interestingly

enough, the subgenus Mactrotoma (typified by Mactra

fragilis Gmelin, 1791), which is the common Caribbean

representative of the Mactridae, was not collected along

the Miskito Coast but was replaced by the nominate ge-

nus.

Although similar to the South American-Brazilian

Mactra iltcringi, M. inceri can be readily distinguished

by the following characters; 1. shell shape — The new

species has a less-elongated, more rounded and truncat-

ed anterior and; because of this truncated appearance,

the umbones of M. inceri are located slightly anterior to

the midbody line, while those of M iheringi are almost

exactly centrally located, directly on the midbod)'; 2.

form of the hinulc — in M. inceri, the lunule is distinctly

depressed, producing a noticeable concavity along the

anterior-dorsal area immediately anterior to the umbo-

nes; in M.iheringi, the lunular area is simply flattened,

producing a straight, sloping surface; 3. paUial sinus — in

M. ihcringi, the sublinguiiorm pallial sinus extends hor-

izontally to, or a Uttle beyond, the midbody Une (Dall,

1897; Weisbord, 1964:381), while in M inccri, the palhal

sinus extends well beyond the midbody Une (Figures 8,

11) occupying fully two-thirds of the shell interior; this

enlarged palhal sinus demonstrates that M. inceri has

longer siphons than does its southern congener and lives

more deeply buried in the substrate.

Micronuuira Dall, 1894

Microuiactra miskito new .species

(Figures 12, 13)

Description: Shell of average size for genus (av. 27

mm), elongated, thin, fragile, with beaks centrally locat-

ed. Anterior dorsal margin of shell, immediately anterior

to beaks, nearly stndght, with only slightest hint of con-

cavity. Anterior end rounded, posterior end pointed but

angularly truncated. Low but sharp angular ridge pres-

ent along entire posterior dorsal edge of both valves.

Beaks and umbonal area of shell exterior characteristi-

cally sculpted with 6-15 large, prominent, undulating

waves. Surface of exterior smooth, shiny, covered with

very fine, closely-packed concentric growth lines, giving

shell faintly silky appearance. Large umbonal undula-

tions also covered by fine concentric growth lines. Very-

fine, thin, evenly-spaced radial lines present on anterior

one-half of shell. Beaks large, prominent, projecting well

above hinge line, roughly trigonal in .shape. Umbones

highly recurved, strongly prosogyrate. Resilifer trigonal

in shape, with apex tipped anteriorly, set in small cup-

shaped chondrophone. Cardinal teeth distinctly "V"-

shaped. Lateral teeth elongated, bladehke. Anterior ad-

ductor scar near extreme anterior margin of shell, oval

in shape. Posterior scar of approximately equal size and

shape, located near extreme posterior shell margin. Pal-

hal fine sinu.soidal in form, with short, rounded, wide,

pallial sinus. Palhal sinus extends horizontally only about

one-third of shell interior Shell exterior and interior uni-

form translucent white. Periostracum thin, smooth and

silky, adherent, pale straw colored, generally missing on

umbones.

Type material: Holotype, (single right valve) (CMNH

47364), length 30 mm, width 19 mm; Paratypes, (5 sin-

gle viilves, CMNH 47352), ranging from length 25 mm,

width 17 mm to length 28 mm. width 19 mm. All type

material from the type locahty.

Type locality: On lower beach flat at low tide, south-

em side of Bragman's Bluff (Station 1), approximately

2.5 km north of Puerto Cabezas, Miskito Coast, Nica-

ragua.

Distribution: At present, known only from the beach

area near Bragman's Bluff, Puerto Cabezas, Nicaragua,

but may possibly be found southward along the entire

Miskito Coast.

Etymology: Named for the Miskito Indians.

Remarks: The finding of a Caribbean Micromactra

species was one of the most interesting and important

chscoveries to be uncovered during my research in Nic-

aragua. Until now, the genus was kiiown from the west-

em Atlantic by only a single cooler-water species, the

Brazilian Province Micromactra janciroensis (E.A.

Smith, 1915), which ranges from Espirito Santo State,

Brazil to Umguay (Rios, 1975:236). On the other hand,

the genus is well-represented in the Eastern Pacific, with

at least five species occurring in the Panamic Province

alone (Keen, 1971:204).

Of the known Micromactra species, both Eastern Pa-

cific and western Atlantic, M. miskito most closely re-

sembles the Pacific M califoniica (Conrad, 1837) which

ranges from the Puget Sound southward to Costa Rica,

and they form another example of a previously-unk-nown

Panamic-Caribbean sibling species pair. The new species

differs from its Pacific an;dogue in being slightly more

elongated \\ith a noticeably more tnmcated posterior

end, in having more prominent, more highly ele\'ated

umbones, and in having a shghtly more concave lunular

area. Otherwise, the two species are verv similar, es-

pecially in having virtually identical patterns of umbonal

undulations.

From the only other Atlantic Micromactra species, M.

janciroensis, M. miskito can be separated by the follow-

ing characters; 1. stiell shape — the new species has a

E. J. Petuch, 1998

Pas^e 41

much more elongated, slender shell than dot-s M jaitci-

rocnsis, and the posterior end of A/, lindac is much more

pointed and sharply-angled than the blunted, more

rounded posterior of its southern congener; 2. undulat-

ing sculpture — the imdulating, wavy sculpture pattern

(the main characteristic of the genus Micwmactra) is

confined to the umboniil region of M. miskito, while on

M.jancirocnsis it extends onto the main shell body, often

covering as much as two-thirds of the shell surface.

Petricohdae

Petricohnae

Petricola Lamarck, 1801

Petricolaria Stoliczka, 1870

Petricola (Petricolaria) donnac new species

(Figures 24, 25)

Description: Shell thin, fragile, small for subgenus

(holot)pe length 28 mm), elongated, subcylindrical in

shape, with dorsal and ventral margins roughly parallel.

Umbones rounded, elevated abo\e hinge line, placed far

forward near anterior end. Small flattened lunule pres-

ent just anterior t(; umbones. Anterior end pointed, pos-

terior end wider and distinctly rounded. Shell e.xterior

covered with numerous fine, closely-spaced radial ribs.

Radial ribs, especiallv those on anterior one-fourth of

shell, become coarser and farther apart toward anterior

end of shell. Radial ribs crossed bv numerous, evenly-

spaced concentric ribs. Intersections of concentric ribs

with large anterior radial ribs producing small spine-like

scales on radial ribs. Umbones rounded, recurxed,

strongly prosogyrate. Hinge fine straight, smooth. Hinge

vidthout lateral teeth. Cardinal teeth paired, tyjiicallv pe-

tricohd, with anterior tooth cun'ed and hookUke, and

with posterior tooth flattened and bifurcated. Pallial line

sinusoidal in form, with long, deep, pointed subfingui-

form pallial sinus. Pallial sinus extends horizontall)' to

shell midline. Shell exterior and interior pale cream-yel-

low, with posterior end stained pale orange. Pale orange

color may be staining from the red clay within which the

animal had bored, and not part (jf the actual shell color

Type material: Holotype (pair) (CMNH 47373),

length 28 mm, width 9 mm.

Type locality: On beach near clay cliffs, approximately

1 km north of main pier of Puerto Cabezas, Miskito

Coast, Nicaragua.

Distribution: At present, known only from the clay

chff and clay reef areas near Puerto Cabezas, Nicaragua.

Etymology: Named for Mrs. Donna ViUa, of Welling-

ton, Florida. Her husband. Dr. Jiiime Villa, collected the

unique holotype and wished to have the species named

for her.

Remarks: Prior to the discovery of Petricola {Petri-

colaria) donnae, the subgenus was known in the western

Atlantic from only a single wide-ranging species, P. (Pe-

tricolaria) pholadifonnis (Lamarck, 1818) (Figure 23).

Although only a single specimen of the new species has

been collected, the new Petricolaria is so different that

there can be no doubt that it represents a previously-

unknown and undescribed animal. The apparent rarity

of the new species is most probably an artifact of col-

lectibility. Judging from the abundance of bivalve bor-

ings in the clay reefs, the species is probably common,

but is deeply buried below the surface and is virtually

inaccessible. Rarely, specimens such as the holotype may

wash out of broken clay blocks during storms.

The new species can be readily separated from its only

western Atlantic congener in the foUowing ways; 1. shell

shape — Petricola donnac is a much shorter, stockier spe-

cies with a proportiouiilly wider, less elongated shefl; the

posterior end of F donnac is much v\ider and n(jticeably

more rounded than the more pointed posterior end of

P. pholadifonnis; the umbones of P. donnae are also pro-

portionalK' narrower and more acute than those of P.

pholadifonnis, and project farther above the hinge line;

2. shell sculpture — the posterior and central radial ribs

of P. donnae are much more numerous and better-de-

fined than those of P. pholadifonnis, while the anterior

radial ribs of P. donnac are not as large nor as heavilv

sculptured as those of P. pholadifonnis (readily seen in

a comparison of Figures 23 and 24).

Of the known American Petricolaria species, P. don-

nae is actually most similar moqjhologicidly to the Pan-

amic Pro\ince P. cognata C.B. Adams, 1852, especially

in overall shell shape and size. The Pacific P. cognata,

however, has much coarser and more heavily-sculptured

anterior radial ribs, much in the manner of P. pholadi-

fonnis. Even with this sculptural difference, P. donnae

is still very similar to P. cognata and can be considered

to be its Atlantic analogue, just as the Panamic P. par-

allcla Pilsbry and Lowe, 1932 is the Pacific analogue of

the Atlantic P pholadifonnis. Interestingly enough, the

normally-connnon Caribbean clay-dwelling P. pholadi-

fonnis has not been reported from the Miskito Coast,

where it appears to have been replaced by P. donnae.

BIOGEOGRAPHICAL IMPLICATIONS

Based upon the faunal evidence presented in this paper

and in pre\ious studies (Fluck, 1905-1906; Petuch,

1981,1987,1988), it is now known that the shorehne and

estuarine areas of eastern Central America, from ap-

proximately Cabo Camaron, Honduras to near Blue-

fields, Nicaragua, house a combined molluscan fauna

that bears onl\- a partial relationship to the siurounding

Caribbean Molluscan Province. Since this new faun;il

subdivision of the Caribbean Pnnince is centered on the

Miskito Coast of Honduras and Nicaragua, I here pro-

pose the name "Misldtia" for this newly-discovered bio-

geographical unit. At present, the biogeographical sub-

divisions of the Caribbean Province, particularly at the

subprovincial level, are still virtuallv unknowni, so 1 pre-

fer to use "Miskitia" as an inforniiil designation — simply

to emphasize the atypical Caribbean nature of the near-

shore molluscan fauna. As the area is better studied, it

Page 42

THE NAUTILUS, Vol. Ill, No. 1

may be proven that Miskitia actually represents a sepa-

rate and discrete niolKiscan suhprovince.

As presentK understood, the tropic;il Americas con-

tain two types of marine moUuscan faunas; a "Caribbe-

an"-tvpe fauna and a "Panamic"-t>pe fauna. A classic Ca-

ribbean fauna contains index genera such as Lithopoma,

Piipciifa. Snuimgdia. Ccucliiiti.s. Chicorctts. FascioJaria,

S.S., Dolicholatints, ConcUa, Cariholiva, Turbiuclla, Cor-

clih/ria. Lindapecten {"Aequipecten" muscosus complex),

Spcnglciio. and in some areas Valuta and Siphocypraca.

A classic Panamic fauna, on the other hand, contains

index genera such as Ancistromcsiis, Trochita, Jenneria,

Malea, Pseudozonaria, Muricanthtts. Northia, Agawnia,

Noetia (s.s.), and Micromactra. These faunal character-

izations were based on assessments of transisthmian as-

semblages (Caribbean and Eastern Pacific) and were re-

ported on by Olsson (1972), Petuch (1982), Radwin

(1969), Vermeij and Petuch (1986), and Woodring

(1966). From the results of these studies, it was gener-

ally accepted that the Caribliean and Panamic moiiuscan

faunas, with the exception of some analogue species

("cognates" of Radwin, 1969), were quite different-ap-

pearing, and that the Panamic fauna was much more

diverse and species-rich (Olsson in Olsson & Petit, 1964:

525). Subsequent works (Petuch, 1987;1988;1990), how-

ever have showm that the Caribbean area, particularly

northern South America and isolated islands, does har-

bor some prexiously-unrecognized "Panamic" ("paciphi-

les" of Woodring, 1973) genera such as Cotonopsis,

Tnincaria, Agaronia, Aphera, Kncfastia, and Cijclothtjca,

as well as supposedly-extinct genera such as Falsihpia,

Conomitra, Picioptijgina, Ccrithioclavo, and Parahorson-

ia. The Caribbean, then, can be seen to be far richer

and more complicated, faunisticallv, than was previously

thought.

With the present discovery of the new western Carib-

bean fauna reported here, the tropical Americas are now

known to contain three "Panamic"-type faunas; an At-

lantic component containing the Brazilian Province (as

defined by Petuch, 1988:166; Harasewych, Petit & Ver-

hecken, 1992) and the Miskitian area of the Caribbean

Province, and a Pacific component containing the classic

Panamic Province and offshore Cocos Island and Gala-

pagos Islands "Subprovinces" (possibly full provinces).

The biogeographical divisions that contain "Panamic" —

type faunas and that show close faunal affinities to each

other are shown in Figure 49, and as can be seen, Mis-

kitia represents a sh;illow water Panamic fauna that has

been left behind in the western Atlantic after the final

closure of the Central American Isthnms in the late Pfio-

cene. A short listing of shallow water and estuarine en-

demics and their living Panamic analogues (cognates)

demonstrates the Panamic- Eastern Pacific nature of the

Miskitian fauna;

Figure 49. Moiiuscan biogeographical areas of the tropical

Americas that sliow close fauna) affinities, particularK- in their

shoreline assemblages. P = Panamic Moiiuscan Pro\ince; M

= Miskitia (svibprovince of the Caribbean Moiiuscan Province

?); B = Brazilian Moiiuscan Province. All three share genera

such as Aoaronia, Noetia (s.s.), and Micromactra. With the

Panamic Province, Miskitia also shares species of the Plicatula

penicillata complex, Pacipecten. and the Petricola (Petricolaria)

cognatn complex.

Agaronia hilli

Canccllaria iiudiainericana

Coiuts pa.sclmlli

Noetia lindae

Pacipecten leticophaeus

Plicatula miskito

Micromactra miskito

Polymesofla placans

Petricola donnae

A. propatula

C obesa

C. regitlaris

N. olssoni

P. tumbezrnsis

P. penicillata

M. califoniica

P. nicaragiiaua

P comata

Miskitia Restricted

Stroinhiis piigilis nicaraguen-

.vi.v

Ficuti villai

Panamic (Pacific)

.S gracilior

F ventricosa

A close relationship to the other Atlantic "Panamic"

fauna, the Brazilian Province, is also readily demonstrat-

ed by the h)llovving Miskitian endemics and their Bra-

zilian an;dogues;

Miskitia Roslrictcd Brazilian

Stroinbiis pugilis nicaraguen- S. pugilis worki

si.s

Agaronia hilli A. travassosi

Cancellaiia mediameriruna C. petuchi

Conns pasclialli C. tostesi

Soelia lindae N. bisiileala

Mactra inceri M. iheringi

Micromactra miskito M janeiroensis

All three "Panamic" faunas can be seen to siiare the

genera Aganmia. Noctio (s.s.), and Microimirtra. Inter-

estingly enough, all three areas also have their own

ineml)er of a Stromhus (s.s.) species complex (S. pugilis

nicaragiicusis Fluck, 1905, .S ptigili.s worki Petuch,

E. J. Petuth, 1998

Page 43

Figure 50. Configurations of the American tropical seas dur-

ing the early Piacenzian Pliocene, showing the cUstributions ot

the subprovinces of the Gatunian Molluscan Provance (shaded

areas) (taken from Petuch, 1988). 1 = Iniperialian Subpro\-

ince, 2 = Limonian Snbpro\-ince, 3 = Esmeraldan Subprov-

ince, 4 = Agueguexitean Subpro\ince. 5 = Gnraban Svibprox-

ince, 6 = Carriacouan Snbpro\ince, 7 = Pimtaga\ikinian Sub-

pro\ince, 8 = Piraban Subprovince, 9 = Camachoan Subprox-

ince. C = region of the Caloosahatchian Molluscan Proxmce

(with four subpro\inces; see Petuch. 1997); PM = region ot

the Protomagellanic Molluscan Province.

1993, and S. gracilior Sowerby, 1825), a CanccWana (s.s.)

species complex (C. mediamciicana n.sp., C. pctiichi

Harasewvch, Petit & Verhecken, 1992, and C. ohcsa

Sowerby, 1832), and Contis (Leptocoinis) species com-

plex (C. paschaUi n.sp., C. tostcsi Petuch, 1986, and C.

regularis Sowerby, 1833).

The shared genera and species complexes of the geo-

graphicallv-separated Panamic and Brazilian Proxinces

and Miskitia reflect a common origin in the Gatunian

Province (Petuch, 1982) (Figure 50) during the late Plio-

cene, prior to the formation of the Panamanian Isthmus

and Amazon Ri\er barriers. Since the final closing of

Panama (VVhitmore & Stewart, 1965:185), the Panamic

Province has retained a nearly intact Gatunian appear-

ance (Vermeij & Petuch, 1986) while the Caribbean

Province has become a mLxture of Gatunian sunivors

and immigrants from the northern Caloos;ihatchian

Province (Petuch, 1988:115-116; 1997). Prior to the dis-

covery of Miskitia, the onlv Atlantic area that retained

any semblance of the Gatunian Province (Piraban Sub-

province) was the Brazihan Province, which shares with

the post-closure Panamic Province additional genera

such as Malea, Northia, and Plcuroploca (s.s). Miskitia,

then, represents yet another pocket of shallow water Ga-

tunian survivors within the western Atlantic, and one

that is contiguous with typical Caribbean assemblages to

the north and south

The ancestr;il fauna of Miskitia is found within the

fossil beds of the Limonian Subprovince of the Gatunian

Prfnince (Figiue 50), a Pliocene biogeographiciil area

that spanned both sides of the then-open Central Amer-

ican Isthmus. A survey of the eastern Costa Rican Limon

and Gatun Formations and vvesteni Costa Rican Ar-

muelles and Charco Azul Formations (hsted by Olsson,

1922; 1942), which all contain tvpical Limonian Gatu-

nian elements, will readilv demonstrate the Gatunian an-

cestrv of the Recent Misldtian fauna. As in the cases of

the Recent Brazihan and Panamic Provinces, Miskitia

also contains analogues with the older Limonian Sub-

province. These inchide;

Miskitian Restricted Limonian

Stroinhwi pii^ihs iiicaragtu'n-

sis

Ficus villai

Agaronia hilli

CanccUaiia iiwdiauwricana

Conus pa.schalli

Noetia lindae

Pacipecten leucophaeus

Micntmacim iiiiskito

S- puffloidcs

F. carbasea

A. costaricensis

C biiiretti

C costaricensis

N. subreversa

P. costaricensis

M. sp. (Arniuelles Form.. Ols-

son, 1942)

These Limonian paleoanalogues appear to be the direct

ancestors of the Miskitian species.

The survival of a "Panamic", Gatunian-derived Mis-

kitian fauna in the Caribbean region is also probably due

to the retention of a Panamic-tvpe coastal habitat within

the western Atlantic. Both the Pacific and Atlantic coasts

of Honduras and Nicaragua are Upified bv sificiclastic,

muddy, terrigenous run-off, high productivity' environ-

ments and both coasts have large mangrove forests with

extensive estuarine areas. Although this tvpe of coasthne

is tvpical of the Panamic Province south ot Mexico, it

occurs onlv sporadicaliv within the Caribbean Basin,

where carbonate environments predominate. To the

north of Miskitia, the extensive coral reef systems of the

Great Barrier Reef of Belize and the Bay Islands of

Honduras predominate. To the south, the open, clean

sand coasts and coral reef systems of Costa Rica, Pana-

ma, and the San Bias Islands (the Blasian Subregion of

Petuch, 1990) represent a mirror image of the carbonate

environments of the Bav Islands. The Miskitian area,

environmentally and sedimentologicaliv, represents an

"island" of muddy nearshore habitats wedged between

these two extensive coral-dominated subregions. This sil-

iciclastic-dominated environment mav have been pres-

ent since before the closure of Panama, allowing the

older Gatunian elements to survive within the Caribbean

Sea even though their congeners died-out in the other

Atlantic Gatunian subprovinces due to lowered produc-

tivity- and increased carbonate build-up (Vermeij & Pe-

tuch, 1986).

The discoveiy of Miskitia further underscores the fau-

na! and temporal heterogeneity of the Caribbean Mol-

luscan Province and, hopefullv, will focus the attention

Page 44

THE NAUTILUS, Vol. Ill, No. 1

of biogeographers on one of the most biotically compli-

cated, but vet une.vjilored, areas of the entire Atlantic

Ocean.

ACKNOWLEDGMENTS

For assistance in the collection of specimens from the

Wdwd Ri\er and Puerto Cabezas areas, I thank Dr. Jaime

D. Villa, Department of Biok)gical Sciences. Florida At-

lantic University', Boca Raton, Florida, Dr. Norman Pas-

chall. Largo. Florida, and Sr. Raimundo Solorzano, Ma-

nagua, Nicaragua. I also thank the Caribbean Conser-

vation Corporation (CCC), Giunesville, Florida, and Dr

Charles S. Luthin, Director of Programs (CCC), for fi-

nancial support and for sponsoring this research. Special

thanks go to IRENA, Managua, Nicaragua, and its di-

rector. Dr. Jaime Incer, for technical, logisticiil, and fi-

nancial support while in the field. For patiently typing

the manuscript, I thank Mrs. Cynthia Mischler, Depart-

ment of Geology, Florida Atlantic University.

LITERATURE CITED

Abbott, R. T. 1974. Anwrican SeasheUs. Second Edition. Van

Nostrand Reinhold Company, New York. 66.3 pp.

Bequaert, J. C. 1942. Cerithidea and Batillaria in the western

Atlantic. Johnsonia 1(5):1-11.

Clench, W. J. and R. T. Abbott. 1941. The Genus Strombus in

the western Atlantic. Johnsonia 1(1): 1-1.5.

Dall, W. H. 1897. List of species collected at Bahia. Brazil by

Dr. H. von Ihering. The Nautilus 10(11): 121-12.3.

Pluck, W. H. 190.5a. Shell collecting on the Mosquito Coast of

Nicaragua. Part I (Introduction). The Nautilus 19(1):8-12.

Fluck, W. H. 190.5b. Shell collecting on the Mosquito Coast of

Nicaragua. Part II. The NauHlus 19(2): 16-19.

Fluck, W. H. 190.5c. Shell collecting on the Mosquito Coast of

Nicaragua. Part III. The Nautilus 19(3):32.

Fluck, W. H. 1905d. Shell collecHng on the Mosquito Coast of

Nicaragua. Part IV The NauHlus (5):55-57.

Fluck, W. H. I905e. Shell collecting on the Mosquito Coast of

Nicaragua. Part V. The Nautilus 19(7):78-S0.

Fluck, W. H. 1906. Shell collecHng on the Mosquito Coast of

Nicaragua. Part VI. The NauHhis 20(1): 1-4.

Harasewych, M. C, PeHt, R.E., and A. Verhecken 1992. Two

new species of Cancellariidae (Gastropoda: Neogastropo-

da) from Brazil. The NauHlus 106(2):4.3-49.

Houbrick, J. R. 1968. A survey of the littoral marine mollusks

of the Caribbean coast of Costa Rica. The Veliger 11(1):

4-23.

Keen, A. M. 1971. Sea Shells of Tropical West America. Second

Edition. Stanford University Press, Stanford, California.

1064pp.

Olsson, A. A. 1922. The Miocene of northern (^osta Rica with

notes on its general stratigraphic relations. Bulletins of

American Paleontology 9(.39):l-.3()9.

Olsson, A. A. 1942. Tertiar\' and Quateniarv fossils from the

Burica Peninsula of Panama and Costa Rica. Bulletins of

American Paleontolog)-, 27(106): 1-106.

Olsson, A. A. 1972. Origin of the exisHng Panamic moUuscan

biotas in terms of tlieir geologic historv and their separa-

Hon by the isthmian land barrier BulleHn of the Biological

Society of Washington 2:117-123.

Olsson, A. A. and TL. McGintv. 1958. Recent marine mollusks

from the Caribbean coast of Panama with the descripHon

of some new genera and species. Bulletins of American

Paleontology .39(177):5-.59.

Olsson, A. A. And RE. PeHt. 1964. Some Neogene Mollusca

from Florida and the Carohnas. Bulletins of American Pa-

leontology 47(2I7):509-575.

Petuch. E. J. 1981. A voluHd species radiaHon from northern

Honduras, with notes on the Honduran Caloosahatchian

Secondary Relict Pocket. Proceedings of the Biological

Society of Washington 94(4): 1 1 10-1 1L30.

Petuch. E. J. 1982. Geographical Heterochrony: Contempo-

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Petuch. E. J. 1987. New Caribbean Molluscan Faunas. The

Coastal Education and Research Foundation, Charlottes-

ville, Virginia. 154 pp.

Pehich. E. J. 1988. Neogene History of Tropical American Mol-

lusks. The Coastal Education and Research FoundaHon,

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Petuch. E. J. 1990. A New Molluscan Faunule from the Ca-

ribbean coast of Panama. The NauHlus 104(2):.57-71.

Petuch. E. J. 1997. Coastal Palcoceanographj of Eastern Noiih

Anwrica {Miocene to Plci.stoccne). Kendall-Hunt Publish-

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Petuch, E. J. and D. M. Sargent. 1986. Atlas of the Living

Olive Shells of the World. The Coastal EducaHon and Re-

search Foundation, Charlottesville, \'irginia. 253 pp.

Radwin. G. E. 1969. A Recent Molluscan Fauna from the Ca-

ribbean coast of Panama. TransacHons of the San Diego

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seum Oceanografico do Rio Grande. Rio Grande do Sul,

Brazil. 331 pp..

Venneij. G. J. and E. J. Petuch. 1986. DifferenHal exHncHon

in tropical American mollusks: endemism, architecture,

and the Panama land bridge. Malacologia 27(1):29— 41.

Vokes, H. E. and E. H. Vokes. 1983. DistribuHon of Shallow-

Water Marine Mollusca, Yucatan Peninsula, Mexico. Me-

soamerican Ecologv- Instihite. Monograph 1:1-183.

Wannke, G. and R. T Abbott. 1961. Caribbean SeasheUs. Liv-

ingston Publishing Co., Narberth, Pennsylvania. 346 pp.

Weisbord, N. E. 1964. Late Cenozoic pelecypods from north-

ern Venezuela. Bulletins of American Paleontology

45(204): 1-564.

Whitmore, F. C, Jr and R. H. Stewart. 1965. Miocene mam-

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185.

Woodring, W. P. 1966. The Panama land bridge as a sea barrier.

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42.5-4.33.

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THE NAUTILUS 111(1):45, 1998

Pase 45

The Protoconch o( Fasti giella carinata Reeve, 1848 (Mollusca:

Cerithiidae)

Jesus Ortea

Laboratorio de Zoologi'a

Facultad de Biologia

Universidad ile 0\iedo

33005 Oviedo, Asturias SPAIN

Jose Espiiiosa

Istitiito de Oceanologi'a

Avenida Priniera 18406 entre 1S4 y 186

Plava, 12100, Havana CUBA

Houbrick, Robertson and Abbott (1987) proxided a de-

tailed description of the shell, operculum, radula and

anatomy of Fa.sti^iclla carinata Reeve, 1848, and con-

firmed the taxonomic position of this previously enig-

matic species in the family Cerithiidae. These authors

also provided information about the ecology and geo-

graphic distribution of this species, which is restricted to

the Great Baliamas Bank and the northwestern coast of

Cuba. The specimens described and figured by Houb-

rick ct al. (1987) lacked not only the protoconch, but

also the first teleoconch whorl. The purpose of this short

note is to describe and illustrate the protoconch of Fas-

tigiella carinata Reeve, 1848.

Young specimens of F. carinata were collected at sev-

eral localities along the northern coast of Cuba. Voucher

specimens are deposited in the Instituto de Oceanologia

and Instituto de Ecologia y Sistematica, in Havana.

The protoconch (Figures 1-2) is white, nearly cyfin-

drical, composed of two whorls, and has a height of 590

|xm. Pronounced iLxial costae are evident within the first

quarter whorl, but disappear after Va whorl. Three to

four spiral cords appear between the first V^ and V2

whorl, becoming well-de\eloped after V2 whorl. Begin-

ning at V2 whorl, small axial plicae appear normal to the

suture, but do not reach the first spiral cord. The tran-

sition to the teleoconch is abrupt and marked b\ the loss

of these axial plicae.

Protoconchs of F carinata are common in the sand at

some localities along northern Cuba (eg. Jibacoa Beach)

(E. Rolan, personal communication). This leads us to

conjecture that this species has lecithotrophic develop-

ment with a brief plank-tonic phase and a high mortafitv-

at metamorphosis. This may contribute to the fimited

geographic distribution of this taxon.

ACKNOWLEDGEMENTS

Special thanks to Dr. Emiho Rolan for providing infor-

mation and the Scanning Electron Micrographs.

LITERATURE CITED

Houbrick, R. S., R. Robertson, and R. T Abbott. 19S7. Anat-

omy and systematic position of Fastigiella carinata Reeve

(Cerithiidae: Prosobranchia). The >4autilus 101(3):101-

110

Figures 1-2. Protoconcfi of Fastigiella carinata. 1. Lateral view, specimen from Esquixal Key, Sagiia la Grande, Cuba. Scale bar

= 100 |xni. 2. Apical view, specimen from Jibacoa Beach, Havana, Cuba. Scale bar = 100 (jlui.

THE NAUTILUS 111(1):46, 1998

Page 46

Notices

THE R. T ABBOTT VISITING CURATORSHIP

The Bailey- Matthews Shell Museum is pleased to invite applications for the 1998 R. T. Abbott Visiting Curatorship.

The Curatorship, established originally in accordance v\ath the wishes of the late Dr. R. Tucker Abbott, Founding

Director of Shell Museum, is awarded annually to enable systematic malacologists to visit the museum for a period

of one to two weeks. Abbott Fellows will be expected, by performing collection-based research, to assist \\ith the

curation of the portions of the museum's Mollusk Collection and to provide one evening talk for the general pubUc.

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THE NAUTILUS

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February 17, 1998

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m8

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THE€9NAUTILUS

CONTENTS

Vohinw IIL Number 2

Febnianj 17, 1998

ISSN 0028-1344

Philippe Bouchet

Geerat Vermeij

Tw(j new deep-water Pseudolividae (Neogastropoda) from

the south-west Pacific

47

Geerat Vermeij

Generic rexision of the neogastropod famiK' Pseudolividae 53

■^vme Biological Lnoorator'

'Voods Hole Oceanography ins^

Library

fEB 2 3 1998

W'.nri'- (loir.. M/\ 0;"-,..|'i

ilulion

THE NAUTILUS lll(2):47-^2, 1998

Page 47

Two New Deep-Water Pseudolhidae (Neogastropoda) from die

Soudi-West Pacific

Philippe Bouchet

Museum national d'Histoire naturelle

55 me Buffon. 75005 Paris FRANCE

E-mail: [email protected]

Geerat J. Vermeij

Department of Geolog\' and Center

for Population Biologv-

Universits' ot California at Davis

Daws, CA 95616 USA

E-mail: \eniieij@geolog\'.ucda\is.edu

ABSTRACT

The new genus Fusulciihis. conchologicallv most smiilar to

Benthobia Dall, 1889 and Zemira H. & A. Adams, 1853, is

erected tor axiallv sculptured species of Pseudolividae with

shouldered whorls and obsolete labral tooth; the columellar

and parietal callus is of verv limited extent, and a parietal rib

at the adapical end of the inner lip is absent. Two new species,

Fuxtdcidus crenatus (t\pe of genus) and F. albtts are described

from bathval (400-.S00 m) hard bottoms at tropical and sub-

tropical latitudes in the southwest Pacific. No post-Paleocene

species of Pseudohxidae are known from the tropical Indo-

Pacific; the habitat of Fusulculus is bathviiietricallv transitional

between those of Benthobia, from abyssal depths, and the \'ar-

ious genera from subtidal waters in southern Australia, South

Africa and Angola.

Key words: Muricoidea, Pseudoli\idae, Fusulculus, labral

tooth, deep-sea. New Caledonia, New Zealand.

INTRODUCTION

Among the manv remarkable mollusks in the deep-water

fauna around New Caledonia are two species that belong

to a pre\iouslv unkiiowii genus of the neogastropod fam-

ily Pseudohvidae. The discoverv of these species docu-

ments the occurrence of the Pseudolividae at tropical

latitudes in the Indo-W'est-Pacific region, whose enor-

mous molluscan fauna is othersvise wholK' lacking in

members of this relict family. Here we introduce the

new genus Fusulculus and describe the Uvo new species

belonging to it.

ABBREVIATIONS USED

LACM Los Angeles County Museum of Natural His-

tory, Los Angeles

MNHN Museum national d'Histoire naturelle, Paris

NM Natal Museum, Pietermaritzburg

NMNZ Museum of New Zealand Te Papa Tongarewa,

WeUington

NZOI National Institute of Water and Atmospheric

Research [formerly New Zealand Oceano-

graphic Institute], WeUington

dd empty shell

Iv collected ahve.

Superorder CAENOGASTROPODA Cox, 1960

Order NEOGASTROPODA Wenz, 1938

Superfamilv Muricoidea Rafinesque, 1815

Family Pseudolividae Fischer, 1884

Genus Fusulculus new genus

Type species: Fusulculus crenatus new species

Description: Small pseudolivid with relatively high-

spired, fusiform shell in which the spire comprises .3.3 to

39% of total shell height; shell consisting of one proto-

conch whorl [in species with non-planktotrophic devel-

opment] and three to six teleoconch whorls; teleoconch

whorls separated bv impressed sutures; base of last

whorl constricted above short siphonal canal; sculpture

of last whorl consisting of a.xial riblets on adapical part

of whorl, crossing finer spiral cords; conspicuous spiral

groove below middle of last whorl, ending in very small

labral tooth at edge of outer hp; below the groo\e, sur-

face adorned with stronger spiral cords; inner side of

outer hp smooth; anterior siphon;il notch very shallow;

fascicle, umbihcal sht, and parietal rib absent; columellar

callus of verv hmited extent, thin; adapical end of outer

lip with obsolete notch. Operculum lanceolate, nucleus

terminal. Radula with broad, sUghtK' concavelv arched

rachicban, with three strong mechallv placed cusps, cen-

tral one stronger; lateral teeth with strong, sohd base,

and short, claw-Uke cusp.

Latin: spindle) and

Etymology: Combination oifusus

sulculus (Latin: small groove).

Remarks: Fusulculus belongs to a group ot Pseudoh-

vidae in which the columellar and parietal callus is of

ver\- limited extent, and in which a parietal rib at the

adapical end of the inner hp is absent. Among Recent

genera, Fusulculus is most similar to Benthobia Dall,

Page 48

THE NAUTILUS, Vol. Ill, No. 2

Figures 1-3. Fti.^ulculiis civiialiis new <;imuis. new species. New Hehriiles Arc. 22°1V'S, ITFISE, 450-.550 ni. 1-2. Holot)pe,

height 16.5 mni; 1. Natnral \ievv; 2. Coated to enhance sculpture. 3. Protoconch, scale bar = 2tW |jini.

1889 (t\pe .species: B. tnjonii Dull, 1889) aiul Z(^mira H.

& A. Adams, 185.3 (t\pe species: Ehtinia aii.stralis G. B.

Sowerby I, 18.33). All three genera are characterized by

a very shallow siphonal notch, a spiral extem;il groove

that is situated just below the middle of the last whorl,

by having the inner side of the outer lip smooth, and by

lacking a lasciolar ridge bordering the anterior notch.

Axial riblets confined to tin- subsutural ramp and shoul-

der are characteristic of Fttsulcuhis as well as of Ben-

tliohia and of the Late Oligocene Australian species Zc-

mira tcsscllatii Tate, 1894 (see Ponder & i:)arragh, 197.5;

Bouchet & VV'aren, 1985). Fttsttlculus further resembles

Bcnthohia in having the labral tooth obsolete, and by

being basallv slightK' constricted; but it differs from Bcn-

thohia h\ lia\'ing shoulilered instead ol evenlv rounded

whorls. In ha\ing a higher spire (spire comprising 33 to

p. Bouchet and G. J. Venneij, 1998

Pay;e 49

39% instead of 25% of shell height), antl a iiiueh less

inflated and more elongate last whorl, and h\ ha\ing fine

instead of obsolete spiral sculpture. Fiisulciilus resem-

bles Zcinira in ha\ing a moderately high spire (spire

about 33% of total shell height in Z. anstrolis) and in

possessing fine spiral sculpture; hut it differs in ha\ing

the last whorl basallv constricted instead of convex-sided

and conical, by having appressed instead of deeply chan-

neled sutures, and in having an obsolete instead of

prominent and sharj) labral tooth. Most species of Zc-

inira, moreover, lack axial sculptme.

In Recent faunas, two other genera with limited callus

development and without a parietal rib are Liiizia Dou-

ville, 1934 (tvpe species: Biuciniim (Ltiizia) costac Dou-

ville, 1934; Miocene, Angola) and yaiidolivo Kilbuni,

1989 (type species; N. caitlinac Killnim, 1989; south-

eastern Africa). Luizia resembles Fusulcuhts in having

axial riblets on the upper part of the whorls, bv having

appressed sutures, and bv the relative Iv high position of

the groove on the last whorl; but it differs from Fiisitl-

ctihis in usuidlv being lower-spired (spire comprising

20% of shell height in L. zebrina (A. Adams, 1855) from

Angola, instead of 33 to 39%), in having the inner side

of the outer lip finely firate instead of smooth, bv having

a distinct labral spine, and in lacking the basal constric-

tion of the last whorl. Naudoliva, like Fustilculus. is rel-

atively high-spired (spire comprising 36% of total shell

height) and basallv constricted, and the inner side of its

outer hp is likewise smooth; but it differs from Ftisul-

culus in having a long labral spine, which is situated at

the end of a shghtlv raised spiral band near the base of

the shell rather than at the end of a groove that is sit-

uated at a high position (jn the whorl. Naudoliva, more-

over, has a siphonal fasciole, which is lacking in Fiifiid-

ctdtis.

The Late Cretaceous to Late Eocene genus Calore-

hama Squires, 1989 (tvpe species: Pscudoliva dillcri

Dickerson, 1914; Cahfornia) is characterized bv a broad-

ly fusiform shell with a moderately low spire (spire com-

prising less than 27% of total shell height), eight to

twelve axial folds often raised as nodes at the shoulder

or periphen, and fine spiral sculpture over the whole

whorl surface, as well as by the absence of a parietal

thickening and of outer-Up Urae. As in Fusidcidiis, the

spiral groove is located at a relativelv high position on

the whorl. Calorcbama chffers from Fusiiladiis by its

lower spire, and by having a conical or evenlv convex

last whorl instead of a basallv constricted bodv vvhorl.

The adal sculpture of Fusiilniliis is finer

Fusidculus crenatiis new species

(Figures 1—4, 6)

Type material: Holotvpe (dd) in MNHN; paratvjies

in NMNZ (1 dd), NM (Idd), LACM (1 dd) and MNHN

(2 Iv, 7 dd).

Type locality: New Hebrides Arc, 22°17S, 17ri8E,

450-550 m [VOLS MAR sta. DW30].

Material examined: Coral Sea. MUSORSTOM 5; sta.

313, 22°24S, 159°33E, 780-930 m, 4 dd.— Lovaltv Ba-

sin. BIOGEOCAL: sta. DW289, 20°36S, 167°o6e,'830-

840 m, 3 dd.— New Hebrides Arc. VOLSMAR: sta.

DW.30, 22°17S, 17n8'E, 450-550 m, 2 Iv, II dd (type

material)— Walhs & Futuna. MUSORSTOM 7: sta.

DW540, 12°27S, 177°28W, 600 m, 1 Iv, 12 dd.; Sta.

DW578, 13°08S, 176°16-W, 640-730 m, 2 dd.; Sta.

DW586, 1.3°11S. 176°13"W, 510-600 m, 2 dd (all

MNHN).

Distribution: SW Pacific, from the Coral Sea to NE

of Fiji, alive in .5.50-600 m, shells to 8.30 m.

Description: Shell thick, solid, ovoid, consisting of 0.9

protoconch and 5.7 teleoconch whorls, protoconch with

smooth glassy shell, teleoconch more chalky, with thin

and fragile intritacalx and strong incremental fines, most

distinct in lower parts of shell surface between ribs. Te-

leoconch whorls convex with a distinctlv concave sub-

sutural ramp occupving about 2/3 of exposed height of

spire whorls. Sculpture consisting of strong axial ribs, 14

per whorl, and weaker spiral cords; one adapical cord

just adjacent to suture, no cord but onlv very indistinct

spiral threads in subsutural ramp, 2 cords below ramp

(5 on penultimate whorl), forming prominent nodules at

intersection with ribs. Last whorl with ca. 10 cords adap-

icallv of groove, those on shoulder stronger. 6 abapically

with interspaces broader than cords; groove narrow and

shaqi, terminating in a short, pointed triangular tooth at

aperture; axial ribs not extending abapically of groove.

Aperture ovate with broadlv open siphonal notch, colu-

mella simple; outer fip not thickened, regularlv convex

except for constriction at level of subsutural ramp. Color

salmon brown with whitish intritacalx on very fresh

shells, fading in dead collected specimens. Operculum

(figure 6) lanceolate, nucleus temiinal. Radula (figure 4)

with broad, slightly concavelv arched rachidian. with

three strong mediallv placed cusps, central one stronger;

lateral teeth with strong, sofid base, and short, claw-fike

cusp.

Dimensions: Holotvi^e height 16.5 mm, diameter 7.9

mm, aperture length 10.4 mm, aperture width 4.0 mm.

Largest specimen, height 18.4 mm.

Remarks: A comparison with F alhus is given under

the latter species.

Fitsidcidiis alhus new species

(Figures 5. 7, 8-14)

Type material: Holotvpe (dd) in MNHN; paratvpes

m NMNZ (1 dd), NM (Idd), LACM (1 dd) and MNHN

(3 Iv, 15 dd).

Type locality: North of New Caledonia, 18°42'S,

163°13E, 668 m [MUSORSTOM 4 sta. DW160].

Material e.xamined: New Caledonia (dl MNHN).

MUSORSTOM 4: sta. DW160, 18°42'S, 163°13'E, 668

m, 3 Iv, 19 dtl (tvpe material).; BIOCAL; sta. DW48,

Page 50

THE NAUTILUS, Vol. Ill, No. 2

Figures 4—5. Radiilaf ol Fusiili-iilti.s specie.s. 4. Fustilailiis

crenatus new species, taken from paiatvpe from t\pe localitrv,

scale bar = 50 |xm. 5. Fitsiilculus albtis new species, taken

from paratvpe from type locality, scal^ bar = 20 \xm.

Figures 6-7. Opercuia of Fiisiilciilus species. 6. Fusiilcuhis

crctuilii.s new species, from paratN-jie from hpe localitv, leni!;th

6.5 mm. 7. Fusulmlns alhus new species, taken from parat\pe

from hpe localitv, lengtb 2.85 mm.

23°()()S, 167°29'E, 775 m, 2 del.; BATHUS 4: sta.

DW917, 18°47"S, 16.3°14E, .397-400 m, 5 dd.— New

Zealand. Three Kings Rise. R.V. Acheron, sta. BS391,

34°()1S, 172°07E, 622 m, 7 dd (NMNZ M34932).; R.V.

Tangaroa, sta. BS633, 34°2()S, 171°48E, 440 m, 4 dd

(NMNZ M100.323).; Sta. BS634, 34°18'S, 171°45'E, 427

m, 2 dd (NMNZ Ml()0.322).; Sta BS896, .34°0rS,

17r45'E, 201-216 m, 1 dd, 1 fragm. (NMNZ

M 100324).; Sta. BS898, 34°0rS, 171°44'E, 206-211 m,

1 dd (NMNZ M100321).; NZOI sta. U602, 3r31S,

172°50'E, 1216-1385 m, 1 dd (NZOI).

Distribution: New Caledonia, alive in 668 m, shells

in 400-775 ni; north of New Zealand, Three Kings Rise,

shells in 211-622 ni, with a single shell, probably trans-

ported vertically, in 1216-1.385 m.

Description: Shell thick, solid, ovoid, consisting of

0.8 protoconch and 3.2 teleoconch whorls, protoconch

with smooth glassy shell, teleoconch more chalky, with

thin and fragile intritacalx and strong incremental lines,

most distiTict in lower parts of shell surface between

ribs. Teleoconch whorls convex with a distinctly con-

cave subsiitural ramp occupying nearly half of exposed

height of spire whorls. Sculpture consisting of low,

broad a-xial ribs and low spiral cords, indistinct except

at their intersection with ribs; one adapical cord just

adjacent to suture, no cord in subsutural ramp, 3 cords

below ramp, most adapical one forming prominent

nodules at intersection with ribs, most abapical one

very indistinct. Last whorl with 4 cords adapically of

groove, 5 abapically; groove very sharp, terminating in

a pointed triangular tooth at aperture; a.xial ribs not

extending abapically of groove. Aperture ovate with

broadlv open siphonal notch, columella simple; outer

lip not thickened, regularlv convex e.xcept for a sUght

constriction at level of subsutural ramp and at level of

groove. Color white. Operculum (figure 7) and radula

(figure 5) as in F. crenatus.

Dimen.sions: Holotvpe height 7.9 mm, diameter 4.0

nun, aperture length 4.8 mm, aperture width 2.15 mm.

Largest specimen^BATHUS 4 sta. DW917], height 10.3

mm.

Remarks: Fiisitlciilii.s alhus differs from F. crenatus by

haxing a smaller adult size, by being white instead of

salmon brown in colour, and bv ha\ing the axial riblets

mainly confined to the shoulder area instead of extend-

ing from the suture to the groove. Fusulculus alhus is in

these respects more like Benthohia than is F. crenatus.

The t^vo species co-occur in the New Caledonia region,

liut are never SMitopic, nor svmpatric.

The material from Three Kings Rise all consists of

empty, sometimes worn or fragmentary shells. General

shi-Il moiphology and proportions are similar to the ma-

terial from New Caledonia. Most specimens ha\e

liroadiv spaced axial ribs and onlv weak spiral cords

adapicallv of the spiral groove, thus the shoulder is not

muricated as in the New Caledonia specimens (figure

14). One sample differs h\ having more numerous axial

p. Boiichet and G. J. Vermeij, 1998

Page 51

Figures 8-14. Ftisiilailns albiis new species. 8. Holot\pe, height 7.9 mm. nortli of New Caledonia, 18°4i2'S, 163°13'E, 668 m.

9. High-spired specimen, height 10. .3 mm, north of New Caledonia, 1S°47'S, 163°14'E, ,397—400 m. 10-11. Low-.spirecl specimen,

height 6.2 mm, south of Ne\K' Caledonia, 2.3°00'S, 167°29'E, 77.5 ni; 11. Protoconch, scale line = 200 (xm. 12-13. Specimens from

Three Kings Rise, height 6. .5 and .5,9 mm respecti\elv, north of New Zealand, 622 m (NMNZ M.349.32). 14. Specimen with weak

spiral sculpture, height 7.1 mm, Nf>rth of New Zealand, 440 m (NMNZ M 100323).

ribs and strong spiral sculpture, extending adapicallv of

the spiral gro()\e (figures 12-1.3). Despite these differ-

ences, we interpret the New Caledonia and New Zea-

land material as probably representing a single species

with discontinuous distribution along the Norfolk

Ridge.

BIOGEOGR.A.PHICAL REMARKS

The family Pseudohvidae was distributed nearly world-

wide during the Late Cretaceous and Paleogene, but it

has become progressively biogeographically restricted

during the Neogene. In Recent seas, the family is rep-

Page 52

THE NAUTILUS, Vol. Ill, No. 2

resented in abyssal and bathyal waters by Benthohia,

which is known from the deep Atlantic, Indian Ocean,

and New Ze;iland (see Bouchet & Waren, 1985; Kantor,

1991). Shallower-water genera are known from New

Caledonia (Fusulcuhis). temperate Australia (Zcmira).

warm-temperate southeastern Africa (Natidoliva) , tem-

perate and tropical western Africa (Pseudoliva Swainson,

1S40; Fitlmciittim Fischer, 1S84; Luizia), and western

warm-temperate North America {Macron H. & A. Ad-

ams, 1853) and the Panamic Province (Tiiumphis Gray,

1857) (see Vermeij, 1998). During Miocene and Plio-

cene time, the family was also represented in temperate

western South America bv Tcstolliiiui Vermeij and

DeVries, 1997.

Beside the abyssal species of Bcutliohia. Fiisulnihis is

the only genus of Pseudolividae known from moderately

shallow water in the vast Indo-West Pacific region since

Piileocene time. The family disappeared from Europe

and Japan after the Late Eocene.

ACKNOWLEDGEMENTS

We thank Bnice Marshall (NMNZ) who brought our at-

tention to the material of Fusulcuhis under his care, and

Anders Waren (Naturhistoriska Riksmuseet, Stockliolm)

who prepared the radulae for scanning electron micros-

copy. The research was funded in part by a grant (NSF

EAR-94-05537) from the National Science Foundation

to Vermeij.

LITERATURE CITED

Bouchet, P. and A. Waren. 1985. Re\ision of the northeast

Atlantic bathyal and abyssal Neogastropoda e.xcludingTur-

ridae (Mollusca, Gastropoda). Bollettino Malacologico,

Suppl. 1:123-296.

Kantor, Yu.I. 1991, On the morphology' and relationships of

some olivifonii gastropods, Ruthenica l:17-.52.

Ponder, W. F, and T. A. Darragh, 1975, The genus Zcmira H,

and A, Adams (Mollusca: Gastropoda). Journal of the Mal-

acological Society of Australia .3:89-105.

Vermeij, G. J. 1998. Generic Revision of the Neogastropod

Family Pseudolixidae, The NauHlus lll(2):5.3-84.

Vermeij, G. J., and T. J De\'ries. 1997. Taxonomic remarks on

Cenozoic pseiidoli\id gastropods from South America.

The Veliger 41:2:3-28. "

THE NAUTILUS lll(2):53-84, 1998

Page 53

Generic Revision of the Neogastropod Family Pseudolividae

Geerat J. Vemieij

Department ot Geologv' and Center

tor Population Biolog\'

University of California at Da\is

Davis. CA 95616 USA

vernieij @geolog\'. ucda\is edvi

ABSTRACT

The neogastropod family Pseudolividae Cossmann, 1901, is a

Late Cretaceous to Recent group of about one hundred species

characterized bv a spiral groove (pseudolivid groove) on the

lower half of the last whorl, a labral tooth on the outer lip at

the end of this groove, a posterior notch at the adapical end

of the outer hp, spiral sculpture increasing in prominence to-

ward the base, and an indeterminate, shaq^-edged, planar outer

lip. A genus-level revision based on shell characters supports

previous anatomical studies in recognizing the Pseudolividae as

a ta.\on distinct from other neogastropod famihes.

I recognize fourteen genera in the Pseudolividae: Benthobia

Dall, 1889 (Recent, low-latitude bathval and abvssal oceans);

Fulmenttim Fischer, 1884 (Recent, West and South Africa); Fii-

sopsis Ravii, 19.39 (Paleocene, Denmark); Ftisulculiis Bouchet

& Vemieij, 1997 (Recent, New Caledonia); Hubachia Etavo

Senia. 1979 (Paleocene. Colombia): Luizia Douville. 1933

(Earlv .Miocene to Recent. West Africa); Macron H. & A. Ad-

ams, 1853 (Early Miocene to Recent, wanii-temperate North

Pacific and Caribbean); Naudoliva Kilbum, 1989 (Recent,

southeastern Africa); Pseudoliva Swainson, 1840 (Earlv Mio-

cene to Recent, West Africa); Sulcobiiccinum d'Orbigny, 1850

(Late Cretaceous (Campanian) to Earlv' Oligocene, wann seas

worldwide); SiilcoUva. new genus (tvpe species: Pseudoliva

monilis Ols.son, 1928; Eocene, Peru); TestaUiuin Vemieij &

De\'ries, 1997 (Earlv Miocene to Late Pliocene, westem South

America); Triumphis Grav, 1857 (Early Miocene?, Recent,

tropical eastern Pacific); and Zemira H. & A. Adams, 1853

(Late Eocene to Recent, Australia). I designate Buccinum fis-

suratimi Deshayes, 1835, the type species of Sulcobuccinum.

of which the following taxa are subjective junior sviionviiis:

Buccinorbis Conrad. 1865; Calorebama Squires, 1989; Pcgo-

comphis Ziiismeister, 1983; and Popcnoeiim Squires, Zinsmeis-

ter & Paredes-Mejia, 1989, The genus St/lianocochUs Melvill,

1903, is a junior subjective sviionvni ot Fidmentum. Pseudoliva

guppyi Mansfield, 1925. from the Late Miocene of Trinidad,

is assigned to Fusulculus with great hesitation. I propose Sul-

cobuccinum midtinodulostnn as a replacement name for Pseu-

doliva chavani Glibert, 1973, non Tessier, 1952. ^tacron

mcleani is a new species described from the Recent fauna of

the Gulf of California.

Genera removed from Pseudolividae include Austrosphnera

Caniacho in Furque & Camacho, 1949 (Late Cretaceous to

Paleocene, Argentina); Seymourosphaera Oleinik & Ziiismeis-

ter, 1996 (Earlv Paleocene, Seymour Island, Antarctic Penin-

sula); and the Late Cretaceous genera Fulfi^erca Stephenson,

1941, Hijdwtnbulus Wade, 1916, and Ptychosyca Gabb, 1877,

all from the Gulf Coast;il Plain ot the United States, These taxa

are not reassigned to other taiiiilies, but belong to basal buc-

cinoids. Niccnm Woodring, 1964, which has sometimes been

considered a subgenus of Triumphis, is a Late Miocene to Re-

cent tropical American genus here assigned to the buccinid

subtamilv Photinae. The Pliocene to Recent South African ge-

nus Melapium H. & A. Adams, 18.53, may belong to the largely

Paleogeiie family Strepturidae,

Key words: Muricoidea, Psuedolividae.

INTRODUCTION

The Pseudolividae is a distinctive family of some one

hundred species of neogastropods, whose origin can be

traced to the Late Cretaceous. The history of this family

contrasts markedly with that of many other neogastro-

pod families including the Muricidae, Fasciolariidae,

CanceUariidae, Mitridae, and Conidae, which Bkewise

originated during Late Cretaceous times (Taylor et al.,

1980; Bandel, 1993). Whereas these latter families un-

derwent spectacular phases of diversification during both

the Paleogene and Neogene, the Pseudohvidae have de-

chned in diversitv and geographical range since their

peak during the Paleogene. The group is represented

today by only about sixteen species, a number that is

one to two orders of magnitude smaller than the living

diversitv- in each of the other famihes.

In order to understand why some clades diversified

during the Neogene while others dwindled in diversitv,

we must specify- the order and timing of evolutionary

branching events, as well as the ecological and geograph-

ical context of evolution and e.xtinction in the contrasting

groups. Such a phvlogenetic understanding, in turn,

hinges critically on a comprehensive account of the tax-

onomy of the groups in question.

The aim of this paper is to present a taxonomic review

of all genera of Pseudolividae and to reassign genera that

have been previously referred to the Pseudolividae or

that were considered to belong to genera here assigned

to that family. In the second part of the study (Vermeij

& Carlson, in preparation), we present a phvlogeny of

Page 54

THE NAUTILUS, Vol. Ill, No. 2

the group together with a narrative evolutionary and bio-

geographit;J hi.stoiT ot the P.seudolividae.

PREVIOUS WORK

A comprehen.sive genus-level review of the PseudoLvi-

dae has not been attempted in more than fiftv years.

Cossmann (1901), Thiele (1929), and Wenz (1938-44)

compiled the genera ot the group, but they did not crit-

ically evaluate the Umits of the taxa they recognized, and

assigned most members of the Pseudolividae to the type

genus Psnidoliva. Adegoke (1977) recognized the diver-

sity of Paleocene and Eocene pseudolivitls was too great

to be accommodated in the genus PscudoUva and its

subgenus Buccinorbis, but he dechned to propose new

superspecific taxa. Although some of these groups were

subsequently named (Zinsmeister, 19(S3; Squires, 1989;

Squires et al., 1989), the naming of fossil members of

the family proceeded piecemeal, and several superspe-

cific taxa named earlier were overl(X)ked. Witli the exception

of Ponder and Darragh's (1975) paper on Zcinira. which

treated fossil as well as li\'ing species, most studies ot

Recent pseudohvids did not incorporate data on fossil

forms.

Varying interpretations of the phylogenetic position of

the Pseudohvidae have contributed to confusion sur-

rounding the taxonomic assignment of several genera, as

well as to difficulties in defining the hmits of the family.

Perhaps because Pscudoliva has a smooth shell superfi-

cially resembling some members of the OliNidae, many

twentieth-century authors followed Swainson (1840) in

placing Pscudoliva in or near the Olividae (see e.g. Mel-

vill, 1903; Thiele, 1929; Wenz, 1938-44; Gilbert', I960;

Tavlor & Sohl, 1962; Sohl, 1964a, b; Ponder & Darragh,

1975; Adegoke, 1977; Ponder & Waren, 1988; Kilbum,

1989). Strongly sculptured genera such as Litizia, Ma-

cron, and Triiimphis were therefore not recognized as

pseudoli\dds, and were instead referred to the Buccini-

dae (Thiele, 1929; Dou\alle, 1933; Wenz, 1938-44; Gli-

bert, 1963; Addicott, 1970; Keen, 1971; Abbott, 1974).

In their treatment of gastropod genera, H. & A. Ad-

ams, (1853) assigned their new genus Zcmira , together

with Pscudoliva and its new subgenus Macron, to the

muricid subfamily Purpurinae, a taxon encompassing

genera that are assigned by modern authors to the mur-

icid subfamilies Rapaninae ami Ocenebrinae (see e.g.

Kool, 1993; Vermeij & Kool, 1994; Vermeij, 1995; Ver-

rneij & Vokes, 1997; Vermeij & Carlson, in preparation).

This assignment was supported by characters of the rad-

ula and by the mistaken assertion that the opercula of

these genera have a lateral nucleus, as do inanv nuni-

cids. Beyrich (1854) referred two small Lattorfian ( =

Priabonian, latest Eocene) species from North Germanv

to the gemis Puqnira, but he did not coiriment on his

reasons lor this assignment. One of these species, P. pus-

ilia, was thought by Beyrich to lack the characteristic

pseudolivid groove. Von Koenen (1889) referred the two

species to Pscudoliva. [lointed out that both exhibited

the characteristic groove, and m;iintained that they have

little in common with Puqmra.

Sowerbv ( 1846) was the first author to propose a clo.se

relationship between Pscudoliva and the Buccinidae. He

considered Pscudoliva to he closely related to Ebunm

Lamarck, 1822 (non Lamarck, 1801), the buccinid genus

knowii todav as Babijlonia Schliiter, 1838. Cossmann

(1901) considered the Pseudolixinae as a subfamily of

Buccinidae and included in it Pscudoliva. Buccinorbis,

and Fubtwntum: but he assigned Zcmira and Macron to

the Latruncuhnae (= Babyloniinae) and Triumphis to

the CominelUnae (= Photinae). Assignment oi Pscudo-

liva, its subgenus Buccinorbis, and related Paleogene

genera to the Buccinidae or Buccinoidea was accepted

isy von Koenen (1889), Fischer (1884), Bohm (1926),

Palmer (1937), Gardner (1945), Golikov & Starobogatov

(1975), Docker/ (1977, 1880), and Squires (1989),

among others. Shell characters formed the basis for most

of these assignments.

Kantor's (1991) anatomical and phylogenetic study

provided strong support for separating the Pseudoli\idae

as a distinct family from otlier oh\iform gastropods (see

also Kantor, 1996). He isolated the group in a new sub-

order Pseudohvoidei, which together with the Muricoid-

ei comprises the order Rachiglossa. Many derived fea-

tures in Ohvidae, including pedal lobes, inner-whorl re-

sorption, posterior mantle tentacle, mobile crescent-

shaped propodiuni, and absence of the operculum, for

example, are absent in Pseudolividae. Character states

of Pseudolividae interpreted to be plesiomoqjhic for

neogastropods include position of radular sac at base

rather than near tip of proboscis, and (in Bcntltobia) the

connection behveen the radular retractor muscle and

the columellar muscle. In PseudoU\idae, the proboscis

is formed bv elongation of the buccal tube, and there is

a well defined Gland of Leiblein; whereas in the Muri-

coidei, which includes the Olividae and Buccinidae, the

proboscis is formed by the elongation of the dorsal wall

of the buccal cavity (Kantor, 1991).

The present study is based on shell characters. Many

authors dismiss shell characters as unreUable tor infer-

ring phvlogeny above the species level on the grounds

that the shell is luore subject to environmental variation

and to CNolutionaiT convergence than are the animal's

internal organs. Not only is this argiunent predicated on

untested and probablv false assumptions, but it also ef-

fectively eliminates phylogenetic assessments ot fossil

forms. There can be no question that data on anatomy

aiul on DNA sequences are immensely valuable in con-

structing classifications and plnlogenies of living .species;

l)ut thev have only limited power in helping paleontol-

ogists make sense of the thversity of shells encountereil

iTi the fossil record. Instead of ignoring shell characters,

we should exaluate the latter carefully The aim of the

present study is therefore to present a unified scheme

of classification of both fiving and fossil pseudoli\ids

based on a detailed analysis of shells.

G. J. Vermeij, 1998

Page 55

MATERIALS AND METHODS

I ha\e exaniineil the hpe species of all but two genera

of the Pseudoluiilae, as well as main' acklitioiiai species

within and outside the family. A complete hst of recog-

nized species of PseudoUvidae, together with material

examined, is gi\en in AppendLx I. Material of species

discussed in this paper hut not belonging to the Pseu-

dolidae is listed in Appendix 2.

ABBREVIATIONS

ANSP Academy of Natural Sciences, Philadelpliia,

Pennsylvania, USA

BMNH The Natural History Museum, London, UK

CAS California Academy ot Science, San Francisco,

California, USA

IGPS Institute for Geology and Paleontolog)', To-

hokii University, Sendai, Japan

IRSNB Institut Roval des Sciences Naturelles de Bel-

giijue, Bnissels, Belgium

LACM Los Angeles County Museum of Natural His-

tory', Los Angeles, California, USA

MNHN Museum national d'Histoire naturelle, Paris,

France

MGS Mississippi Geological Sun'ey, Jackson, Missis-

sippi, USA

NM Natal Museum, Durban, South Africa

PU Purdue University, West Lafavette, Indiana

USA

PRI Paleontological Research Institute, Ithaca,

New York, USA

RGM Nationaal Museum voor Natuurlijke Historic,

Leiden, The Netherlands

SAM South African Museum, Capetown, South Af-

rica

SCO PI Museo Nacional de Chile, Instituto de Palon-

tologia, Santiago, Chile

TMM Texas Memorial Museum, Austin, Texas, USA

UCMP University of Cahfoniia Museum of Paleon-

tolog)-, Berkeley, California, USA

USNM United States National Museum of Natural

History, Washington. DC, USA

GJV Vermeij collection

YPM Yale Peabody Museum, New Haven, Con-

necticut, USA

SYSTEM ATICS

Class Gastropoda

Subclass Proscjbranchia

Order Neogastropoda

Superfamily Muricoidea Rafinesque, 1815

Family Pseudolividae Fischer, 1884

Diagnosis: Shell verv small to large, maximum height

5 to 100 mm, ovate to fusiform; spire consisting of six

or fewer teleoconch whorls; last whorl comprising 60%

or more of total shell height; spiral sculpture usually

present, consisting of a deep groove (pseudolivid groove)

on basal half of last whorl, and of cords or threads that

increase in degree of expression abapically; axial sculp-

ture, when present, consisting of eight or more ribs per

whorl, restricted to upper part of whorl; outer lip thin,

shaip at edge, the adaxial laver not reflected over the

edge to the abaxial side; pseutlolivid groove terminating

at edge of outer lip in blunt or sharp, ventrally directed

tooth (labral tooth); outer lip often crenulated below la-

bral tooth by terminations of grooves between basal

cords; edge of outer lip planar, without sinuses except

for adapical notch where outer lip joins penultimate

whorl; inner Up smooth, adherent throughout its length

to penultimate whorl, weaklv concave, sometimes with

weak fold at entrance of siphonal canal, sometimes with

parietal tooth at adapical end; columellar and parietid

callus variably expressed; li\ing species with persistent

periostracum.

Included genera: Bcnthohia Dall, 1889; Buccinorbis

Conrad, 1865; Fiilnientiim Fischer, 1884; Fti.sopsi.s Ra\m,

1939; Fiisiilailus Bouchet & Vermeij, 1997; Luizia Dou-

ville, 1933; Huhaclua Etavo Serna, 1979; Macron H. &

A. Adams, 1853; P.sciidoUva Swainson, 1840; Sulcobuc-

ciruim d'Orbigny, 1850; Suholiva new genus; Tesfallium

Vermeij & DeVries, 1997; Triumphis Grav 1857; Z^'inira

H. & A. Adams, 1853.

Stratigraphic range:

to Recent.

Late Cretaceous (Campanian)

Discussion: Characters: None of the shell charac-

ters of the Pseudoli\adae is by itself unique to the family,

but the constellation of characters senes to set the

group apart from other neogastropod clades. Perhaps

the most distmctive feature of the Pseudohvidae is the

so-called pseudolivid groove, a spiral depression on the

lower part of the whorl that terminates in a \entrallv and

often adaxialK- projecting tooth (labral tooth) on the out-

er edge of the outer lip. The groove and tooth are ob-

solete in Bcnthohia and Macron Uvidits. and absent in

Triumphis.

A labral tooth at the end of a groo\'e is not uni(jue to

the Pseudolividae. It has also evolved numerous times

in the Muricidae (many members of the subfamilies

Muricinae, Ocenebrinae, and Rapaninae), Buccinidae

(Pisaniinae of the genus Canthanis Roding, 1798, and

related taxa; and Siphonaliinae of the Eocene genus

Fhoracanthtis Cossmann & Martin in Martin, 1914),

Echinofulgurinae (members of the genera Coniiilina

Conrad, 1853, and Protobusi/con Wade, 1917), Fascio-

lariidae (the genus Opcatostoina Berrv, 1958), Turbinel-

lidae (the genus Ccratoxancus Kurotla, 1952), Olividae

(many members of the subfamily Ancilhnae), and various

other buccinoids such as the Cretaceous genera Bucci-

nopsis Conrad, 1857, and Odontobasis Meek, 1876, as

well as the Recent Tritonidca dcntata Schepman, 1911

(see e.g. Sohl, 1964a, b; Bouchet & Waren, 1985; Vig-

non, 1931; Vokes, 1990, 1992; Vermeij & Kool, 1994;

Vermeij & Vokes, 1997; DeVries & Vermeij, 1997).

The simple, sharp-edged outer lip of fully grown

Page 56

THE NAUTILUS, Vol. 111. No. 2

pseudolivids does not differ from that of ontogenetically

vounirer indixidual.s, and therefore indicate.s indetermi-

nate growth aec-ording to the criteria .set out liv Vermeij

and Signor (1992). The adult outer lip oi' Triumphis and

of several large Eocene species of Sulcobticciniim is of-

ten adapicalK- more e.xtended than is that of individuals

at earlier stages of growth, so that the spire comes to

ha\e a strongly conca\e lateral profile; hut there is no

evidence that growth in the spiral direction ceases at

maturit)-, as would be the case in shells with determinate

growth. Manv pseudoh\ids (especially Eocene species of

Sttkobucciman) develop massive columellar and parietal

callus deposits in adulthood, but agiiin there is no evi-

dence that growth at the outer Up ceases.

The indeterminate growth pattern of pseudolivids ap-

pears to be the plesiomoiphic condition in neogastro-

pods. It characterizes all Cretaceous huccinoids as well

as Cenozoic Melongenidae, TurbineUidae, Fasciolari-

idae, Olividae (AncilUnae), and many clades within Mur-

icidae, Buccinidae, and Conoidea, among others (see

Vermeij & Signor, 1992).

The edge of the outer lip of PseudoUvidae usually lies

in a plane. Its contour is interrupted only b\' the emar-

gination at the siphonal canal and bv the adapical notch

(see below). A planar outer hp also characterizes many

Muricidae and Conidae. In manv other neogastropods,

the edge of the outer lip is medially convex downward,

the conve.x sector being flanked adapically and some-

times abapicallv by a shidlow, downwardlv concave sinus.

Variations on this more convex outer Up are found

among Buccinidae, Mitridae, Costellariidae, Melongen-

idae, Nassariidae, Fasciolariidae, TurbineUidae, and

manv other groups.

In fully grown Pseudolividae, the smooth inner (ad-

axial) layer of the outer lip does not extend across the

edge to the outer (abaxial) side. As a result, the edge is

sharp and not glazed. This configuration occurs also in

Muricidae, pisaniine Buccinidae, and Conidae, among

others. The edge of the outer lip is polished smooth by

the inner layer extending across it in most photine and

babvloniine buccinids, nassariids, mitrids, and melon-

genids.

Nearly all members of the Pseudolividae are charac-

terized by a posterior (or adapical) notch of the outer

lip. This is an indentation where the outer lip joins the

penultimate whorl at the adapical end of the aperture.

A comparable notch occurs in many gastropods, includ-

ing many muricine and rapanine muricids, columbeOids,

photine buccinids, melongenids, nassariids, mitrids, vol-

utids, oli\i(ls, and conoideans. Within Pseudolividae, a

few genera (Bcntliobia. Fii.sulculii.s. Sandoliva. and some

large inilividuals ol Triuntphi.s) apparently lack a poste-

rior notch.

Spiral sculpture in the Pseudolividae is usuallv better

expressed on the lower part of the whorl than on the

upper portion. As a result, crenulations at the ends of

spiral grooves are most strongly expressed on the basal

sector of the outer lip below the pseudolivid groove.

This character state occurs widely among neogastropods

including Columbellidae, mam Buccinidae (Siphonali-

inae, Photinae), Nassariidae, many Fasciolariidae, Cos-

telariidae, Mitridae, Harpidae, and manv Conoidea. In

.some Melongenidae, this pattern is superimposed on a

spiral sculpture of keels or folds that decrease in prom-

inence toward the base. In muricids and in the buccinid

subfamilv Pisaniinae, the strongest spiral sculptural ele-

ments alwavs occur on the upper part of the whorl near

the shoulder, decreasing both toward the base and from

the shoulder to the suture above.

In Pseudolividae bearing axial sculpture, the axial el-

ements are confined to the part of the last whorl above

the pseudoli\id groove. The restriction of axial sculpture

to the upper part of the whorl is very widespread in

Cretaceous neogastropods (Sohl, 1964a, b), and also

characterizes many clades in the Buccinidae, Fasciolar-

iidae, Nassariidae, Columbelhdae, and Conoidea, among

others.

The inner lip of Pseudofividae is smooth and always

adlierent above the siphonal fasciole, that is, its edge

merges imperceptibh with the outer surface of the pre-

ceding whorl on which the inner lip encroaches during

growth. Even when a thick callus is formed on the inner

hp, as in species of Sulcobiiccinum and Psctidoliva, the

abaxial edge of the callus is rareK distinct. A smooth,

adherent inner fip also characterizes Melongenidae, ba-

sal Conoidea, and many groups within Buccinidae, Mur-

icidae, and Fasciolariidae (Fusininae). In most Oli\idae,

including the Ancilhnae, the inner lip is sculptured by

thin, often very numerous spiral riblets; but a smooth

inner hp occurs in the ancilfine genus Ebunm Lamarck,

1801.

In most Pseudolividae, the columella terminates abap-

icaUy in a simple point on the left (adaxial) side of the

siphonal canal or notch. This condition is widespread in

Cretaceous buccinoids as well as in Cenozoic Buccinidae

(Buccininae, Volutopsiinae) and basal Conoidea. In the

pseudolivids Macron and Triumpliis. however, there is a

weak fold on the columella at the upper entrance of die

siphonal canal. The abapical end of the columella there-

fore appears tnmcated in these genera. A basal colu-

mellar fold is verv widespread in neogastropods includ-

ing Muricidae, Fasciolaiiidae, and many Buccinidae (Pis-

aniinae and Siphonaliinae), but the tnincation observed

in Macron and Triinnphis is less common, being best

knowni in the Nassaiiidae and the buccinid subfamily

Photinae (see Allmon, 1990).

The protoconch in most pseudohvids is paucispiral,

here defined as ha\ing fewer than three whorls. The onlv

genera with a multispiral protoconch are Bcnthobia and

Fusopsis. Multispiral protoconchs are extremely wide-

spread among Cretaceous buccinoids, and hkely repre-

sent the primitive state of neogastropods in general and

of the Pseudolividae in particular.

ConipariM)ns with other Ncog^astropods: Some Pseu-

dolividae superficially resemble the Ohvidae in having a

posterior (adapical) notch, a relatively smooth shell, a

low spire, and thick c;dlus deposits on the columellar and

G. J. Vermeij, 1998

Page 57

parietal regions. Memhers ot the olhid suhfaniilv Aneil-

linae turther resemble pseudolhitls in usuallv haxing a

basal spiral groove (aneillicl grooxe) and correspomling

labral tooth. The ancillid groove is fonned at the adap-

ical margin of a broad, flat, somewhat raised zone (an-

cillid band ) which is situated above the siphonal fasciole

and which lacks the spiral cords or threads characteristic

ol pseudoliNids. The labral tooth ot Ancillinae lies at the

entrance of the siphonal canal, whereas in PseudoUvidae

it is situated a short distance abo\'e the entrance of the

canal, from which it is separated b\ a short crenulated

sector of the outer lip. C)h\idae linther ditler from Pseu-

doh\idae in ha\ing most or all of the shell coxered with

a callus glaze, which is deposited by pedal lobes that are

absent in Pseudolividae (Kantor, 1991).

Gardner (1945) thought that the Eocene genus An-

cillop.sis Conrad, 1865 (tvpe species: Ancillario altilis

Conrad, 18.32) might be closelv related to Psciidoliia.

This genus is characterized by an ovate, smooth, basallv

unconstricteil sheU, bv a smooth concave columella, a

posterior notch, and a distinct tasciolar bantl, abo\e

which a spiral groove ends in a slight tooth-like or lobe-

hke protrusion at the outer Up. Ancillopsis differs from

Pseudoli\idae by having the spire covered with a glaze

of callus, b\- the absence of a pseudoli\id groo\e, and bv

lacking spiral and axial sculpture. Allmon (1990) showed

that Ancillopsis does not belong with BtiUia Gray in Grif-

fith & Pidgeon, 18.34, in the nassariid subfamilv Dorsan-

inae, where P;ilmer (1937) and Palmer anil Brann (1966)

had placed it, because it lacks the terminal columellar

fold characteristic of Nassariidae. Ancillopsis mav be a

divergent member of the AncUlinae.

The familv Strepturidae Cossmann, 1901, has occa-

sion;illv been subsumeil under, or sviion\iiiized with, the

Pseudohndae (see e. g. Ponder and Waren, 1988). Based

on the tvpe genus Strcpsidtira Swiiinson, 1840 (tvpe spe-

cies: S. turgida (Solander in Brander, 1766), the Strep-

turidae have in common with the Pseudolividae the fol-

lowing characters: spiral sculpture increasing in promi-

nence toward the base, a,xial sculpture most prominent

on upper part of whorl, and presence of small adapical

notch. The Strepturidae differ, however, bv the absence

of a pseudolivid grooxe and labral tooth, and bv ha\ing

the inner shell laver reflected shghtlv o\er the outer lip's

edge, which is therefore smooth and polished. Unhke

most Pseudoh\idae, the Strepturidae have a verv strong

fold on the columella at the entrance of the siphonal

canal. The genus Mclapiiun H. & A. Adams, 18.53, has

often been included in either the Strepturidae (Coss-

mann, 1901) or Pseudolividae (Thiele, 1929: VVenz,

1938-44: Kilbuni, 1989). As discussed later in this paper,

I tentati\elv support Cossmann's (1901) assignment of

Mclapium to the Strepturidae. It this proves to be cor-

rect, then the anatomical studv of Mclapium h\ Kantor

(1991) would implv that the Strepturidae, which are

known mainK' from the Paleogene, are relativelv primi-

tive neogastropods related to the OUvidae.

Several members of the Melongenidae bear a close

resemblance in shell characters to the Pseudoli\idae.

Neogene to Recent genera such as Melongena Schu-

macher, 1817, and Volema Rochng, 1798, share with

pseudolixids a low-spired, indeterminateK' growing shell,

a smooth columella without a basal fold, an adlierent

inner lip, a thick persistent periostracum, and a distinct

adapical (posterior) notch in the outer hp. They differ

from pseudoIi\ids In- lacking a basal groove and corre-

sponding labral tooth as well as in the development ot

spiral sculpture. In melongenids, grooves ending in cren-

ulations at the outer lip increase in prominence from the

shoulder to the base, as in pseudolivids, but spind keels

or strong cords that often bear nodes or spines are most

prominent on the upper part of the whorl.

In the Earlv Miocene (Aquitanian and Burdigahan)

Mclongcna lainci (Basterot, 1825) from southwest

France, the outer lip of the adult shell usuallv has a

downiwardlv convex protnision. In a few indi\iduals, this

protnision, which corresponds to the basal constriction

of the last whorl, bears a very weak, blunt labral tooth

(for further discussion of this species see PevTot, 1927;

Vignon, 1931).

The genus Conmlina Conrad, 1853, contains several

Late Paleocene and Eocene species with a "pseudoLvid"

groove and corresponding labral tooth. These include

the t\pe species, C. minax (Solander in Brander, 1766)

from the Middle and Late Eocene of Europe and C.

annigera (Conrad, 1833) from the Middle Eocene of the

southeastern United States. Like members of the Pseu-

doli\idae, Conudina is characterized bv indeterminate

growth, a shaip-edged outer Hp, adherent smooth inner

lip, and outer-lip crenulations tliat become more pro-

nounced abapiciillv. It differs from Pseudohvidae by the

absence of an adapical notch and bv the presence of two

prominent spiral rows of nodes, and from Eocene pseu-

doli\ids b\' the presence of a parietal rib and a basal

columellar fold. Petuch (1994) interpreted Conmlina as

an ancestor, or an early member of his subfamily Echin-

ofulgurinae of the familv Melongenidae. The earhest un-

doubted Conudina. C praccursor Cossmann, 1902,

dates from the Thanetian stage of the Late Paleocene of

the Paris Basin in France.

Protobusi/con Wade, 1917, mav be an even earlier ge-

nus in this group. The two Late Cretaceous members of

this genus from the Gulf Coastal Plain of the United

States (see Sohl, 1964a) bear a labral tooth at the end

of a groove on the lower part of the last whorl. Sugges-

tions that Protobusi/con (tvpe species: Biisi/con creta-

ccum Wade, 1917) belongs to the TurbineUidae (Bandel,

1993) or Ficoidea (Riedel, 1994) seem far-fetched. Tur-

bineUids (especially Paleogene species) lack the parietal

rib characteristic oi Protohiisi/con. Ficoids, which Riedel

(1994) argues are basal neogastropods, have internal

shells without a labral tooth. The spines or nodes on the

upper part of the whorl of Protobtistfcon indicate that

the shell was probablv external. Like Conudina, Proto-

busycon differs from Paleogene pseucloli\ids by the

presence of a parietal rib and the absence of a posterior

notch. In contrast to the situation in Pseudolividae, axial

sculpture in Protobusijcon extends to the siphonal canal.

Page 58

THE NAUTILUS, Vol. Ill, No. 2

which is very long. Spiral .sculpture gener;dly decreases

in strength adapicallv on the last whorl of Protobusi/con

as it does in pseiidoli\ids, but two widely separated rows

of nodes (one at the shoulder and one just alxne the

tooth-bearing groove) are superimposed on this pattern,

as they are in Comiilina. The single Paleocene species

o{ Protobusijcon (P. judithac Saul, 19S8, from the "Mar-

tinez" stage of Cahfornia) ma\ not ha\'e possessed a la-

bral tooth, but the genus could nonetheless have given

rise to, or be the sister group of Coniulina. In any case,

the labral tooth of this group probalily evolved indepen-

dently from that in the Pseudolixidae.

The genus Lacinia Conrad, 1853, comprises Middle

Eocene shells remarkably similar in form to contempo-

raneous species of the pseudolivid genus Siilcobuccinum

(see below). Like Sulcohucciuiim. Lacinia has a low-

spired, ovate, basally unconstricted, thick, indetermi-

nately growing shell with a distinct adapical notch and

extensive columellar and parietal c;xllus. Lacinia alvcata

(Conrad, 1833) from the Gosport Sand (Middle Eocene)

of Alabama, the type species of Lacinia, is a large species

(maximum height 115 mm) characterized by a ventral

protmsion on the outer lip about one-third the distance

from the base of the shell. This protrusion corresponds

to a very shallow, very broad spiral depression on the

last whorl. Below this protrusion, the outer hp is finely

crenulated. Unhke Paleogene pseudolivids, Lacinia has

a distinct fold at the base of the columella. Moreover,

this spiral sculpture is better expressed on the upper

part of the whorl, where it consists of four broad, round-

ed folds. Lacinia is represented by two species in the

Gosport Sand of Alabama, L. alvcata and the much

smaller L. claibonienesis Palmer, 1937, a species with

axial ribs on the spire whorls. Vredenburg (1922) has

named three species of Lacinia from the Late Eocene

and Ohgocene of Burma and India. The Late Eocene

Nigerian Athlcta lugardi Newton, 1922, the t)'pe o( Lac-

cinum Eames, 1957, is very similar to Lacinia except that

it lacks any trace of spiral sculpture.

Palmer '( 1937) and Wenz ( 1938-1944) assigned Lacin-

ia to the Buccinidae, but the genus could equally well

be included in the Melongenidae. It is even possible that

Lacinia and Laccinum are divergent members of the

Pseudolividae. For the time being, I consider their tax-

onomic placement uncertain.

There is also a considerable resemblance in shell char-

acters between Pseudolividae and the buccinid subfam-

ily Babyloniinae, as exemplified by the Eocene to Recent

genus Babylonia Schliiter, 1838 (type species: B. spirata

(Lamarck, 1822)). Like manv Pseudolividae, Babylonia

is characterized by a thick periostraciun, canaliculate su-

ture, relatively low spire, planar simple outer lip, a pos-

terior notch, a well developed siphonal fasciole, smooth

columella without basal fold, and a basally unconstricted

last wliorl. Babylonia differs from Pseudolividae by hav-

ing the glos.sy inner (adaxial) layer of the outer lip re-

flected over the edge of the outer Up, so that the edge

is smooth and polished rather than shaqi; and by lacking

basal spiral sculpture as well as a pseudolivid groove and

labral tooth.

Members of the buccinid subfamily Pisaniinae have in

common with pseudoli\ids a short spire, adherent peri-

ostracum, sharp-edged outer lip, smooth columella, and

axial sculpture that is most strongly developed on the

upper part of the whorl. Pisaniines differ from pseudo-

hvids bv having the spiral sculpture increasing in prom-

inence from the base to the shoulder (adapicallv) rather

than from the shoulder to the base (aliapically), by hav-

ing the edge of the outer lip ventrallv convex instead of

planar, and by lacking a posterior notch. Some Pisani-

inae, such as the genera Canthanis Boding, 1798, and

Pollia Gray in Sowerby, 1833, have a small labral tooth

at the end of a spiral groove on the lower half of the

last whorl, but this groove is a derived feature in the

Pisaniinae and is therefore convergent with the labral

tooth and groove of pseudoUvids.

Shells of the buccinid subfamihes PhoHnae and Si-

phonaliinae ihffer from those of pseudoli\ids In ha\ing

the edge of the outer hp polished rather than sharp in

the fully grown shell, and usually by having a basal con-

ca\'e sector or sinus on the outer lip. Above this sinus,

the outer lip of these buccinids is mechallv convex, and

there is commonly a broad posterior sinus on the outer

hp above the level of the shoulder Some members of

the Photinae, such as Northia, have a posterior notch at

the upper end of this sinus. The siphonaliine genus

Phoracanthus Cossmann & Martin in Martin, 1914 (type

species: Siphonalia ickei Martin, 1914; Nanggulan beds,

Eocene of Java) has a basal spiral groove encUng in a

labral tooth. Sculpture consists of fine spiral threads

bundled into cords, and of widely spaced axial ribs on

the upper part of the whorl (see Martin, 1914). It is

likely that the labral tooth of Phoracanfluts is convergent

with that of Pseudolixidae. A few species of Photinae

also ha\e a labral tooth. They include members of the

genera Cominella Gray, 1850 (type species. Cominella

maculosa (Maitvn, 1784)) and loscpha Tenison-Woods,

1879 (type species: /. tasmanica Tenison-Woods, 1879).

In Cominella acutinodosa (Reeve, 1846) from Western

Australia, a labral tooth is situated at the end of an en-

larged central cord on the last whorl. In loscpha tas-

manica from southern Australia and /. ^landiformis

(Reeve, 1846) from New Zealand, a blunt labral tooth is

formed at the center of the convex sector of the outer

hp and does not correspond with either a cord or a

groove. These labral protiiisions are therefore not ho-

mologous with those of Pseudohvidae.

Finally, the Muricidae chffer from the Pseudolividae

In having the spiral sculpture increasing in prominence

from the base to the shoulder rather than from the

shoulder to the ba.se, and (in Paleogene forms) by lack-

ing a posterior notch. In early muricids, moreover, axial

sculpture almost always extends from the suture to the

base, and is not confined to the upper part of the whorl.

No muricids have the thick, persistent periostracum t\p-

ical of pseudolivids.

G. J. Vermeij, 199S

Page 59

KEY TO GENERA OF PSEUDOLIVIDAE

la Protocoiuli iiuiltispiral (tliree whorls or more)

2

Ih Prototonch paucispiral (less than three whorls)

3

2a Shell fusiforiii, relati\elv slender (heijiht-to-di-

ameter ratio 2.0); spiral sculpture consisting of

fine cords Fiisopsis

2b Shell inflated (height-to-diameter ratio 1.4 to

1.6); spiral sculpture obsolete Bcnthohia

3a Inner (adaxial) side of outer lip lirate (bearing

spiral riblets) 4

3b Inner side of outer lip smooth or unknown ... 6

4a Shell moderatelv slender (height-to-tliameter

ratio 1.7 to 1.9); axial sculpture present on up-

per part of whorls; pariettd rib absent; abapical

end of columella pointed Lttizia

4b Shell relatixelv broad (height-to-diameter ratio

1.4 to 1.8); axial sculpture absent or confined to

earlv whorls; parietal rib present; abapical end

of columella with distinct fold 5

5a Pseudohxid groove and labral tooth absent; axid

sculpture present on earlv whorls; outer lip

adapicallv extended at matiuit^' Tiiuiuphis

5b PseudoU\id grooxe and labral tooth present (but

obsolete in Macron lividus); axial sculpture ab-

sent; outer hp not adapicallv extended at ma-

turitv Macron

6a Pseudolivid groove situated low on last whorl;

parietal rib present 7

6b PseudohNid groove situated high on last whorl;

parietal rib absent 9

7a Spiral sculpture present on whole surface of last

whorl; axial sculpture present on early whorls,

or ex-pressed as strong growth lines on last

whorl; columellar callus thin, of limited extent

Tcstalliuni

7b Spiral sculpture on last whorl absent; axial

sculpture absent; columellar callus thick, exten-

sive S

8a Parietal rib rotmded, not markedly constricting

adapical end of aperture Psciuloliia

8b Parietal rib lamella-Uke, sharp-edged, separating

long adapical channel from rest of aperture . . .

Fiibncnttnn

9a Siphonal fasciole absent; siphonal notch verv

shallow 10

9b SiphoniJ fasciole present; siphonal notch deep . . 11

10a Suture linear, appressed; base of last whorl con-

stricted; labral tooth obsolete; axial sculpture

present on upper part of whorl Fusulcuhis

10b Suture deeply channeled; base of last whorl not

constricted; labral tooth prominent; axial sculp-

ture usuallv absent Zctnira

11a Shell high-spired (last whorl comprising 60 to

70% of total shell height); pseudolixid groove

indistinct; labral tooth present; last whorl con-

stricted at base; columellar callus thin 12

lib Shell high-spired to low-spired; last whorl usu-

alK not constriited at base; pseudoliNiil groove

and labral tooth distinct; columeUar callus well

developed, thick 13

12a Adapical notch absent; axial sculpture absent or

consisting of short ribs at shoulder . . . Nauiloliia

12b Adapical notch weak but present; axial sculpture

consisting of narrow peripheral band of beads

on last whorl Sulcoliva

13a Deep groo\e situated immediately below sharp-

K' angled sln)ulder; aperture verv wide

Hitbachia

13b Shoulder variably developed, without deep

groove immediatel)' below it; aperture narrower

Stilcohucciniim

Genus Fscudoliva Swainson, 1840

(Figures 10-12)

T\pe species: Biiccinum phimhctnn Chemnitz ( =

Biiccinuui crassiim Gmelin, 1791) by monotypy.

Sj-nonyms:

Gastriciitini Sowerby, 1842 (type species: B. phimbeum Chem-

nitz), non Modeer, 1793, nee Sowerbv. 1S46.

Diagnosis: Shell moderately large, maximum height

50 mm, ovate to globose; protoconch paucispiral; spire

moderatelv low to verv low, last whorl comprising 78 to

90% of total shell height; last whorl rounded abo\'e, not

constricted basallv; spiral sculpture confined to early te-

leoconch whorls, absent on last whorl; axial sculpture

absent; pseudolixid groove situated low on last whorl;

outer lip with distinct labral tooth, wthout basal cren-

ulations; inner side of outer lip smooth; posterior notch

deep; aperture ovate, its height-to-breadth ratio 2.1; col-

umellar and parietal callus moderatelv thick, well de\'el-

oped ventrallw but not extending onto spire; parietal rib

strong, rounded; siphon;il fasciole low, rounded; anterior

notch deep; umbilicus absent.

Stratigraphic and geographic distribution: Early?

Miocene, Madeira; Early Pliocene to Recent, southern

Africa.

Discussion: Although Swainson (1840: 82, 306) pro-

posed the name Pseudoliva for the West African species

we know today as Pseudoliva crassa (Gmelin, 1791),

Pseudoliva was interpreted broadl)' by paleontologists,

who included in it a diverse array of gastropods with a

basal groove and corresponding labral tooth. I follow Kil-

bum (1989) in restricting Pseudoliva to a small group of

warm-temperate to subtropical West African species

characterized by a non-umbilicate, smooth or obs(jletely

spiralK' threaded, ovate shell with a posterior notch, pa-

rietal rib, well developed callus, and smooth inner (ad-

axial) side of the outer lip. Pseudoliva differs from Sul-

eohuecinuin hv ha\ing the pseudolixid grooxe and labral

tooth situated lower on the whorl, b\ the presence of a

rounded parietal rib at the adapical end of the inner lip,

and by being essentiallv smooth, there being no trace of

Page 60

THE NAUTILUS. Vol. Ill, No. 2

axial sculpture. Sulcobuccinum kitsoni (Newton, 1922)

from the Ameld Formation (Middle Eocene) of Nigeria

shows a trace of the parietal ril), and may be transitional

between Sulcobuccinum and Pscudoliia. In the related

genus Fulmcutuin, the parietal rib has become e.xagger-

ated as a plate, which sets a long, channel-like adapical

section off from the rest of the aperture. Pscudoliia dif-

fers from TcstaUium bv lacking spiral sculpture on the

last whorl and b\ not having the last whorl constricted

at the base.

I recognize three species in the genus Pseudoliva (see

Appentlix). The tvpe species, P. crassa, is k-nown only

from the coast of Angola. The largest specimen 1 have

seen (height 49.0 mm) comes from the Baia dos Tigres.

Kensley and Pether (1986) described a very closely re-

lated if not identical form as Pseudoliva lutulenta from

the 50 m level (Earlv Pliocene; J. Pether, personal com-

munication) at Hondeklip, on the west coast of South

Africa. The earliest member ot the genus appears to be

P. orbiffujana Maver, 1864, from deposits of probably

Earlv or Middle Miocene age on Madeira.

Kilbum (1989) svnionvinized SijhanocochUs MeKill,

1903, with Pseudoliva. Its type species, Pseudoliva an-

cilla Hanley, 1859, from theAgulhas Bank off South Af-

rica, differs from Pseudoliva bv its high spire and adap-

ically extended aperture. As discussed below under Ful-

inentuin, I consider Sijlvanocochlis a synonym of Ful-

mc7}fum.

Genus Fulinciitum Fischer, 1884

Type species: Buccinum sepimcntuui Rang, 1832.

Svnonyin:

Sijlvanocochlis Melvill, 190.3 (type .species; Pseudoliva ancilla

Hanley. 1859).

Diagnosis: Shell of moderate size, maximum height

40 mm, ovate; spire moderately low, last whorl compris-

ing 75 to 88% of total shell height; last whorl evenly

rounded above, weakly or not constricted basally; spiral

sculpture usually absent, sometimes expressed as very

weak threads below pseudolivid groove; axial sculpture

absent; pseudolivid groove situated low on last whorl;

outer lip with distinct labral tooth, below which small

crenulations may appear; inner side of outer Up smooth;

posterior notch present; aperture ovate (F sepimentum)

to elongate-ovate (F ancilla). its height-to-breadth ratio

2.4 to 3.0; adapical end of aperture extended as narrow-

channel between outer hp and a prominent, projecting,

plate-hke parietal ridge at posterior end of inner hp; col-

umellar and parietal callus well developed ventrallv, but

not extending onto spire; siphonal iasciole low, rounded,

.sculptured bv five weak spiral threads; anterior siphonal

notch deep; umbihcus absent

Stratigraphic and geographic distribution: Recent,

tropical West Africa and temperate South Africa.

Discussion: Fischer (1884: 632) introduced Fiduicn-

tum as a subgenus of Macron for Buccinum sepimentum

Rang, 1832, a Recent species from tropical West Africa.

Most later authors have treated Ftdmentum as a full ge-

nus or as a subgenus of Pseudoliva. The exaggerated,

plate-like parietal ridge, which sets apart a narrow adap-

ical extension from the rest of the aperture, readil\- chs-

tinguishes Fulmcntum from Pseudoliva. Fidmentum .se-

pimentum ranges from Cotonou (Togo) to southern

Angola. The largest specimen I have seen (height 20.4

mm) is from Pointe Noire, Congo (MNHN).

Melvill (1903: 325) named the genus Sijlvanocochlis

for Pseudoliva ancilla Hanley, 1859, a relatively large

(maximum height 40 mm), high-spired species from the

Agulhas Bank off South Africa. Kilbum (1989) consid-

ered Sijlvanocochlis a synonym of Pseudoliva, correctly

pointing out that spire height in Pseudoliva is quite vari-

able and therefore an unrehable criterion for the rec-

ognition of genera. My e.xamination of Pseudoliva ancilla

prompts me to assign this species to Fulmcntum. Like

F sepimentum, the South African species has a drawn-

out adapical extension of the aperture, set off from the

abapical m;iin portion of the aperture bv a flange-hke

parietal lamella. Fulmcntum ancilla differs from F se-

pimentum by being about hvice as large and bv not being

constricted basally.

Genus Sulcobuccinum d'Orbigny, 1850

Type species: Buccinum fissuratum Deshaves, 1835,

here designated (Figures 1-3).

Synonyms:

Buccinorbis Conrad, 1S6.5 (tvpe species: Buccinum vetustum

Conrad. 1833, subsequent designation bv Cossmann,

1901; Figures .5-7).

Proocoinptus Zinsnieister, 1983 (tvpe species: Molopophortis

howardi Dickerson, 1914, bv original designation).

Calorehama Squires, 19S9 (t\pe species: Pseudoliva dilleri

Dickerson, 1914, by original designation).

Fopenoeinn Squires, Zinsnieister & Paredes-Mejia, 1989 (type

species: Popenoeum nuiritimus Squires, Zinsnieister & Pa-

redes-Mejia, 1989, bv original designation).

Diagnosis: Shell very small to large, maximum height

4.2 to 66 mm, ovate to globose; protoconch paucispiral;

spire relati\-ely high to low, last whorl comprising 63 to

100% of total shell height; last whorl rounded or shoul-

dered above, usually not constricted at base; spiral sculp-

ture variablv expressed; axial sculpture consisting of

eight or more ribs per whorl, confined to part of whorl

above pseudolivid groove, and often forming nodes or

spines at shoulder and at intersections with spiral cords;

axial sculpture confined to spire whorls in some species;

pseudoliviil groove at relativelv high position, not less

than one-fourth from abapic;il end ot last whorl; outer

lip with distinct labrd tooth, below which are three to

fi\-e crenulations corresponding to grooves between basal

cords; iTuier side of outer lip smooth; posterior notch

present; aperture ovate to ovate-elong;ite, its height-to-

l)re;idth ratio 2.0 to 3.8; columellar and parietal callus

\;iri;il)ly expres,sed, often thick and extending onto spire

in huge Eocene species; parietal ridge absent; siphon;il

G. J. Vermeij, 1998

Page 61

Figures 1-3. Suleobuccinuin fissiimtum (Deshayes, 1835); Chalons-sur-\esle, Manie, France; Thanetian; MNHN. Shell height

38 mm.

Figure 4. Hitbnclua raiitirolobac Eta\o Senia, 1979; Paleocene, Colombia; UCMP 16014. Oblique \ie\v of ape.x; total shell height

30 mm.

Figures 3-7. Buccinorhis vctustn (Conrad, 1865); Little Stave Creek, Alabama; Gosport Sand (Middle Eocene); GJ\'. Shell height

36 mm.

Figures 8-9. Sulcoliin tnonilis (Olsson, 1928); Pariiias Sandstone (Earlv Eocene, Pern); PRI 3671. Shell height 17 mm.

Figures 10-12. Psaiiioliin cras.sfl (Gmelm. 1791); West Africa; CAS 063184. Shell height 25 mm.

Figures 13-15. Luizia zcbhna (A. Adams, 1855); Ilha de Luana, Angola; MNHN. Shell height 14.2 mm.

Page 62

THE NAUTILUS, Vol. 111. No. 2

fasciolar ridge present or obsolete, bounded above by

keel when present; anterior siphonal notch usually deep;

umbilicus present or absent.

Stratigraphic and geographic distribution: Late

Cretaceous (Campanian and Maastrichtian) of Europe,

North Africa, southwestern Asia, and western South

America; Early Paleocene to Late Eocene (Danian to

Priabonian) of Europe, western and eastern North

America, South America, North and sub-Saharan Africa;

Earlv Paleocene of India; Late Eocene to Early Oligo-

cene of Japan; PEarly Oligocene of Peru.

Discussion: The taxonomy, relationships, and generic

nomenclature of Late Cretaceous and Paleogene pseu-

dolivids remain matters of controversy and imcertainty.

All species of this age are characterized by a pseudohvid

groove that is situated at a relatively high position on the

whorl, and by the absence of a parietal rib. In these

important respects, they differ from the type genus

Pscudoliva. In addition. Cretaceous and Paleogene pseu-

dolivids usually show traces of iL\ial sculptiue, although

it may be confined to apical whorls, especiallv in Eocene

species. There is great variation in the degree of devel-

opment of spiral and axial sculpture, in the presence or

absence of a siphonal fasciole, in the height of the spire,

and in the development of a sulisutural channel. Many

species have nodes or even spines at the shoulder or at

the adapical ends of axial ribs, but in some species the

upper part of the whorl is roimded and therefore un-

shouldered, and nodes are not developed.

Several authors have attempted to subdivide this

group into genera, subgenera, or species groups. Among

Paleocene and Eocene species with axial sculpture on

the last whorl, Adegoke (1977) recognized but did not

name a group of Paleocene species based on Pscudoliva

rogersi Adegoke, 1977, from the Ewekoro Formation of

Nigeria. It is characterized by reticulate sculpture, in

which several rows of nodes form where strong spiral

cords intersect axial riblets. Zinsmeister (1983) proposed

the genus Pegpcomptus for a very similar species, Mol-

opophonis howardi Dickerson, 1914, from the Santa Su-

sana Fomiation of southern Califcjrnia and the Sepultura

Formation of Baja CaUfornia Norte, Mexico. Mv exam-

ination of this Paleocene species shows that it lacks an

umbilicus, whereas Pseudoliva rogersi has a wide, deep

umbilicus (Adegoke, 1977).

Stjuires (1989) distinguished a second group of axiallv

ribbed Paleocene and Eocene pseudoli\ids as his new

genus Calorebaina, for which he chose Pscudoliva dillcri

Dickerson, 1914, as type. He characterized Calorchama

as having an angulated, noded shoulder, shallow ap-

presscd suture, pseudolivid groove situated relativeh'

high on the last whorl, and variably developed spiral

sculpture that coarsens toward the base. Squires includ-

ed in this group two species (C unicarinata and C. fuh-

erculifcra) from the Paleocene of the Gulf C(xistal Plain

of the United States, as well as five subspecies of C

dilleri from the Eocene of western North America. Kase

(1990) added his new speiies C. crctacca from the Late

Cretaceous (Campanian to Early Maastrichtian) of Ja-

pan. He included this species in Calorchama because of

its resemblance to C. dillcri inomata, which like C. crc-

tacca lacks shoulder nodes.

At least two of the characters of Calorchama men-

tioned by Squires (1989) are not generically diagnostic.

All pseudoli\ads have spiral sculpture that coarsens to-

ward the liase, and a nodose shoulder, which occurs in

man\' Late Cretaceous and later pseudoli\'ids, is often

variably expressed during ontogeny (Squires, 1989). A

third character, the appressed suture, occurs in Pego-

coniptus as well as in several slender species of the Pscu-

doliva kocucni group (see below), in which the shoulder

is not nodose.

Still another group that was recognized in this early

welter of pseudolivids was Popenocum (Squires et al.,

1989), This genus, tvpified bv P maritimus, was char-

acterized as having taliulate whorls, sul)sutural channel,

axial ribs terminating as tubercles at the shoulder, a rel-

atively thick columellar callus, and a well developed si-

phonal fasciole. Besides the t\pe species and its subspe-

cies P. III. hajaciisis. Squires et al. (1989) included a wide

variety of Cretaceous to Eocene species: P. hocascrpcii-

tis, P. chavani (GUbert, 1973, not Tessier, 1952; see be-

low), P. mutahilis and its subspecies wood.si, P. prima, P.

robustuin, P.scaliiia. and P. subcosfata (see Appentlix 1).

Kase (1990) subsequently assigned the Cretaceous .spe-

cies P. praccursor and P. zittcli to Popenocum. Squires

et al. (1989) explicitly excluded P. iiuitabilis douvillei

from Popenocum, apparently because of the extraordi-

nary development of callus on the spire of that large

Early Eocene form from Peru. My examination of the

holotypes of Pscudoliva mutahilis douvillei and P. m.

U()()f/.s(' reveals broad similarities in shell shape and in

the development of axial sculpture, and supports Ols-

son's (1928) suggestion that these two forms are closely

related to each other and to P. m. mutahilis. Pseudoliva

mutahilis douvillei has more prominent spines, which

are directed abaxiallv. than does P. m. wood.si. whose

spines are adapicalK' directed.

Clibert (1973) defined an informal group of axiiilly

ribbed Paleocene pseudolivids as the group of Pscudo-

liva kocneni. Members of this European group are char-

acterized by veiy small, slender shells with a linear rath-

er than canahculate suture. Most species, including P.

kocneni, P. elisae, and P. tcimicostata, lack .shoulder tu-

bercles, but P. curvncostata does possess tubercles (see

C;libert. 1973). Some of these species mav be .smiountos

of each other, and some may pro\e to belong to Fusop-

sis, which is distinguished from other axially ribbed Pa-

leogene pseudolivids bv having a multispiral rather than

paucispiral protoconch.

Manv Late Cretaceous and Paleocene pseudolivids

with axial sculpture on the last whorl were left out of

these formal and informal groups. Some species com-

bine features of several nominal groups. Buecinum /i.s-

surutum I^eshaves, 18.35, from the Late Paleocene (Tha-

netian) of the Paris Basin, for example, has a canaliculate

suture like that n{ Pojiciioeum. but unlike Popenocum it

G. J. Vermeij, 1998

Page 63

has a rounded shoulder lacking nodes or tnliereles. Spi-

ral sculpture appears to be confined to the area helovv

the pseudolivid grt)ove, and is expressed at the edge of

the outer lip as three crenulations. A siphontil tasciole is

absent, as in Calorebamo dilleri kirln/i and Puiyura pus-

ilia. Biicchmm lauduncnsis Defrance, 1826, also from

the Thanetian of the Paris Basin, is ven.- similar to B

fissiiratiun, liut its sutin-e is appressed rather than can-

aliculate, and there is a low, distinct siphonal tasciole.

Moreover, the nominal groups differ in characters that

\ar\' intraspecificalK and even during individual ontog-

eny. Wrigley (1941), for example, argued con\incinglv

that a single Late Eocene species from northwestern

Europe (Germany, Belgium, and England) is represent-

ed bv a continuous series of forms between the types

that have been c;illed Piiijiura nodtilusa. in which eight

to ten axial ribs per whorl are enlarged by two rows of

nodes, and P. pusilla, in which the axial riblets are

smooth (see Beyrich, 1854). This species also varies in

the expression of spiral sculpture below the pseudolivid

groove (Wriglev, 1941). In manv species, spiral sculpture

covers most or all of the surface of the last whorl in

ontogeneticallv voung individuals, but it becomes re-

stricted to the base in ontogenetically older ones. When

describing Psciidolha lihi/ca, Quaas (1902) noted that

young, stronglv sculptiued slender, high-spired individ-

uals have a quite different appearance from larger in-

dividuals of the same species, in which the lateral profile

of the relatively low spire is concave. Squires ( 1989) ob-

served ontogenetic changes in spiral sculpture and in the

presence of a subsutural sweUing in species that he as-

signed to Calorchama.

It is therefore difficult to delineate well defined

groups based on such characteristics as spire height, ex-

pression of spiral and ;L\ial sculpture, and presence of

nodes among Cretaceous and Paleogene pseudolivids

with a.\ial sculpture. Although several hneages may be

represented in this group, I prefer to treat the earlv,

a.xially ribbed pseudolivids with a paucispiral protoconch

as members of a single genus-level group.

In some Paleocene and Eocene pseudolivids, axial

sculpture is confined to the spire whorls or is occasion-

ally absent. Palmer (19.37) used the taxon Buccinorbi.s

Conrad, 186.5, to encompass this group. Like Cossmann

(1901) and Adegoke (1977), she regarded Buccinorbis as

a subgenus or section o( PsciidoUva. Fischer (1884) and

Thiele (1929) .sviionvinized Buccinorbi.s with Pscudoliia.

whereas Squires (1989) and Vermeij and DeV'ries (1997)

considered it to be a distinct genus. Conrad (1865:21)

introduced Buccinorbis as a subgenus of Sulcobuccinum

without diagnosis or comment. He included in it all east-

em North American pseudolivids described up to 1865,

incluiling strongly sculptured species like Pscudoliva

tubcrculifcra as well as the nearly smooth Buccinum ic-

tustum. Cossmann (1901) designated B. vetustum Con-

rad, 1833, the last species Conrad (1865) listed as be-

longing to Buccinorbis, as tvpe species.

The only character distinguishing Buccinorbis from

most other early pseudohvids is the absence of axial

sculpture on the last two teleoconch whorls. Species vary

in the presence or absence of spiral sculpture on the

upper part of the whorl, in spire height, in the presence

or absence of an imibilicus, and in the development of

columellar and parietal callus deposits. As pointed out

by Olsson (1928), Palmer (1937), and Gardner (1945),

these attributes vars' during ontogenv, just as the ex-

pression of axial sculptiue does. Ontogenetically young

shells tend to be high-spired, to be axially sculptured, to

have spiral cords or threads distributed over the whole

surface of the whorl, to have an open umbilicus, and to

have a relativelv limited callus. The loss of axial sculpture

likelv occurred in several Paleocene lineages, and by it-

self does not constitute sufficient grounds for distin-

guishing Buccinorbis from other earlv pseudolivids with

a paucispiral protoconch, high position of the pseudoli-

vid groove on the whorl, and absence of a parietal rib.

I therefore treat most of these species as belonging to a

single genus.

The oldest av;ulable generic name for this group of

pseudohvids is Sulcobuccinum d'Orbigny, 1850. This

name has been overlooked by most authors, perhaps in

part because Fi.scher (1884), Cossmann (19(3l), Thiele

(1929), and Wenz (1938-44) synonymized it with Pseii-

doliia without iliscussion. D'Orbigny (18.50:.303) com-

pared his genus with Buccinanops d'Orliignv, 1841 (Nas-

sariidae, Dorsaninae; see Allnion, 1990), and listed three

species as belonging to it: Buccinum fi.ssuratum Deshay-

es, B tiara Deshaves, and B. scmico.statum Deshayes.

Among these, B. scniico.statum is a junior synonym of

Buccinum lauduncnsc Defrance (see Gilbert, I960; Vil-

latte, 1970), and B. tiara is a junior synon)an of Stni-

thiolaria prima Defrance (Villatte, 1970). Coquand

(1862) added a fourth species, S. michclini; and Conrad

(186.5) introduced Buccinorbis as a subgenus of Sulcob-

uccinum. Neither d'Orbigny (18.50) nor any subsequent

author designated a type species of Sulcobuccinum. Be-

cause Buccinum fi.ssuratum Deshayes, 18.35, was the first

species listed by d'Orbignv ( 18.50), I here designate it as

the tvpe species of Sulcobuccinum d'Orbigny, 1850.

Several genera have in common with Sulcobuccinum

the presence of axial sculpture on the last whorl. These

include Bcnthobia. Fusopsis, Fusulculus, Hubachia, Lui-

zia, and Sulcoliva. As thscussed further under Fusopsis,

Sulcobuccinum differs from that Paleocene group mainly

bv having a paucispiral protoconch rather than a multis-

piral one. Most species of Sulcobuccinum are not basally

constricted, in contrast with Fusopsis. The study of pro-

toconchs may reveal a diversit)' of types among species

currently assigned to Sulcobuccinum. In addition, fur-

ther work mav show that Fu.sopsis is conchologicallv in-

distinguishable from Bcnthobia. Sulcobuccinum differs

from Fusulculus bv the presence of a posterior notch on

the outer hp and bv' lacking the basid constriction of the

last whorl. It differs from Luizia bv the absence of rib-

lets on the inner side of the outer hp, and from Sulcoliva

by having a tleep pseudolivid groove and bv not having

axial ribs confined to a narrow peripheral band.

In reviewing the species here assigned to Sulcobuc-

Page 64

THE NAUTILUS, Vol. Ill, No. 2

cinum, I have uncovered a case of homonyiny. Gilbert

(1973) named Psendoliva chavani from the Calcaire de

Mons (Danian, Earlv Paleocene) of Belgium. Unfortu-

natelw this name i.s preoccupied by Psendoliva (Bucci-

norhis) chavani Tessier, 1952, from the Maringot de

Balling (Paleocene) of Senegal. I therefore rename Gli-

berts species Sulcobuccinum multinodulosiim, new

name, to emphasize the large number (seventeen to

nineteen) of axial riblets forming nodules below the su-

ture.

Nearly all Late Cretaceous species of Sulcobuccinum

are known from the Old World (see Appendix 1). These

include S. aaronsohni and S'. moabiticum trom the Sen-

onian oi Palestine, S. cretaccum from the Campanian to

Early Maastrichtian of Japan, S. praecursor And S. zittcli

from the Senonian of Hungary, S. sofegginense, S. libi/-

cum, and S. subcostatiim paucicostatum irom the Maas-

trichtian of North Alrica, and S. ambiguiim irom the

t)pe Maastrichtian in the Netherlands and perhaps the

Maastrichhan of Tunisia (for species accounts see Quaas,

1902; Per\inquiere, 1912; Blanckenhom, 1927; Rossi

Ronchetti. 1959; Kase, 1990). Although I ha\e not ex-

amined these Late Cretaceous species, the differences

among them seem slight, and there may be many fewer

species than there are av;ulable names. The onlv New

World members oi Sulcobuccinum irom the Late Cre-

taceous are Olsson's (1934) Psendoliva sp. from the

Maastrichtian of northern Peru, represented by a single,

poorly preserved specimen, and Muniz's (1993) Psen-

doliva bcllccompta. a cancellate species from the Gra-

mame Formation (Campanian) of northeastern Brazil.

Oddly enough, the genus is unknown from the Creta-

ceous of sub-Saharan Africa, Madagascar, and North

America. All Cretaceous species have a relativelv high

spire, the last whorl comprising 80% or less of the total

shell height, and all ha\e well developed axial sculpture

on the spire whorls as well as the last whorl.

During the Paleocene, the genus Snlcobnccinnm was

widelv distributed in warm seas, with an apparent center

of diversity around the North Atlantic (see Appendix 1).

Species such as S. koencni from Denmark, S. multino-

dulosum from Belgium, S. rosenkrantzi from Austria,

and S. minntissimum from Morocco are high-spired, ax-

ially ribbed, very' small forms not exceeding 7 mm in

shell height. By contrast, the European S. prima reaches

a height of 65 mm and has a spinose shoulder.

Several Old World Paleocene species show trends to-

ward a reduction in iLxial sculpture. One such group of

species is represented in the Ewekoro Formation of Ni-

geria by S. adelekei, S. f. funkcanum, and S. f. omatnm.

Adegoke (1977) characterized this group as high-spired,

spirally sculptured, and having a narrow elevated callus

and a rounded rather than shouldered upper part of the

whorl. These Paleocene species are small (maximum

height of S. adelekei, USNM 174S53, 12.0 mm). In the

Ameki Formation (Middle Eocene) of Nigeria, this

group is represented bv the larger S. kitsoni (maximum

height 47.4 mm). This is a high-spired .species (last whorl

comprising 70 to 80% of total shell height) with a pari-

etal .swelling at the adapical end of the inner hp. Related

Eocene species include S. thielei and S. Icutucini from

Namibia and S. eschi from Cameroon. Bohm (1926) de-

scribed S. thielei as having an umbihcus, a moderatelv

high spire (last whorl comprising 84%- of total shell

height), relatively large size (maximum height 50 mm),

and spiral sculpture persisting on the last whorl onlv

near the suture and below the pseudolivid groove. Be-

sides being slightlv lower-spired, S. thielei appears to be

very similar to S. kitsoni and probablv' represents die

same species, which would then take the name S. kitsoni

(Newton, 1922).

A second group in which axial sculpture has become

reduced is represented in the Ewekoro Fonnation of

Nigeria by S. guineense. This species has a minute, very

low spire, a well developed callus extending on the spire,

a distinct siphonal fasciole, and axial sculpture consisting

of fifteen to sixteen ribs on spire whorls but absent on

the last two whorls. The slightlv older S. chavani from

the Danian (Earlv Paleocene) of Senegal is verv similar,

but its axial sculpture extends to the last whorl ( see Tes-

sier, 1952). Snlcobnccinnm gnincense and the S. adelekei

group represent two apparently independent transitions

from typical Snlcobnccinnm in which axial sculpture con-

tinues to adulthood, to the Bnccinorbis phenotxpe, in

which the last one or two whorls are free of axial sculp-

ture.

Eocene representatives oi Sulcobuccinum were highly

diverse in iorm and size. Strongly tuberculate or spiny

species include S. mntabile and its subspecies S. ;;?.

woodsi and S. m. donvillei from the Early and Middle

Eocene of Peru (see Olsson, 1928; Squires ct al. 1989)

and S. michclini from the Middle Eocene (Suessonian)

of North Africa (ior discussion see Salvan, 1954). SuKill,

slender, axiallv ribbed species include S. hussoni from

the Early Eocene oi southern Tunisia (see .Albanesi &

Busson, 1974) and S. pusilhtm from the Lattoriian (latest

Eocene) of northwestern Europe. The name Pnrjntra

pusilla Bevrich, 1854 (pp. 14.3-144) has page priority

over the sviionvmous P. nodulosa Bevrich, 1854 (p. 144),

and must thereiore stand as the vahd name for this

small, sculpturally variable species. Relatively low-spired

species with a tuberculate shoulder and variablv devel-

oped axial sculpture on the last whorl include S.

schweinfurthi from Cameroon (see Oppenheim, 1904)

and the forms that Squires (1989) subsumed under the

name Calorebama dillcri from the Earlv to Late Eocene

(Ypresian to Priabonian) of western North America.

These western North American forms overlap greatly in

form, sculpture, and stratigraphic distribution (Squires,

1989), and mav well represent a single, variable, strati-

graphicallv long-lasting, widelv distriliuted species whose

earliest v;ilid name is Sulcobuccinum lineatum (Gabb,

1864). A large (perhaps 50 mm high), axiallv ribbed spe-

cies without a tuberculate shoulder from the latest Eo-

cene (Lattorfian) of Helmstedt, Germanv, was described

as Psendoliva nubs on the basis of one poor specimen

bv von Koenen (1889). The (hstinctive Middle Eocene

.S eoronaria from Peni is characterized bv large size

G. J. Vermeij, 1998

Pase 65

(height 50 mm), a high spire, and a nodose, shaqoly

shouldered last whorl (see Olsson, 1930). Several species

groups ot Sulcohucciniim lack ;L\ial sculpture on the last

one or more teleoconch whorls, and were therefore pre-

viously assigned to the genus, subgenus, or section Buc-

cinorhis. Be.sides S. hitsani, which I discussed above, this

group includes man\ species in North and South Amer-

ica, Europe, and one in Japan.

The Old World group of slender species ot the Biic-

ciuorhis phenotype, tvpified by S. adclckci and S. kit.wni

(see above), is represented in the early Eocene ot the

Gulf Coastal Plain of the United States by S. ostronipis

from Alabama and b\' the Texan sub.species S. o. pauper.

These forms have a high spire (last whorl comprising

67% to 76% of total shell height), a narrow aperture,

and an obsolete pseudoli\id groo\'e. The last whorl is

rounded abo\e and e\enK' tapered below, and lacks an

umbihcus and siphonal tasciole. It is possible that some

of these characters are those of juvenile specimens, but

another possibilit\- is that the S. ostranipis group is ple-

siomoiphic with respect to features ot the spire and

base.

The species group of S. vctustiim (tspe species of Biic-

cinorbis) occurs on the Gulf Coastal Plain of the south-

eastern United States (see Palmer, 1937; Dockeiy, 1980).

Sulcobucciiuim vvtiistiim, from the Gosport Sand (Mid-

dle Eocene) of Alabama, has an appressed suture, closed

umbihcus, low spire (last whorl comprising more than

90% of shell height), and obsolete spiral cords on the

upper part of the last whorl. Grooves betv\'een fi\'e cords

belou' the pseudoli\id groo\e end in shaq^ crenulations

at the outer hp. In my material from Little Stave Creek,

Alabama, most indi\iduals ha\e a relatively restricted

callus, but one large mdivitUuil (height 34.1 mm) has the

spire nearK' obliterated h\ a massive callus deposit. Stil-

cobiiccinuiii caiinatum and S. pcrspcc'tivum are related

species with an open umbilicus. Although Palmer (1937)

regarded them as forms of vcttistum, mv examination of

this group reveals subtle but consistent differences. Sul-

cobticcinuiu carinatitiu from the Stone City and Cook

Mountain Formations of Texas (Middle Eocene) and S.

pcrspectivum from the Moodvs Branch Formation of

Mississippi (Late Eocene) are somewhat higher-spired

(last whorl comprising 81 to 83% ot shell height) than

is S. vcfustum. The last whorl ot S. caiinatum is some-

what less slender (whorl diameter-to-whorl height ratio

0.93) than that of S. perspectivum (ratio 0.86), but the

sheO of S. caiinatum has a somewhat more slender ap-

pearance because the voidest point of the last whorl lies

at a relativelv lower position and because the shoulder

is less wide than the periphery in S. carinatuni Other

species in this group include S. linosum, S naunm. and

S. santandcr from the Gulf Coastal Plmn, and the non-

umbihcate S. obtusum from France. Docken,- (1980)

speculates that the thick callus deposits of S. santandcr

from the Bashi Marl Member of the Hatchetigbee For-

mation (eark- Eocene) of Mississippi indicate a gravita-

tionallv stable shell in a wa\e-swept, shallow-water en-

\ironment. Similar callus deposits are developed in co-

occurring species of AnciUopsis and in several solutid

gastropods.

Vermeij and DeViies (1997) reviewed a group of large

(height up to 50 mm), nonumbilicate, smooth species

from Peru, m which the outer hp is adapicall)- extended

beyond the tip of the spire in mature shells. This group,

\\hich niav have extended into the Earlv Oligocene (T.

J. DeVries, personal communication), includes P. retu-

sum and its subspecies S. r samanicum and S. r man-

corensc, as well as S. vientoensc and a form comparable

to S. retusum from Colombia and Curafao (see also

Clark & Durham, 1946: Jung, 1974).

One species of the smooth Buccinorbi.s phenotvpe is

known from Japan. Although originallv described as a

species of Oiiliaulax bv Nagao (1924), the affinities of

S. japonicum with Pscudoliva were recognized bv Nagao

(1928) by virtue of the presence of a pseudoli%id groove.

Honda (1991, 1994) records this species, in which axial

sculpture is absent and spiral sculpture is confined to

the zone below the pseudoli\id groove, from Late Eo-

cene (Takashiman and Okinoshiman) and Earlv Ohgo-

cene (Mazean and Funazuan) deposits.

Genus Hubaclua Etayo Senia, 1979

(Figure 4)

Type species: Htdmchia ramirolobac Etavo Sema,

1979, bv original designation.

Diagnosis: Shell medium-sized, maximum height

about 30 mm, o\'ate; protoconch unknown; spire mod-

erateK' low to x'erv low, last whorl comprising 81 to 100%

of total shell height: last whorl marked b\' low, sharply

angulate shoulder, immediately below which is a deep

spiral groove: between upper groove and pseudohvid

groove, last whorl evenlv rounded; base of last whorl not

constricted: spiral sculpture consisting of fine spiral

threads on spire whorls, and of eight weak cords below

pseudohvid groove on last whorl; axial sculpture con-

fined to spire whorls, consisting of about 15 irregular,

rounded, low ribs; pseudohvid groove situated high on

last whorl, just below its middle; outer hp unknown; ap-

erture broadlv ovate, its height-to-breadth ratio approx-

imatelv 2.0; columellar and parietal callus thick, well de-

veloped ventrallv, but not extending onto spire; umbili-

cus absent.

Stratigraphic and geographic distribution: Paleo-

cene, Colombia.

Discussion: Etavo Sema (1979:37) introduced Huba-

clua as a monoKpic genus of the familv Olividae from

the Paleocene of Colombia. He noted that Hubaclua cht-

fers from the olivid genus Olivancdlaiia d'Orbignv,

1839, by having axial sculpture on the spire whorls and

by having basal cords on the basal half of the last whorl.

Etavo Sema (1979) iilso noted the presence of a pseu-

dohvid groove, and remarked on the similariu of Hu-

bachia to Pscudoliva, from which it differs bv the pres-

ence of axial sculpture on the spire whorls.

Page 66

THE NAUTILUS, Vol. Ill, No. 2

Poor preservation of the available specimen.s pre-

cludes detailed comparison with other genera, and forc-

es me to consider the assignment and status o( Huhachio

as provision;il. The verv deep spiral groove immetliatelv

below the sharply angled shoulder distinguishes Huha-

chia from other pseudoli\'id genera. Hubachia has an

unusually broad aperture and low spire, antl has a round-

ed periphery well below the shoulder. It superficially re-

sembles some species ot Siilcohucrimiin in having axial

sculpture confined to the spire whorls, but tlixerges from

species of Sulcobuccinum in which axial sculpture is ab-

sent on the last whorl by the shaip shoulder and accom-

panving groove.

Hubachia differs from Sitlcobiicciiuiin by ha\'ing axial

sculpture confined to the early teleoconch whorls, by the

exceptionally low spire, and by the presence of a very

deep groove just below the sharply angulate shoulder

The periphery of the last whorl is evenly rounded in

Hubachia. The aperture of Hubachia is relatively very

broad compared to that in species of Sulcobuccinum and

Buccinorbis. Hubachia thffers further from Buccinorbis

by the character of the shoulder and its accompanying

groove.

Genus Benthobia Dall, 18S9

Type species: Benthobia tnjonii Dall, 1889, by origi-

nal designation.

Synonym:

Nux Barnard, 1960 [Type species: Nux alabaster Baniard, 1960

(= Benthobia trieimia Dall, 1SS9)].

Diagnosis: Shell small, maximum height 13 mm, thin;

prf)toconch multispiral; spire moderately low, last whorl

comprising 75 to 77% of total shell height; last whorl

inflated, evenly rounded above, not constricted basally;

spiral sculpture obsolete; axial sculpture consisting of nu-

merous very fine, short riblets confined to subsutural

ramp and shoulder; pseudtjlivid groove very shallow, sit-

uated just below middle of last whorl; outer lip with

obsolete labral tooth; inner side of outer lip smooth; pos-

terior notch absent; aperture ovate, its height-to-breadth

ratio 2.0 to 2.3; columellar and parietal callus thin, of

very fimited extent; parietal ridge absent; siphonal fas-

ciolar ridge absent; anterior siphonal notch obsolete;

umbilicus absent.

Stratigraphic and geographic distribution: Recent,

bathyal and abyssal eastern and western Atlantic Ocean

from 38°N to 33°S; southwestern IndiaTi Ocean; off New

Zealand; 800 to 4380 m.

Discussion: Dall (1889:131) originally placed his new

geiHis Benthobia in the family Cancellariidae because of

its superficial resemblance to Adnu-tc Kroyer in MoUer,

1842; but he ex|:)ressed doubts about this assignment in

view of the absence of an:itomic;il intoniiiition. The ab-

sence of columellar folds ;ilso inijilicd lliat Beulhohia

was unlikely to belong to the (Cancellariidae. Bouchet

and Waren (1985) figured the niduhi of Benthobia ;nid

transferred the genus to the olivid subfamily Pseudoli-

vinae. Kantor's (1991) anatomical work subsequently

provided further support for this reassignment.

As presently understood, Benthobia is a monotvpic ge-

nus of Recent bathytil and ab\ssal gastropods. Anatom-

ically, Benthobia is plesiomorphic in having the radular

sac situated at the base of the proboscis and b\' retaining

the connection between the radular retractor muscle

and columellar muscle (Kantor 1991).

Among living Pseudofividae, Benthobia is unique in

having a multi-spiral protoconch, which Bouchet and

Waren (1985) link to planktotrophic development. This

character is shared with the Paleocene genus Fttsopsis.

Benthobia and Fu.sopsis have many features in common,

and further work may indicate that Fusopsis represents

an early appearance of Benthobia. Both genera have ax-

ial sculpture confined to the upper part of the whorl.

Fu.sop.'ii.s differs from Benthobia mainly by having a

shghtly constricted base.

Another similar genus is Fu.sulcuhi.s. which hke Ben-

thobia has a very shallow anterior siphonal notch Both

genera lack a posterior notch and parietal rib. Fiisulculus

differs from Benthobia by being basally constricted. The

spire of Fu-mlcuhts is higher, and spiral sculpture is well

developed on the base. Fu.sulcuhi.s bears a small but dis-

tinct labral tooth as well as se\eral crenulations below

the tooth on the outer lip, whereas in Benthobia the

tooth is obsolete and crenulations are absent.

The genus Zemira resembles Benthobia in lacking a

fasciole and basal constriction and in having a very sh;il-

low anterior siphonal notch. It differs by having well de-

veloped spiral sculpture, bv possessing a distinct labral

tooth, and (in the tvpe species) bv lacking anv trace of

a\i;il sculpture.

Genus Fu.sopsis Ra\ni, 1939.

Type species: P.seudoUva (Fu.sop.si.s) canaUfera Ra\Ti,

1939.

Diagnosis: Shell ven,' small, maximum height 3.5 mm,

relatively high-spired; last whorl comprising 70% of shell

height; protoconch multispiral; shell fusiform, constrict-

ed at base of last whorl; teleoconch whorls convex, sep-

arated by deep, cauidiculate suture; spiral sculpture con-

sisting of 10 threads on last whorl; axial sculpture con-

sisting of 12 riblets on upper part of last whorl; pseu-

doli\id groove situated just below middle of last whorl,

forming small labral tooth at edge of outer lip; innbilicus

lacking (surmised from description of tvpe species bv

Ravni, 1939).

Stratigraphic and geographic distribution: Paleo-

cene of Demnark.

Discussion: I^avii (1939:76) introduced Fti.sop.si.s as a

subgenus of Pseudoliia in the course of describing F«'»-

(lohia catudijera from the Paleocene of C'openhagen,

Denmark Although he did not provide a cfiagnosis for

the subgenus, he nuule it clear that Fu.sop.si.s differs from

G. J. Vermeij, 1998

Page 67

Psciidoliva s.s. bv having a multispiral rather than a pau-

cispiral protoconth. Ravn (1939) was apparently un-

aware of the genus Bentliohia. whicli hke Fiisopsis has

a multispiral protoconeh, hut which was then classified

in the Cancellariidae.

Without the protoconeh, Fii.sopsis would be indistin-

guishable from other relativek high-spired species of

small, Paleocene pseudolivids that 1 assign here to the

genus Sidcobiiccinum. In fact, Ravn (1939) pointed out

that PscudoUia hoencni Ra\Ti, 1939, which co-occurs

with Fusopsis canalifcra. is transitional to more typical

"Pscudolivci" (= Sidcohuccinitiii as used in this paper).

Fusopsis canalifcra differs from most species of Sitlco-

buccinum by being basally slightly constricted. It is pos-

sible that several species here tentatively assigned to Sid-

cobuccimtm will prove to belong to Fusopsis, which, as

discussed under Benthobia, mav itself fall as a junior

synon\in of Benthobia.

Genus Sulcoliva, new genus

(Figures 8-9)

Type species: Pscudoliva monilis Olsson, 1928.

Diagnosis: Shell moderately small, ma.ximum height

27.4 mm, narrowly ovate to biconic; protoconeh un-

known; spire high, last whorl comprising 59 to 75% of

total shell height; sutures appressed; rounded above,

slightly or not constricted basallv; spiral sculpture con-

sisting of 14 to 16 weak threads below pseudoUvid

groove on last whorl; axial sculpture consisting of 20 to

25 low, weak beads on periphery of last whorl, not visible

on spire whorls; pseudolivid groove at high position on

last whorl, expressed as stepwise basal narrowing rather

than as a groove; outer lip with small labral tooth; pos-

terior notch weak; aperture elongate-ovate, its height-to-

breadth ratio 2.4 to 3.0; columellar and parietal c;illus of

very limited extent; siphonal tasciolar ridge low, round-

ed; anterior siphonal notch deep; umbilicus absent.

Stratigraphie and geographic distribution: Earlv

to Late Eocene, Peru.

Included species: Pscudoliva monilis Olsson, 1928;

Pscudoliva modcsta Olsson, 1928.

Discussion: I propose the new genus Sulcoliva for

tvvo unusual pseudolivids from the Eocene of Peru de-

scribed bv Olsson (1928). Sulcoliva resembles Sulcobuc-

cinum, but is distinguished by the high spire and by the

presence of a peripheral band of fine beads. In Sulco-

buccinum. axial sculpture, when present, normally ex-

tends from the suture to a point above the pseudolivid

groove; whereas in Sulcoliva the axial riblets are con-

fined to the periphery and cannot be seen on the spire

whorls. Moreover, whereas the pseudohvid groove in

Sulcobuccinum is deep, that of Sulcoliva has an adapical

but not an abapical edge. The groove is therefore more

like a step, a sudden, very shght abapical narrowing of

the whorl at or just below mid-whorl height. Sulcoliva

also resembles the Recent genus Naudoliva. Both gen-

era have a high-spired shell with an ill-defined pseudo-

livid groove and a distinct labral tooth. Naudoliva chffers

from Sulcoliva by lacking axial sculpture and by the ab-

sence of a posterior notch.

The genus Sulcoliva contains two species from the

Eocene of northern Peru. These are S. monilis. the tvpe

species, from the Parinas Sandstone (Early Eocene), and

the higher-spired, somewhat larger S. modcsta from Ols-

son s (1930) Talara Formation (Late Eocene, Bartonian).

Genus Naudoliva Kilburn, 1989

Type species: Naudoliva caitlinac Kilburn, 1989, by

original designation.

Diagnosis: Shell thin, ovate, of medium size, maxi-

mum height 29 mm; protoconeh paucispiral; spire rela-

tivelv high, last whorl comprising 64% to 69%- of total

shell height; suture appressed or impressed; last whorl

with concave subsutural ramp, a rounded or weakly an-

gulated shoulder, and an unconstricted base; spiral

sculpture consisting of verv fine threads on spire whorls

and on base of last whorl; axial sculpture absent {N. cait-

linac) or consisting of short, rounded ribs at shoulder

above peripherv- {N. vorstcri); outer hp planar, with long

labral tooth; inner side of outer fip smooth; posterior

notch absent; aperture elongate-ovate, its height-to-

breadth ratio about 2.7; columella and parietal callus

thin, not extensive; columella without fold at base; pa-

rietal rib absent; siphonal fasciole low, rounded, sculp-

tured with verv' fine spiral threads; umbihcal slit ver\'

narrow or absent; anterior notch deep: periostracum ab-

sent.

Stratigraphie and geographic distribution: Late

Miocene, Trinidad; Recent, Inchan Ocean coast of South

Africa.

Discussion: Kilbimi (1989) proposed Naudoliva to ac-

commodate N. caitlinac, a distinctive pseudohvid char-

acterized by a high spire, small umbilical slit, and very

fine spiral but no axial sculpture. The shell has an un-

usually long labral tooth, situated at the end of shallow,

broad pseudolivid groove on the lower third of the last

whorl. There is no periostracum. Kilburn noted that the

radula has a unicuspidate lateral plate, as in Benthobia

(Kantor, 1991), and that the head tentacles are bilobed,

as in Luizia zcbiina (Kantor, 1991). Pscudoliva, by con-

trast, has a bicuspidate lateral plate and single head ten-

tacle. The latter character also occurs in Zcmira (see

Ponder & Darragh, 1975; Kantor, 1991). The absence of

a posterior notch on the outer lip is a character that

Naudoliva has in common with Benthobia and Fusul-

culus Naudoliva differs from these genera by having a

long labral tooth, deep anterior siphonal notch, and a

low but distinct siphonal fasciole. It differs from Luizia

by having a smooth instead of hrate inner side of the

outer hp, and bv lacking a posterior notch.

The type species, N. caitlinac, is from Transkei, South

Africa. Its shell is verv shghtlv constricted at the base,

Page 68

THE NAUTILUS, Vol. Ill, No. 2

and lacks axial sculpture. Naiidoliva vorstcri. recently

described by Lussi (1995) from Algoa, Bay, differs from

N. caitlinae by lacking the basal constriction and by hav-

ing about ten short avial ribs at the shoulder These ribs

do not extern! up to the sutiue or dowii to the gently

rounded peripherv, and become obsolete near the outer

lip. The absence of the periostracum characterizes both

species, and readilv distinguishes this unusual genus

from other South .Africa pseudolivid (see also Lussi,

1995).

Mansfield (1925) described Pseudolwa guppi/i on the

basis of a single, exquisitely preserved specimen from

the Spring\ale Formation (Late Miocene) of Trinidad.

Mv examination ot the holot\pe reveals that the shell is

moderately high-spired, the last whorl comprising 73%

of total shell height. The last whorl is not constricted at

the base; it is sculptured by very faint spiral threads, and

below the pseudoli\id groove by more prominent spiral

cords. The low shoulder is marked h\ sLxteen short, weak

axial ribs. The shallow pseudoli\id groove is situated just

below the middle of the last whorl, and ends in a rela-

tively long, sharp labral tooth. The inner side of the out-

er Up is smooth. The aperture is ovate, its height-to-

breadth ratio being about 2.3. .A posterior notch, parietal

rib, siphonal fascicle, and umbilicus are lacking. The an-

terior siphonal notch is deep.

I assign Mansfield's species proxasionally Naiidoliva. It

is ven. similar in form and sculpture to iV. vorsteii. from

which it differs in having a larger number of axial ribs

(sLxteen versus ten), a broader aperture (apertural

height-to-breadth ratio 2.3 instead of 2.7), and a smaller

shell (10.4 versus 29.4 mm). The only significant feature

of Pscudoliva guppi/i that is inconsistent with assignment

to \audoliva is the absence of a siphonal fasciole, which

is low and w^eak in Naiidoliva. In the absence of a fas-

ciole, P. giippiji resembles Fusulculiis, with which it ad-

ditionally has in common the numerous short axial ri-

blets and the absence of a posterior notch. P.seudolivo

guppiji differs from Fusidculiis, however, in not being

constricted at the base, in having a long labral tooth, a

much deeper siphonal notch, a relatively broader aper-

ture, and a shorter spire. It is possible that the absence

of a siphonal fasciole in the Miocene fossil as well as in

species of Fitsidcuhis. Bcnthohia. and Zcmira is a juve-

nile or neotenic feature. For now, it seem best to assign

Pseudoliva guppyi to Naiidoliva, despite the great geo-

graphic distance between Trinidad and the South Afri-

can localities where the other hvo species of Natidoliia

occur

Natidoliia guppyi is only one of two pseudohvids

known from the tropical western Atlantic Neogene, the

other being Macron con.stricttis from the early Miocene

of Venezuela. As noted by Mansfield (1925), there is no

h\ing or fossil tropical American species that is e\en re-

motely similar to this unusual late Miocene .species from

Trinidad.

Genus Liiizia Douville, 19.33

Type species: Biiccininn (Liiizia) co.stac Douville,

1933, by original designation.

Diagnosis: Shell small, ma.ximum height 22 mm;

ovate; protoconch paucispiral; sutures appressed; spire

moderately high, last whorl comprising 77 to 83% of

total shell height; last whorl shouldered above, not con-

stricted basally; spiral sculpture consisting of \erv fine

threads over whole shell surface; axial sculpture consist-

ing of 14 to 17 riblets on upper part of whorl, forming

nodes at shoulder; pseudofixid groove situated high on

last whorl, about one-third the tlistance from the ante-

rior end of the shell; outer hp planar, with small, distinct

labral tooth; inner side of outer lip with 11 to 12 Hrae;

posterior notch shallow; aperture narrowK elongate, its

height-to-breadth ratio 3.-3; columellar and parietal cal-

lus thin, of very small extent; parietal rib absent; siphonal

fasciole bounded posteriorly by low keel; anterior notch

deep; umbilicus absent.

Stratigraphic and geographic distribution: Early?

Miocene to Recent, Angola.

Discussion: Douxille (1934) proposed Liiizia as a sub-

genus of Buccinum for a small gastropod from the Mio-

cene (probably Burdigahan, Early Miocene) of Angola.

Although he noted the presence of a pseudoli\id-hke

groove on the lower part of the last whorl. Douxille

(1934) assigned Liiizia to Buccinum because of its weak

axial riblets and spir;il threads. In the Recent fauna, Liii-

zia is represented by Pseudoliva zebrina A. Adams, 1855,

from Angola (Figure 13-15). This species has a small

shell (maximum height 19.2 mm) with a relatively high

spire (last whorl comprising 77 to 83% of total shell

height), 14 to 17 weak axial riblets forming nodes at the

shoulder crossed by very weak threads. The inner side

of the outer lip bears 11 to 12 weak firae.

The eenus Luizia closely resembles Naiidoliva, Fu-

sulciilus, and slender species o{ Sulcobuccinum. It differs

from all these by having the inner side of the outer Up

Urate (sculptured bv small riblets) instead of smooth.

Luizia differs from Fii.siilciiliis and Naiidoliva h\ having

a weak posterior notch, and from Fu.sulculii.s h\ the pres-

ence of a siphonal fasciole and hv lacking a basal con-

striction on the last whorl.

Genus Fii.siilcttlu.'i Bouchet & Vermeij, 1998

Type species: Fu.sulciilu.s crcnatu.s Bouchet & Ver-

meij, 1998, by original designation.

Diagnosis: Shell relatively small, maximum height

18.4 mm, fusiform; protoconch paucispinil; sutures ap-

pressed; spire relatively high, last whorl comprising 61

to 67% of total shell height; last whorl v\ith concave sub-

sutural ramp, a rounded shoulder, and basal constriction;

spiral sculpture consisting of up to 15 weak cords be-

hveeu shoulder and ba.se, sometimes obsolete; axial

sculpture consisting of 14 to 16 riblets extending from

the suture to a point just below the shoulder; pseudolivid

groove at lip convex veutrallv, with veiv weak lalual tooth

and five weak basal crenulations; inner side of outer lip

smooth; posterior notch obsolete; aperture narrowly

G. J. Vermeij, 1998

Page 69

ovate, its height-to-breadth ratio 2.6 to 3.2; coluinellar

and parietal calhi.s thin, of \erv limited extent; parietal

rib ab.sent; .siphonal fasciole and umbilicus absent; an-

terior notch shallow.

Stratigraphic and geographic distribution: Recent,

bath\al southwestern Pacific from Coral Sea and New

Caledonia to north of New Zealand.

Discussion: The genus Fiisulciilus resembles Beutlio-

hia and Zi'inira in ha\ing a \en' shallow anterior siphonal

notch, a limited e.xtent of callus, a high position of the

pseudoli\id groove on the last whorl, and in lacking a

parietiil rib at the adapical end of the inner lip. Fusnl-

ciiliis differs from these genera bv being constricted at

the base, and from the hpe species of Zt'iuira bv ha\ing

well developed axial sculpture. Naiuloliia and Fiisulcii-

lus have in common the absence of a posterior notch on

the outer lip, but Fiisiilailus is distinctlv constricted at

the base, whereas Nmidoliva has httle or no basal con-

striction. Moreover, Nauiloliia has a low, rounded si-

phonal fasciole and a deep siphonid notch, features lack-

ing in Fiisidcithts.

Bouchet and Vermeij (1997) describe i\vo species, F.

crcnotus and F. olbiis, from bath\al waters in the south-

western Pacific. These species are the onlv pseudoUvids

Ii\ing in the tropical Indo-VVest Pacific.

Genus Zcmira H. & A. Adams, 1853

Type species: Ebiinia aiistralis G. B. Sowerbv, 1833,

by monotvpy.

Synonym:

Ebumopsis Tate, 1889 (t\pe species: Ebumopsis aulacoessa

Tate, 1SS9, = Zcmira australis. fide Ponder & Darragli,

1975)

Diagnosis: Shell relativelv small, maximum height 21

mm, ovate; protoconch paucispiral; spire moderatelv

high, last whorl comprising 66 to 68% of total shell

height; sutures deeplv channeled; last whorl rounded

above, not constricted basallv; spiral sculpture consisting

of as manv as 25 weak flattened cords between suture

and base of last whorl, the six to se\'en cords below pseu-

dohxid groove widest; axial sculpture absent; pseudohvid

groove situated high on last whorl, just below its mid-

point; outer lip with distinct labral tooth and basal cren-

ulations; inner side of outer lip smooth; posterior notch

present; aperture ovate, its height-to-breadth ratio 1.9 to

2.2; columellar and parietal callus thin, of very hmited

extent; parietal rib absent; siphonal fasciole absent; an-

terior siphonal notch obsolete; umbilicus usually absent.

Stratigraphic and geographic distribution: Late

Eocene to Recent, Australia,

Discussion: Ponder and Darragh (1975) have fully re-

viewed the living and fossil species of TiCmira and pro-

vided anatomical details tor the two living species, Z.

australis (northern New South Wales to Tasmania) and

Z. hoclalla (southern Queensland). H. & A. Adams

(1853) proposed Zcmira as a member of the muricid

subfamiK- Puipurinae because of characters of the rad-

ida. Fischer (1884) regarded Zcmira as a section of Ma-

cron in the Buccinidae. Cossmann (1901) assigned it to

the buccinid subfamily Latnmculinae (= Babvloniinae).

Poniler and Darragh s (1975) anatomic;d work revealed

similarities to Pscudoliva, and suggested that the ratlular

similarities with PurjDurinae (= Ocenebrinae and Rapan-

inae) result from inheritance of plesiomorphic character

states (see also Kantor, 1991).

Zcmira resembles Fusiilculus in its relatively high

spire, position of the pseudohvid gnxjve, lack of a si-

phonal fasciole, and obsolete anterior notch. It differs

from Fusulcultis by lacking the basal constriction, by

having a deeply channeled suture, and bv lacking axial

sculpture in most species. The labral tooth is distinct in

Zcmira but obsolete in Fiisulculus. Zcmira differs from

Bcnthflbia bv having a higher spire, a much less inflated

last whorl, well-developed spiral sculpture, a posterior

notch, and a labral tooth. Axial sculpture, which is pres-

ent as short riblets on the upper part of the whorl in

Bcnthobia, is lacking in Zemira .

The tvpe species of Zcmira , Z. australis. is interpret-

ed by Ponder and Darragh (1975) as a long-ranging,

variable species. It includes Ehiirnopsis aulacoessa Tate,

1889 and Zcmira praccursoha Tate, 1888, wliich are

heavily corded and smooth forms respectively from the

Balcombian (middle Miocene) of Victoria. The spire of

Z. australis is generally higher than in other species of

the genus. Axial sculpture is absent in Z. australis and

in its presumed progenitor, Z. australis antecursoria,

from the Longfordian and Batesfordian (early Miocene)

of Victoria; but it is present in Z. tcsscllata, the Late

Oligocene ancestor of the Z. australis lineage from the

Jan Juc Formation of Victoria (Ponder & Darragh,

1975). An umbilicus is absent in Z. a. australis, but is

variably developed in the other species assigned by Pon-

der and Darragh (1975) to Zcmira . The earliest repre-

sentative of Zcmira recognized b)' Ponder and Darragh

(1975) is an undescribed species from the Glen Aire

Clay (Late Eocene) of Victoria. This species lacks axial

sculpture and has a wide umbilicus.

Genus Tcstallium Vermeij & DeVries, 1997.

Type species: Gastridium ccpa Sowerbv, 1846, by orig-

inal designation.

S>'nonyms:

Gastridium Sowerbv, 1846 (tvpe species: Gastridium cepa

Sowerbv, 1846), fuwi Modeer, 1793, nee Sowerby, 1842.

Diagnosis: Shell large, maximum height 57 mm,

broadly fusiform; protoconch unknown; last whorl

rounded or shouldered above, constricted basallv; spiral

sculpture variably expressed, consisting of cords that are

strongest below pseudolivid groove; axial sculpture usu-

ally absent, or when present is confined to spire whorls

(T! valuta); pseudohvid groove at low position on last

whorl; outer Up planar, with distinct labral tooth and

Page 70

THE NAUTILUS, Vol. 111. No. 2

about five basal crenulations; posterior notch present;

aperture ovate, its height-to-breadth ratio 1.9 to 2.4; col-

umellar and parietal callus of liniitetl extent but may be

thick; parietal rib present; siphonal hisciole strong,

bounded above by prominent keel; anterior notch deep;

umbilicus present or absent.

Stratigraphic and geographic distribution: Early

Miocene to Late Pliocene, temperate western South

America.

Discussion: Vermeij and DeVries (1997) have given a

full account of the svnonvnnv and composition of the

genus TcstaUutm from the Miocene and Phocene of tem-

perate western South America. The name Gastridium

Sowerby, 1846, which has been used previously for these

species (Sowerby, 1846; Phihppi, 1887; Herm, 1969), is

preoccupied by Gastridium Modeer, 1793 (Conidae) and

Gastridium Sowerby, 1842 (a synonym of Pscudoliva).

Von Ihering (1907) referred species of Sowerby 's (1846)

Gastridium to AustrocomincUa von Ihering, 1907, which

he proposed as a subgenus oi ComincUa Gray, 1850. All-

mon (1990) and Beu & Maxwell (1990) have pointed out

that ComincUa {AustrocomincUa) fucgcnsis von Ihering,

1907, the type species of AustrocomincUa from the Rio

Turbio and Loreto Formations (Earlv? Miocene of Ti-

erra del Fuego), is closely related to or belongs in the

same genus as does ZclandicUa subnodosa (Hutton,

1877), the type species of ZclandicUa Finlay, 1926, from

the Altonian (Early Miocene) of New Zealand. Von Iher-

ing (1907) mistakenl)' beUeved that the South American

species of Gastridium Sowerby, 1846, lack a pseudohvid

groove. The inclusion of these species in Austrocomi-

ncUa, which indeed does lack a pseudolivid groove, was

therefore unwarranted. Species of Gastridium have also

occasionally been confused with members oi the ocene-

brine muricid genus Chorus Gray, 1847 (tvpe species:

Chorus giganteus (Lesson, 1831)), from Chile. Both gen-

era have a labral tooth corresponding to a basal groove,

but Chorus lacks the posterior notch and parietal rib

characteristic of TestaUium, the replacement name for

Gastridium Sowerby, 1846. In Chorus as in other mur-

icids, the spiral sculpture of the last whorl increases in

prominence from the base to the shoulder rather than

from the shoulder to the base as in TestaUium and other

pseudoUvids. The cords of Chorus are widely separated

and rounded, not crowded as in TestaUium.

TestaUium differs from Psendoliva bv haxing the last

whorl covered with fine spiral sculpture instead of being

smooth, by having the base of the last whorl distinctly

constricted (that is, with a concave lateral profile) instead

of being evenlv tapered, by having a prominent, keel-

like siphonal fasciole instead of a low, rounded one, bv

the very fimited develcjpment of callus, and by having

the lower part of the outer hp distinctly crenulated. It

differs from Suleobuceirium by the lower position of the

pseiidoli\nd groove on the last whorl, bv the limited de-

velopment of callus, and by tlie presence of a parietal

rib at the adapical end of the inner Up. TestaUium is verv'

similar to Macron. Whereas the inner side of the outer

lip of TestaUium is smooth, that of Macrtm is sculptured

with riblets (lirae).

There is considerable intraspecific variation in the de-

gree of development of the shoulder In some forms of

T ccpa and in T. vohita, the shoulder may be developed

as a raised ridge in the mature shell; whereas in other

specimens of T. ccpa, the upper part of the whorl is

rounded (.see Venneij & DeVries. 1997).

Included in TestaUium are T! ccpa from the EarK' Mio-

cene of southern Peru to southern Chile, T. vohita from

the Heath Formation (PEarlv Miocene) of northern

Peru, and T c.scalonia from the Late Phocene of Chile.

TestaUium vohita, which Olsson (19.32) originally de-

scribed as a species oi Acanthiza (Chorus), differs from

the other two species by having axial ribs on the spire

whorls. Olsson's Acanthiza was presumably a misspelfing

of Acantluna, another ocenebrine muricid genus with a

labral tooth (see Vermeij & DeVries, 1997).

Genus Macron H. & A. Adams, 1853

Type species: PscudoUva (Macron) hcUettii A. Adams,

1854 (= Bueeinum aethiops Reeve, 1847).

Synonym:

Macronisais Thiele, 1929 (T\pe species: Macron Uvidus A. Ad-

ams. 1S55).

Diagnosis: Shell small to large, maximum height 20

to 100 mm, ovate; protoconch paucispiral; spire relative-

ly high, last whorl comprising 68 to 74% of total shell

height; sutures appressed to deeply impressed; last

whorl rounded or shouldered above, httle or not con-

stricted basallv; spiral sculpture consisting of weak to

very prominent cords decreasing in strength from base

to shoulder; axial sculpture absent; pseudolixid groove

situated low in last whorl; outer lip planar, usualK' with

distinct labral tooth (obsolete in M. Uvidus), strongly

crenulated below; inner side of outer lip hrate; posterior

notch weak; aperture ovate in most species, its height-

to-breadth ratio 1.9 to 4.1 (narrowly elongate in M. mer-

riami): columella v\ath weak fold at entrance of siphonal

canal; columellar and parietal callus of very fimited ex-

tent; siphonal fasciole prominent, bounded above bv

keel; anterior notch deep; umbilicus present or ab.sent.

Stratigraphic and geographic di.stribution: Early

Miocene to Recent, warm-temperate northeastern Pa-

cific; PEarh- Pliocene, Japan.

Discussion: Macron has remained a taxonomic enig-

ma ever since H. & A. Adams (1853:132) introduced it

as a subgenus of Pscudoliva in the muricid subfamily

Puipiuinae. Adams (1855) and Sowerby (1859) main-

tained this assignment. Fischer (1884) referred Macron

to the Buccinidae, along with other Pseudoli\adae. Coss-

mann (1901) and MeKill (1903) refined this assignment

bv referring the genus to the Latnmculinae or Ebumi-

nae (= Babyloniinae), and noted th;it. whereas Macron

has an unguiculate operculum with a terminal nucleus,

Pscudoliva has a puipiuoid operculum with a laterally

G. J. Vermeij, 1998

Page 71

placed nucleus. Grant and Gale (1931) re-examined the

opercula of the two genera and concluded that hoth are

characterized by a terminal opercular nucleus. The\

therefore reaffirmed the close relationship hehveen Ma-

cron and Pscudoliva. Thiele (1929) described the radula

of M. trochlea (Grav) (= M. aethiops) and of A/, liiidus,

and assigned the genus to the Buccinidae. In this as-

signment, he was followed bv Wenz (1938—14), Glibert

(1963), Adchcott (1970), Keen (1971), and Abbott

(1974).

Yuri Kantor (personal communication) suggests that

the radula of Macron is verv' similar to that of the Nas-

sariidae. As illustrated bv Thiele under the name M.

trochlea, the rachichan tooth is multicuspidate with small

cusps, and the lateral teeth are tricuspidate. The pseu-

dolixid radula, hv contrast, either has a multicuspidate

rachidian and unicuspidate marginal teeth or a tricus-

pidate rachidian with very large cusps and bicuspidate

margined teeth (Kantor, 1991). The shell characters of

Macron, however, differ widely from those of Nassari-

idae. The genus Na.s.sarius Dunieril, 1906, and related

taxa (see CemohorsW, 1984) are characterized by de-

terminate growth, and by a polished outer lip with an

anterior sinus. Macron, by contrast, has a simple, shaqi-

edged, outer hp indicating indeterminate growth and

lacking am* trace of an anterior sinus.

The pseudolivid affinities of Macron are strongly sup-

ported bv shell characters. These include a thick peri-

ostracum, pseudohvid groove and corresponding labral

tooth (obsolete in A/, lividus), shaqi-eilged simple outer

hp, posterior notch, and spiral sculpture most stronglv

expressed near the base. The only similar genus that has

tracbtionally been referred to the Buccinidae is Trium-

phis, which differs from Macron chieflN' h\ the absence

of the pseudolivid groove and labral tooth. With the

transfer of Triitinphis to the Pseudohvidae (see below),

there is no other buccinid genus that closelv resembles

Macron.

Macron is most similar to Tcstallinni. from which it

differs principally in haxing the inner side of the outer

lip sculptured b\' nine to seventeen riblets (lirae). In Tes-

t allium, the inner side of the outer lip is smooth.

The middle Miocene species Macron mcrriami from

the Temblor Formation of southern California may rep-

resent an evolutionary link with Testallium. This species

has \er\' weak spiral sculpture, a rounded (not shoul-

dered) last whorl, appressed suture, weakly constricteil

base, and an unusually narrow aperture (height-to-

breadth ratio 4.1). Because the aperture of the holotype

is filled with matrix, the presence of Urae on the inner

side of the outer hp cannot be ascertained. The .species

could therefore be assigned to either Testallium or Ma-

cron. 1 retain it in Macron on the basis of its geographic

provenance (California instead of western South Amer-

ica).

Another species. Macron hatimanni. ma\' Unk Macron

with the genus Triumphia. This species, described bv

Hertlein and Jordan (1927) from the San Ignatio For-

mation (middle to late Miocene; see Smith, 1984) of

Baja Cahfomia Sur, Mexico, resembles Triumphis in

having a raised, keel-like shoulder, but differs from that

genus by having six to ten low, flattened cords on the

last whorl and by the presence of a blunt but distinct

labral tooth at the end of a pseudoh\id groove. Addicott

(1970) pro\isionallv assigned M. haiimanni to Niceina,

a taxon that has occasionalh' been considered a subgenus

of Triumphis but that is here assigned to the buccinid

subfamily Photinae (see below). Niccma lacks the labral

tooth and pseudoli\id grotne of Macron. I therefore

agree with Hertlein and [ordan (1927) that their species

should be referred to the genus Macron.

The type species of Macron, M. aethiops, is the larg-

est, most heavily sculptured hving member of the Pseu-

doIi\idae, attaining a height of 100 mm. The expression

of spiral sculpture abo\e the pseudolixid groo\e is highly

variable e\'en within populations. In t\pical M. aethiops,

seven thick, flattened or rounded cords with very deep

interspaces he between the pseudoh\id groove and the

shoukler. In the hcllettii form, only the lowest one or

two of these cords are expressed. All forms have a deeply

impressed, canahculate suture, and all have fifteen or

more hrae on the inner side of the outer lip. In the hving

fauna, M. aethiops occurs on the Pacific coast of Baja

Cidifomia from just south of Ensenada to just south of

Bahia Magdalena. There appears to be a geographically

isolated population in the Gulf of Cahfomia. The species

is not represented on the southern part of the Baja Cal-

ifornia peninsula. M\' examination of a specimen re-

ported by Mulliner (1996) from Isla Danzante in the

Gulf of Cahfomia indicates that this specimen falls well

v\ithin the hmits of variation of the species as a whole.

With onK' one cord above the pseudoli\id groove, the

specimen from Isla Danzante belongs to the kellettii

form of M aethiops. It is unusual among pseudolivids in

that the left (leading) edge of the inner lip has a chstinct

margin. In the fossil record, M. aethiops occurs in the

lower Olcese sand (middle Miocene) and the San Pedro

beds (Earlv Pleistocene) of southern Cahfomia (Grant

& Gale, 1931; Adchcott, 1970).

Thiele (1929) proposed the subgenus Macroniscus for

Pscudoliva (Macron) livida A. Adams, 1855, a small spe-

cies from the Pleistocene and Recent of southern Cah-

fomia. It chffers from other species of Macron by its

small size (maximum height 18.6 mm), appressed rather

than channeled suture, obsolete pseudoli\id groove and

labral tooth, and \'erv weak basal spiral cords. Although

M lividus does not closely resemble the large, strongly

sculptured Al aethiops, several species are moqjhologi-

cally intermechate. These include M. orcutti, M. mcleani

n. sp. (see below), and an earlv Miocene species. Macron

constrictus, described by Gibson-Smith, Gibson-Smith,

and Vermeij (1997) from the Cantaure Formation of

Venezuela. This less sculptured group does not merit

distinction as a subgenus or genus Macroniscus, which

1 therefore consider a subjective s\iion\iii of Macron.

Besides the h\ing and fossil species of Macron from

the warm-temperate northeastern Pacific and the early

Miocene species from Venezuela in the western Atlantic,

Page 72

THE NAUTILUS, Vol. HI, No.

Figures 16-17.

height 23.1 mm.

Macron mclcnni new species; Bahia Gonzaga, Gulf Coast, Baja Calitoniia Norte, Mexico; LACM .37-119. Shell

there is one species from the earlv Pliocene of Japan

that has been tentatively referred to Macron. Chinzei

(1959) named "Macron" nipponensis on the basis of a

single, thick-shelled specimen from the Kiibo Formation

of Honshu. It is characterized by thirteen spiral cords

on the last whorl and five cords on spire whorls, the

cords increasing in strength toward the constricted base.

Chinzei (1959) mentions neither a pseudolivid groove

nor a labral tooth. The outer hp, whose inner side is

ornamented with ten denticles, is exceptionally thick and

is posteriorly notched. An umbiUcus is absent. "Macron"

nipponensis differs from Macron hv the absence of a

labral tooth and pseudoli\iil groo\e and bv the presence

of denticles rather than brief lirae on the inner side of

the outer lip. If the species belongs to the Pseudolixidae.

it may have paralleled Triiunphis in having lost the pseu-

dolixdcl groove and labral tooth.

For the time being, I retiiin M. nipponensis in the

genus Macron.

Macron mclcani. new species

(Figures 16-17)

Diagnosis: Macron with smooth, rounded whorl, ap-

pressed suture, nine hrae on inner side of outer lip, nar-

row aperture, and fibrous periostracum.

Description: Shell ovate-fusiform, consisting of about

fi\e teleoconch whorls; spire moderately high, last whorl

comprising 68% of total shell height; suture appressed;

last whorl gently and evenly rounded on upper part, veiy

weakly constricted at base; spiral sculpture consisting oi

three low cords on ba.se, above which is a distinct (pseu-

dolivid) groove; iLxial sculpture absent; outer lip planar

with three strong crenulations on anterior sector below

a blunt labral tooth; inner side of outer lip with nine

brief lirae; posterior end of aperture with parietal rib;

inner lip appressed, with a distinct fold at entrance of

siphonal canal; aperture elongate, its height-to-breadth

ratio 3.1; siphonal fasciole low, rounded; anterior si-

phonal notch deep; umbificus absent; periostracum fi-

brous; shell brown, interior of aperture white.

Holotype: Height 23.1 mm, chameter 11.9 mm, ap-

erture height 15.6 mm, aperture width 5.1 mm. L.\CM

2812.

Type locality: Willard Point, Balria Gonzaga, Baja

California No'rte, Mexico, 29°48' N, 114°23' W, two to

three fathoms in sand.

Etymology: I take great pleasure in naming this spe-

cies for James H. McLean, great authority^ on the living

mollusks of the northeastern Pacific, who brought this

species to mv attention.

Remarks: Macron mclcnni is most similar to M livi-

dus. Both species have a distinctl)' fibrous periostracum,

a broadly fusiform shape, obsolete spiral sculpture, and

an appresseil suture. Macron mclcani differs from M.

liiidu.s bv having a chstinct rather than an obsolete labral

tooth, bv having nine strong lirae instead oi nine to

twelve vei-v fiiint lirae on the inner side of the outer lip,

and bv having a nnich narrower aperture (apertural

h(iglit-to-wi(lth ratio 3.1 as compared to 2.3 to 2.5 in M.

liii(lus). Other relatively smoodi species of Macron (M.

orcutti and the early Miocene species being described

bv C.ibson-Smith et al., 1997) have a relativelv broader

G. J. Vermeij, 1998

Pasje 73

aperture and a larger number of lirae (thirteen to sev-

enteen) than M. mclcani.

Genus Triiiinplii.s Grav, 1S57

Tjpe species: Biicciniiiii distoiium Wood, 1828, by

monotypy.

Diagnosis: Shell large, maximum height 6.3 mm,

ovate; protoconch paucispiral, consisting of 2.25 to 2. .5

smooth whorls; spire relatively high, last whorl compris-

ing 66 to 7.5% of total shell height; spire whorls flat-

tened, sutures appressed; last whnrl rounded above in

young stages, but with a strong, ridge-hke shoulder dur-

ing final growth stages; basallv not constricted; spiral

sculpture consisting of two or three cords on spire

whorls, one or tvvo low, rounded, below shoulder on last

whorl, and five stronger basal cords on last whorl; axial

sculpture confined to spire whorls, consisting of ten

rounded ribs per whorl; pseudolivid groove and labral

spine absent: outer lip more or less planar, its base

sharplv crenulated at terminations of groo\es between

basal cords; posterior notch weak, often obsolete in ma-

ture shells; inner side of outer lip with ten to eleven long

lirae; aperture ovate to elongate-ovate, its height-to-

breadth ratio 2.1 to 2.8; posterior end of aperture con-

stricted by massive, rounded parietal ridge; columella

with three or four superficial ridges on upper part, and

with weak fold at entrance of siphonal canal; columellar

and parietal callus often thick, but of ven' limited extent;

siphonal fasciole low, rounded; anterior siphonal notch

deep; umbilicus absent.

Stratigraphic and geographic distribution: Early

Miocene, southern California; Recent, tropical eastern

Pacific.

Discussion: All pre\ious authors who have concerned

themseKes with the classification of Tritiinphis ha\e as-

signed it to the Buccinidae. When introducing Tiium-

phis, Gray (1857:15) referred his new genus to the Pis-

aniana (= Pisaniinae), a group which, along with other

buccinid subfamilies, he assigned to a greatlv expanded

Muricidae. Fischer (1884) treated it as a subgenus of

Cominclla Gray, 18.50. Cossmann (1901) affirmed this

assignment by referring the genus to the subfamilv

Cominellinae (= Photinae). Thiele (1929) and Wenz

(1938—44), however, considered Tiiiimphis a subgenus

or section of Caiithanis Roding, 1798, a genus usualK'

assigned to the subfamily Pisaniinae (see CemohorskT,

1971, 1975). Keen (1971) recognized it as a full genus,

but she did not specif\' where in the familv it belongs.

Assignment of Tiiiiinphis to the Photinae is precluded

on three grounds. A heavy periostracum of the kind seen

in Tiiumphls is not known in any member of the Pho-

tinae, in which the periostracum is either absent or ver\

thin. The outer lip of Triiimphis has two characteristics

inconsistent with assignment of the genus to Photinae.

In Triiimphi.s, the edge of the outer lip is sharp, and the

smooth surface of the inner side of the outer lip does

not cross the edge to the outer side. In most Photinae,

by contrast, the edge of the outer lip is polished, because

the inner layer minutely extends across the edge to the

outer side. The anterior part of the outer lip above the

siphonal notch lacks a sinus in Triumphh. Instead, this

part of the lip is planar and bears verv strong crenula-

tions. In most Photinae, the outer hp is mediallv convex,

and there is a variably developed concave sinus between

the convex sector above and the anterior siphonal notch

below. This sinus is so prominent in some genera that it

is occasionalK' referred to as a stromboid notch to em-

phasize the superficial resemblance with the anterior

part of the outer lip in the Strombidae. The sinus in the

Photinae is not strongly serrated as is the basal sector of

the outer fip in Triumphis. Triiimphis superficially re-

sembles some members of the Pisaniinae, including

Cantlmnts and related genera, in possessing a thick, per-

sistent periostracum, and by having an unreflected,

shaip-edged outer lip; but it differs in characters of shell

sculpture and in the profile of the outer lip. In Triiim-

phis. the strength of spiral sculpture increases from the

shoulder to the base. This condition is typical of the

Pseudohvidae as well as of many other neogastropod

groups including the buccinid subfamihes Photinae and

Siphonaliinae, and the families Nassariidae, Fasciolari-

idae, Costellariidae, and clades within the Toxoglossa,

among others. This condition contrasts with the situation

in the Pisaniinae as well as in most Muricidae, in which

spiral sculptural elements increase in prominence from

the base to the whorl shoulder. The outer lip of imma-

ture and most mature Triumphis is characterized by a

notch at the junction with the penultimate whorl. Such

a notch is absent in Pisaniinae. In the latter subfamily,

the outer lip has a con\ex profile in its medial sector,

and a broad, shallow sinus on its upper half. The outer

lip of Triumphis is strongly crenulated at its edge only

along its lower half, as in most other Pseudolividae,

whereas in the Pisaniinae it is crenulated along its entire

length.

I assign Triumphis to the Pseudohvidae because of the

following combination of characters: persistent perios-

tracum, planar shaip-edged outer lip without anterior

sinus, presence of posterior notch, spiral sculpture

strongest near base, and outer lip crenulated at ends of

basal spiral grooves. Triumphis clearly diverges from

most other pseudolixids b\' the absence of a pseudoli\id

groove and corresponding laliral tooth. The point along

the edge of the outer lip at which crenulations begin,

however, corresponds to the position of the pseudofivid

groove and labral tooth in such genera as Macron and

TcstaUium Thieles (1929) description of the rather gen-

erahzed radula of Triumphis is also consistent with

placement of Triumphis in the Pseudolividae. Anatomi-

cal data and molecular sequences will be needed to con-

firm the phvlogenetic affinities of Triumphis.

Addicott (1970) referred a single, poorly preserved

specimen from the Middle Miocene Olcese Sand of

southern CaUfomia to the genus Triumphis. If additional

material confirms this assignment, Triumpliis would have

Page 74

THE NAUTILUS, Vol. Ill, No. 2

a record dating back to the middle Miocene. The only

other species, T. distotia, is knowm onlv from the Recent

tropicid eastern Pacific, where it ranges from El SaKador

to northern Peni.

Kilbum and Tankard (1975) described Triumphis di-

lemma from early Pbocene strata in Cape Province,

South Africa (see also Kenslev and Pether 1986). Else-

where (Vermeij, 1997), I ha\e shown that this species is

not closely related to Triumphis, and I proposed a new

genus for it in the buccinid subfamily Photinae.

TAXA EXCLUDED FROM PSEUDOLIVIDAE

Several genera and species included by previous authors

in the Pseudolividae or Pseudolivinae or in taxa here

assigned to that family do not belong in this group. Some

of these are treated below. A few western North Amer-

ican Paleogene species assigned to Pscudoliva by previ-

ous authors were already excluded by Squires (1989) and

will not be discussed further here.

Material e.xamined is Usted in Appendix 2. I provide

diagnoses for Melapiiim, Nicema. and Scymourosphaera,

because important shell characters of these genera have

not been noted previousK. My conclusions about the ge-

nus Austrosphaera are surmised from Oleinik and Zins-

meister's (1996) paper The Cretaceous genera Fidgcrca.

Hydrotribiihis, and Pti/chosyca were so well described

by Sohl (1964a, b) that thagnoses for them are superflu-

ous. The important shell attributes of these genera are

mentioned below in the discussions of these taxa.

Genus Nicema Woodring, 1964

Type species: Nicema amain Woodring, 1964, by

(jriginal designation.

Diagnosis: Shell moderately large, maximum height

57 mm, broadly fusiform: protoconch paucispiral; spire

consisting of five to seven teleoccjnch whorls; spire mod-

erately high, last whorl comprising 68 to 82% of total

shell height; sutures appressed, increasingly deep in later

growth stages; last whorl rounded or weakly shouldered

above, the shoulder a raised ridge in mature specimens

of some species; last whorl basidly constricted; spiral

sculpture consisting of about seven weak cords on spire

whorls, and of up to seven cords on and just above bastd

constriction of last whorl; axial sculpture consisting of

eight to ten rounded folds on spire whorls, obsolete or

absent on last one or two whorls; pseudolivid groove and

labnd tooth absent; outer lip medially convex downward,

with adapical concave sinus and weak posterior notch;

lower sector of outer hp crenidatetl by ends of grooves

between basal cords; upper concave sector of outer lip

with smooth edge; inner side of outer lip briefly lirate

in adult; aperture elongate-ovate, its height-to-breadth

ratio 2.8 to 2.9; inner hp smooth, adherent or very slight-

ly erect; columellar callus very limited in extent; colu-

meUa smooth, concave, with basal fold at entrance of

siphonal canal; parietal rib present; siphonal fasciole

prominent, sculptured with spiral threads and bounded

adapicallv by keel; siphonal canal dorsally recurved, with

deep anterior ncjtch; umbihcus absent or represented by

very narrow slit.

Stratigraphic and geographic distribution: Late

Miocene to Recent, tropical eastern Pacific; Late Mio-

cene, Atlantic Panama.

Discussion: Woodring (1964) proposed the genus Ni-

cema lor several late Miocene buccinids from tropical

America. Besides the type species, N. amara from the

Gatun Formation of Panama, he included iV. fiiitiifera

(Grzybowski, 1899) from northern Peru and N. predis-

torta (Marks, 1951) from the Daule beds of Ecuador

Olsson (1964) considered Nicema as a subgenus of Nor-

thia Gray, 1847, and added his new species N. mictiia

and N. venada from the Miocene of Ecuador Keen

(1971) included Nicema as a subgenus ofTriiimphi.s, and

added stiU another species, N. subrostrata (Wood, 1828)

from the tropical eastern Pacific. Finally, Addicott (1970)

tentatively referred Macron harimauni Hertlein & Jor-

dan, 1927, to Nicema. I have retained the latter species

in Macron. All authors e.xcept Olsson (1964) considered

Nicema to be closely related to Triiimplti.s.

Despite its morphological similarih to Triumphis. Ni-

cema may belong to the buccinid subfamiK' Photinae

rather than to the Pseudolividae. Nicema resembles

Triumphis in having a low spire (last whorl comprising

79 to 82% of total shell height in N. .subrostrata. 68%

in IV. noctua, l'i% in T. distorta), axial sculpture confined

to the spire whorls, an adapical notch, strong rounded

parietal rib, adherent smooth inner hp, firate outer hp,

and a persistent periostracum. The most conspicuous

chfferences reside in the spind sculpture and the shape

of the outer hp. In Nicema, the constricted basal part of

the last whorl is adorned with fine threads, and stronger

cords occur above this sector In Triumphis. the strong-

est sculpture (of five cords) occurs on the Tuiconstricted

base. The outer hp of Triumphis is essentially planar in

profile, whereas that of Nicema is medially convex and

adapicallv characterized by a broad, concave, smooth-

edged sinus below a shallow notch where the outer lip

joins the penultimate whorl.

Characters of the outer lip and the exteniid sheU

sculpture support Olsson's (1964) suggestion that Nice-

ma is related to Norihia and other genera in the bnc-

ciniil subianiiK Photinae. The onlv characteristic incon-

sistent with this assignment is the absence of a sinus on

the lower h;df of the outer lip. Such a sinus characterizes

most photines. but not Nicema. Anatomical observations

on the li\ing .V. subrostrata will be needed to confirm

my assignment of Nicema to the Photinae.

In adthtion to the species assigned to Nicema by

Woodring (1964), Okson (1964), and Keen (1971), the

genus Nicema includes Clavella sohda Nelson, 1870,

from the Late Miocene of northern Peni (see also Ver-

meij & DeVries, 1997). This species, which Olsson

(19.32) assigned to the muricid subgenus Acanthiza

(Chonis), has eight to ten strong axi;d ribs on the spire

G. J. Vermeij, 199S

Page 75

whorls, a weaklv constricted liase, strong fasciolar ridge,

and narrow umbiliciil chink, has the shoulder r;iised into

a ridge in the adult stage. Olsson (1932) perceived a

pseudolivid groove, and compared his specimens of N.

solida to Gastrid'utm (= TcstaUium) ccpa and Acanthiza

{Clionis) valuta Olsson, 1932, a species here assigned to

TcstaUium. Mv examination of the holot\pe of ClavcUa

solida failed to reveal a pseudolivid groove, and Nelson

(1870) did not mention such a feature in his accurate

original description of the species. Instead, the last whorl

shows an abrupt ahapical narrowing near its base. This

abrupt narrowing is not like the deeply concave basal

constriction of Niccma suhrostrata. in part because the

siphonal canal t)( ClavcUa solida is not dors;dl\- recurved

as it is in other species of Niccma. but in other respects

Nelsons species resembles members ot Niccma.

Jung (1995) named the genus Judaphos for/, impar-

abilis Jung, 1995, a Late Miocene species from the Pa-

cific coast of Costa Rica. Judaphos cIoseK' resembles Ni-

ccma but differs from the latter genus bv being more

slender and ha\'ing a less prominent siphonal iasciole. It

is possible that Judaphos forms a link between Northia

and Niccma.

Genus Mclapium H. & A. Adams, 1853

Type species: Pinila lincata Lamarck, 1822, subse-

quent designation by Cossmann, 1901.

Diagnosis: Shell large, maximum height 100 mm, glo-

bose; protoconch large, paucispiral; spire \ery low, last

whorl comprising 97% or more of total shell height; last

whorl e\'enlv roimded above, stronglv constricted basal-

Iv: .spiral sculpture consisting of about six low cords on

bas;il constriction; axial sculpture consisting ot distinct

growth lines; pseudofivid groove and labral tooth absent;

outer hp simple, gently and evenly convex; inner side of

outer hp smooth; posterior notch \\eak but present; ap-

erture ovate, its height-to-breadth ratio 2.1; columella

with strong fold at entrance of siphonal canal; inner lip

adherent; adapical end of inner lip with strong parietal

tooth; siphonal Iasciole well developed, bounded adapi-

cally by prominent keel; siphonal canal dorsalK' stronglv

recurved; anterior siphonal notch deep; umbilicus ab-

sent; parietal and columellar callus well developed, ren-

dering apertural side of last whorl flattened, but callus

not extenchng onto spire.

Stratigraphie and geographic distribution: Early

Phocene to Recent, southern Africa.

Discussion: The genus Mclapium was placed in the

Strepturidae by Cossmann (1901) and in the Oh\idae

(Pseudolivinaej by Thiele (1929), Wenz (1938-44), and

Kilbuni (1989). A stud\' of shell and anatomical charac-

ters persuaded Kantor (1991) to place Mclapium in its

own family Melapiidae. According to Kantor (1991), Mc-

lapium differs from Pseudoli%idae by the absence of ac-

cessory salivary glands and of the operculum, and by

having the opening of the radular diverticulum to the

buccal cavity at the tip rather than at the base of the

proboscis. The accessory esophageal gland oi Mclapium

is h\pertrophied.

The shell of Mclapium also differs from that of the

Pseudolividae. The pseudolivid groove and labral tooth

are absent in Mclapium, and the base of the columella

is marked by a prominent fold at the entrance of the

siphonal canal. The siphonal canal, moreover, is long and

strongly recurved dorsally.

Given the characters of the shell o( Mclapium, I ten-

tatively agree with Cossmann (1901) in assigning the ge-

nus to the Strepturidae. Mclapium differs from the tvpe

genus Strepsidura Swainson, 1840, by the absence of

axial sculpture, and by the presence of a strong parietal

tooth. A considerable stratigraphie gap exists between

the last appearance of Strepsidura in the Late Eocene

and the first Phocene appearance oi Mclapium. Consid-

ering the poor pre-Phocene fossil record in South Africa,

such a stratigraphie hiatus does ik )t constitute strong ev-

idence against a phvlogenetic hnk between the Paleo-

gene Strepturidae and the Late Neogene Mclapium. If

Mclapium does prove to belong to the Strepturidae, the

name Melapiidae Kantor, 1991, would fall as a subjective

junior .svnonviii of Strepturidae Cossmann, 1901.

The genus Mclapium is represented by two hving spe-

cies, M. lincatum from the temperate coast of South Af-

rica, and M. datum (Schubert & Wagner, 1829) from

Natal and Mozambique. Kenslev and Pether ( 1986) ad-

ditionally recognize two fossil species, M. patcrsouac

Bullen Newton, 1913, from the Neogene of Bredasdorp;

and M. hawthonici Kensley & Pether, 1986, from the

30 m and 50m levels (Early to Middle Phocene) at Hon-

deklip Bay, Cape Province, South Africa.

Genus Austrosphacra Camacho ;//

Camacho, 1949

Fur

qut

&

Type species: Austrosphacra glabra Camacho in

Furque & Camacho, 1949.

Discussion: Oleinik and Zinsmeister (1996) have giv-

en a brief account of this Late Cretaceous and earhest

Paleocene genus from Patagonia and Tierra del Fuego,

Argentina. The shell is thick, low-spired, and semi-ovate,

with short, shallow anterior and posterior canals and a

poorly developed siphonal fasciole. Although spiral

sculpture, a pseudohvid gr(jove, and a labral tooth are

absent, Oleinik and Zinsmeister (1996) assigned A(/.s-

trosphacra with doubt to the buccinid subfamily Pseu-

dohvinae. I have not examined any material oi Austro-

■'iphacra, but on the basis of great similarity with Sci/-

mourosphacra (see below), I tentatively exclude Ai/.sfro-

sphacra from the Pseudolividae. Just where this genus

should be assigned reniiiins unclear.

Genus Sci/mourosphacra Oleinik & Zinsmeister, 1996

Type species: Sci/mouro.^phacra hulloidcs Oleinik &

Zinsmeister, 1996, by original designation.

Page 76

THE NAUTILUS, Vol. Ill, No. 2

Diagno.si.s: Shell ol niecliiiiii .size, maxiimini height 41

mm, broadly fu.siform; protoconth large, paucispir;il.

.smooth; teleoconch consisting of four to five whorls sep-

arated by impressed suture; spire low, last whorl com-

prising 79 to 94% of total .shell height; last whorl evenly

rounded on upper part, weakly constricted at base; spiral

sculpture consisting of very weak, low, fine cords, best

expressed on spire whorls and on upper part of last

whorl, but often absent on last whorl and not expressed

on base; axial sculpture absent: pseudolivid groo\'e and

labral tooth absent; outer lip of mature shell abaxiallv

expanded, weakly con\ex, slightly thickened and smooth

on inner side; posterior notch weak but present; aper-

ture broadly ovate, its height-to-breadth ratio 1.9 to 2.3,

adapicallv narrowed, but outer lip not adapically extend-

ed; columella rounded, its profile sigmoid, smooth, with-

out basal fold; parietal callus ventrally thick, extending

partly onto spire in some species; inner lip adherent;

parietal rib absent; siphonal fasciole absent or low and

rounded; anterior siphonal notch very shallow; umbificus

absent.

Stratigraphic and geographic distribution: Early

Paleocene, Seymour Island. Antarctic Peninsula.

Di.scu.ssion: The above diagnosis is based on my ex-

amination of paraty'pes of S. buUoidcs and all the other

three members of the genus, very Idndly sent to me by

Anton Oleinik. When they proposed the genus Scijmou-

wsphacra. Oleinik and Zinsmeister tentatively assigned

it to the buccinid subfamily Pseudolivinae, although they

noted that a pseudolivid groove and labral tooth are ab-

sent. Two characters are inconsistent with assignment of

Sci/nioiirosphacra to the Pseudoli\idae. Spiral sculpture

is best developed on the upper part of the whorl, and

therefore does not increase in prominence abapically as

it does in the Pseudolividae. The outer hp of Sct/mou-

rosphaera is abaxiallv expanded and is somewhat thick-

ened on its smooth inner side; it therefore indicates de-

terminate growth. Pseudohvids never show abaxial ex-

pansion of the adult outer lip and apparently continue

to grow in the spiral cbrection throughout life.

Just how Seijmourosphacro shoukl be classified re-

mains unclear. Despite superficial resemblance to Mc-

lapium, Sci/mourosphaera is unlikely to belong to the

Strepturidae. Members of that family have spiral sculp-

ture increasing in prominence abapically, have a strong

basal columellar fold, and display indeterminate growth.

Seyinourosphaera may belong to a large Paleogene com-

plex of buccinoid gastropods with low-spired shells that

have no Neogene or Recent relatives.

Oleinik and Zinsmeister (1996) named four species of

Scymuuroaphacra, all from a restricted interxal of the

Early Paleocene Sobral Formation and the uppermost

part of the underlying Lopez de Bertodano Formation

of Seymour Island, Antarctic Peninsula All the species

are extremely similar to each other, varving only in spire

height and in the degree of persistence of the verv' weak

spiral sculpture on the last whorl. I strongly suspect tliat

the three species confined to the Sobral Formation (S.

(Icprcs.sa. S. subiilohosa. and S. btilloides) are a single

species, which would then be known as S. huUoidcfi^ The

fourth and oldest species (S. cicvata), known from the

uppermost Lopez de Bertodano Formation as well as

from the Sobral Formation, differs from the others by

having a higher spire of five instead of four whorls. It

may be a distinct species.

Genus Fidgcrca Stephenson, 1941

Type species: Fidficrca venii.sfa Stephenson, 1941 ( = ?

Psctidoliva? aticnuata Wade, 1926; see Sohl, 1964b), by

original designation.

Discussion: Three genera from the Late Cretaceous

of the Atlantic and Gulf Coastal Plain of the United

States have often been included in the Pseudolividae,

probably because they possess a labral tooth (see Sohl,

1964a, b). One of these is the Santonian to Maastrichtian

genus Ftd^crca. The labral tooth is formed at the end

of a fasciolar band at the tapering, unconstricted base of

the gentK' rounded last whorl. The spire is high, the last

whorl comprising 50% or more of the total shell height.

Sculpture consists of fine spiral and axial elements. Sohl

(1964a, b) gives a thorough over\iew of the group.

Exclusion of Fidgcrca from the Pseudolividae is sup-

ported by several characters, including the high spire

and the formation of the labral tooth at the termination

of a fascif)lar band instead of at the end of a groove

above the fasciole. Where Fidgcrca should be assigned

remains unresolved. Ponder (1973) suggested an affinity

with the Colubrariidae (now often considered part of the

buccinid subfamily Pisaniinae; see Beu & Ma^cwell,

1987), perhaps because of the elongate fusiform shape,

high spire, and finely cancellate sculpture of Fuljyrca.

The labral tooth formed at the end of the fasciolar band

would, however, be unique among Pisaniinae. For the

present, I prefer to keep the familial assignment of Ful-

gerca open.

ACKNOWLEDGMENTS

I thank J. Cooper, H. Vermeij, and E. Zipser for tech-

nical assistance; J. H. McLean (LACM) and G. Rosen-

berg (ANSP) for bringing important specimens to my

attention; and W. D. Allmon (PRI), W. Blow (USNM),

P Bouchet (MNHN), A. D Hondt (IRSNB), D. Dockery

(MGS), T Goshner (CAS), L. S. Groves (LACM), M. G.

Ilarasewvch (USNM), A. W Janssen (RGM), R. N. Kil-

burn (NM), J. Lang (TMM), D. R. Lmdberg (UCMP),

Y. Noda, (IGPS), A. Oleinik (PU), G. Rosenberg

(ANSP), and J. D. Taylor (BMNH) for u.se of their col-

lections and loans of specimens. Yuri Kantor made ex-

tremely helpful comments on the manuscript. The re-

search was funded in part by a grant (NSF EAR-94-

05537) from the National Science Foundation to G. J.

Vermeij.

G. J. Vermeij, 199S

Page

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APPENDLX 1

Genera and species of PseucloLi\idae, together with a list

of material examined for this study. The genus in which

the species was described is given in parentheses ii it

differs from the genus to which it is assigned here.

Benthobia Dall, 1889

B tiyonii Dall 1889, type of genus: = Ntix alabaster Bar-

nard, 1960; Lacuna cossmanni Locard, 1897 (see

Bouchet & Waren, 1985). Distribution: Recent,

hath\al and abyssal western and eastern Atlantic,

southwestern Indian Ocean, southwestern Pacific.

Material examined: MNHN (Azores, Gulf of Mex-

ico); USNM 717352 (Tulean Madagascar), 824174

and 832910 (off Virginia).

FiihiHutuin Fischer, 1884

F. ancilla (Hanley, 1859) (Pseiidoliva), type of Sylvano-

cochlis Melvill, 1903. Distribution: Recent, Agulhas

Bank, South Africa.

F. sepimentiim (Rang, 1832) {Biicciniim). type ot Fulmen-

tiiiiL Distribution: Recent, West Africa (Togo to An-

gola). Material examined: MNHN (Cotonou,

Pointe Noire, Cap Esterias, Luanda); CAS 063185

(West Africa).

Fusopsis Ra\ii, 1939

F. canalifrra (Ra\ii, 1939) (PseudoUva), type of genus.

Distribution: Paleocene, Copenhagen, Denmark.

Fusulctilus Bouchet & Vermeij, 1997

F. albus Bouchet & Venneij, 1997. Distribution: Recent,

bathval. New Caledonia and north ol New Zealand.

Material examined: MNHN

F. crcnaius Bouchet & Vemieij, 1997, type of genus. Dis-

tribution: Recent, bathval. Coral Sea to northeast of

Fiji. Material examined: MNHN (New Caledonia).

Huhachia Etayo Sema, 1979

H. ramirolobae Etayo Sema, 1979, type of genus Distri-

bution: Paleocene, Guajira, Colombia. Material ex-

amined: UCMP 16014, 16105

Liiizia Douville, 19.33

L. costae (Douville, 1933) {Bucriniim). type of genus; =

L. costae var. hrevis (Dou\ille, 19.33) Distribution:

Early? Miocene, Angola.

L. zebrina (A. Adams, 1855) iFsciitlolita). Distribution:

Recent, Augola. Material examined: MNHN (Lu-

anda, Mo^amedes).

Macron H. & A. Adams, 18.53

M acthiops (Reeve, 1847) (Bucciniim). = Pseiidoliva kel-

lettii A. Adams, 1855 (type of genus). Purpura troch-

lea Gray, 1839, auct. non Buccinum trochlea Bnigui-

ere, 1789;

? Macron wrightii H. Adams, 1865, .'' M. stereoghjpta

Sowerby, 1882 (see MeKill, 1903). Distribution:

middle Miocene (Olcese Sand), Califorma; Pleisto-

cene to Recent, Pacific co;ist of Baja California and

Isla Danzante, Gulf of California (see Grant & Gale,

1931; Addicott, 1970; Mulliner 1996). Material ex-

amined: GJV' (Guerrero Negro); Mulliner collection

(Isla Danzante); USNM 55665 (Cedros Island),

60074 (San Quentin Island). 60075 (San Quentin

Bay), 102256 (Ballena Bavi, 105428 (Scammons La-

goon), 105429 (Abreojos), 105432 (San Ignacio),

105433 (Manuel Lagoon), 105434 (Point Abreojos),

253114 (Lagoon Heads), 265240 (Cedros Island),

265601 (San Benito Island), 265613 and 265771

(Point Abreojos), 332389 (Magdalena Bay), 852828

(San Ignacio Lagoon), 1499926 (Gulf of California).

M. constrictus Gibson-Smith, Gibson-Smith & X'emieij,

1997 Distribution: early Miocene (Burdigalian,

Cantaure Formation), X'enezuela. Material exam-

ined: NMB

M. hartmanni Hertlein & Jordan, 1927. Distribution:

middle to late Miocene (San Ignacio Formation),

Baja California Sur, Mexico. Material examined:

CAS 61714.07.

M. lividus (A. Adams, 1855) (Psmdoliva). t\pe of ^ta-

cronisciis Thiele, 1929. Distribution: Pleistocene to

Recent, southern California. Material examined:

GJ\' (La Jolla).

M. mcleani Vermeij, 1997. Distribution: Recent, Gulf of

California. Material examined: LACM 2812

M. merriami Arnold, 1907. Distribution: middle Mio-

cene (Temblor Fonnation), Califonna. Material ex-

amined: USNM 164982.

PM. nipponensis Chinzei, 1959. Distribution: Pliocene

(Kubo Formation), Honshu, Japan.

M oraitti Dall, 1918. Distribution: Pacific coast, Baja

California Sur, Mexico Material examined: LACM

79-26.27 (Punta Marquez), 71-.3.26 (Punta Abreo-

jos).

Naudol'wa K.ilbuni, 1989

N. caitlinae Kilbuni, 1989, tvpe of genus. Distribution:

Recent, Transkei, South Africa. Material Examined:

NM E5750A-107.

N. guppiji (Mansfield, 1925) {PseudoUva). Distribution:

Late Miocene (Spring\ale Fonnation), Trinidad. Ma-

terial examined: USNM 352664

N. vorstcri Lussi, 1995. Distribution: Algoa Ba\-, South

Africa. Material examined: ANSP 399956.'

Pscudoiha Swainson, 1840

P. crassa (Cinieliii, 179U [Buccinum), t\pe of PseudoUva;

= P strialula .\. Adams, 1854 (see Mekill. 1903).

Distribution: Recent, southeni Angola Material

examined: MNHN (Baia dos Tigres, Ilha de Luan-

da); CAS 063184 (West Africa).

P lutulenta Kensley & Pether, 1986 Distribution: early

Pliocene (50-nieter level), Hondeklip, Cape ProWnce,

South Africa. =? P. crassa (see text under PseudoU-

va).

G. J. Vermeij, 1998

Page 81

P. orbij^nijana Mayer, 1864. Distribution: Early? Mio-

cene, Madeira.

SiilroL

)ut(nuiin t

■OrbiCTiiv, 1850

S anroiisolmi (Blaiickenhoni, 1927) (Psetidoliva). Distri-

bution: Late Cretaceous (.Senonian), Palestine.

S. aflelckci (Adegoke, 1977) [Pscitdoliva). Distribution:

Paleocene (Ewekoro Formation), Nigena. Material

examined: USNM 17485.3.

S. ambifi^mim (Binkhorst, 1861) (Pi/niln). Distribution:

Late Cretaceous (Maastnchtian), Netherlands and '■'

Tunisia (see Per\inquiere, 1912).

S. bellecompttim (Muniz, 1993) (Psetidoliva). Distribu-

tion: Late Cretaceous (Campanian) (Gramame For-

mation), northeastern Brazil.

S. bocaserpentis (Maur\', 1912) (Psetidoliva). Distribu-

tion: Paleocene (Soldado Fomiation), Trinidad.

S. breve (Doncieax, 1908) (Psetidoliva). Distribution: Late

Paleocene (Thanetian: Spaniacian), southern France.

S. briarti (Vincent, 1928) (Psetidoliva). Distribution:

Early Paleocene (Danian: Calcaire de Mons), Bel-

gium (see Glibert, 1973). Material examined:

IRSNB 5467 (Mons) and unnumbered specimen

(Ludanoyka).

S. btissoni (Albanesi in Albanesi & Bus.son, 1974) (Pseti-

doliva). Distribution: Early Eocene (Series 6b),

southern Tunisia.

S. carinattim (Conrad, 1860) (Psetidoliva). Distribution:

Middle Eocene (Laredo, Stone City, Cook Mountain

Formations, Archusa Marl), Gulf Coastal Plain (see

Palmer & Brann. 1966; Dockerv, 1980). Material ex-

amined: ANSP 70.32

S. chavani (Tessier, 1952) (Psetidoliva). Distribution: Pa-

leocene (Marigot de Balhng), Senegal.

S. conifonne (Oppenheim, 1904) (P.setidoliva). Distribu-

tion: Eocene, Cameroon.

S. eoronaria (Olsson, 1930) (Psetidoliva). Distribution:

Middle to Late Eocene (Saman and Talara Forma-

tions of Olsson, 19.30), Peni Material e.xamined:

PRI 24221.

S. cretacetim (Kase, 1990) (Calorebatna). Distribution:

Late Cretaceous (Late Campanian to Early Maas-

tnchtian: Izumi Group), Japan.

S. ctiixicostattim (Briart & Comet, 1870) (Psetidoliva).

Distribution: Earlv Paleocene (Danian: Calcaire de