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Full text of "The Nautilus"

THE NAUTILUS 



Volume 111, Number 1 
Januanj 29, 1998 
ISSN 0028-1344 

A quarterly devoted 
to malacology. 



Marine Bicloc 
Woods Hole Or. '- 



FEB 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 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: LEAL@WATER.NET 

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. 
bieler@fmnh.org 



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. 
mikkel@amnh.org 



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\vise@hmns.org 



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- 



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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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Robertson, R. 1985. Four characters and the higher category 
svstematics of gastropods, American Malacological Bulle- 
tin, Special Edition No. 1:1-22. 

Sheehan, D. C. and B. B. Hrapchak, 1980. The theorv and 
practice of histotechnology, 2ntl ed. C. \'. Mosbv Com- 
pany, St. Louis, 481 pp. 

Sinione L. R, L. 1995. Ris.si'clln ornala. a new species of Ris- 
soellidae (Mollusca: Gastropoda: Rissoelloidea) from the 
southeastern coa,st of Brazil. Proceedings of the Biological 
Society- of Washington, 108:560-,56. 

Thiele, J. 1912. Die Antarktischen Schnecken und Muscheln. 
Deutsche Siidpolar E.xpedition 13, Zool. 5 (1913) 
f;isc.2(1912):18.3-286, pls.11-19. Berhn. Georg Reimer. 

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): 











F coin- 








F 


lilini 


inunis 


F lin 


1. 


prominent 1° spnul cord 




+ 


- 


- 


2 


2° and 3° spiral cords 




+ 


+ 


- 


3. 


dark longitiidinal flamnniles 




+ 


- 


-1- 


4. 


wiiitc spots on spiral cords 




+ 


+ 


+ 


5. 


pale subsntural area 




+ 


- 


- 


6. 


dark siphonal end 




+ 


- 


- 



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 



C. medi- 


C reti- 




americana 


culata 


C, pet 


+ 


- 


-1- 


+ 


- 


-1- 


+ 


- 


4- 



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 



1 < 


> r 

■J 


\ 






--N_ 


5-> 




W /^^Yc 


















> 






Y^^p^ 


< 


^ 




\\ ^ 


K^ 


8 


1 


PM 




/ 


'^ 



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. 

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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 
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@ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). 



THE NAUTILUS 



Volume 111, Number 2 
February 17, 1998 
ISSN 0028-1344 

A quarterly devoted 
to malacology. 



FEB 23 



m8 




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Dhision of Mollusks 
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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: bouchet@cimrsl.mnhn.fr 



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 
Mons), Belgium (see Ghbert, 1973). Material ex- 
amined: IRSNB .5469, .5470 

S. dechorilatiini (White, 1887) (Harpa). Distribution: Pa- 
leocene (Maria Farinha Formation), Brazil, ? Early 
Paleocene (Mangot de Balhng), Senegal (see Tessier, 
1952). ? = S. scalina (see Palmer & Brann, 1966). 

S. elegans (Hislop, 1860) (Psetidoliva). Distribution: Ear- 
ly Paleocene (Danian: Intertrapine beds), Nagpur, 
India Material examined: BMNH GS10297, 
10298, 10299. 

S. elisae (Briart & Comet, 1870) IPseiidoliva). = P lii- 
dovicae. P ^ro.ssecostata, and P elongata. all ot Briart 
& Comet, 1870 (see CLbert, 1973). Distribution: 
Early Paleocene (Danian: Calcaire de Mons), Bel- 
gium. Material examined: IRSNB 547I-.5474. 

S. eschi (Oppenhenu, 1904) [Psetidoliva). Distribution: 
Eocene, Cameroon; ? Middle Eocene (Ameki For- 
mation), Nigeria. 



S. fisrheri (Mayer-E\iiiar, 1887) (Psetidoliva). Distribu- 
tion: Late Eocene (Bartonian), Thun, Switzerland. 

S. fisstirattim (Deshayes, 1835) (Bticcintim), t\pe ot Stil- 
cobuccimim Distribution: Late Paleocene (Thane- 
tian), Paris Basin, France. Material examined: 
MNHN (Chenay); RGM .58771 (Chenay). 

S, fiinkeantun fiinkeantnn (Adegoke, 1977) (P.setidoliva). 
Distribution: Paleocene (Ewekoro Formation), Ni- 
geria. 

S.f. or^wtiim (Adegoke, 1977) (Psetidoliva). Distribution: 
Paleocene (Ewekoro Formation), Nigeria. 

S. gtiineense (Adegoke, 1977) (Psetidoliva). Distribution: 
Paleocene (Ewekoro Fomiation), Nigeria. Material 
examined: USNM 174854, 174855. 

S, hoivardi (Dickerson. 1914) (Molopophonis). type of Pc- 
gocoinpttis Zinsmeister, 198.3; = Psetidoliva reticulata 
Waring, 1915 (see Zinsmeister, 1983). Distribution: 
Late Paleocene (Santa Susana and Sepultura For- 
mations), southern California and Baja California. 
Material examined: LACM IP 22330. 22376, and 
22688 (Santa Susana Formation, Simi Hills), 

S. iminense (Moret, 1938) (Psetidoliva). Distribution: Pa- 
leocene, Sud-Atlas, Morocco (see SaKan, 1954). 

S. japonictim (Nagao, 1924) (Orthatilax). Distribution: 
Late Eocene to EarK' Oligocene (Takashiman, Oki- 
noshiman, Funazuan, and M;izean faunas), KntisIiu. 
Japan (see Honda, 1991, 1994). Material examined: 
IGPS 8068, 8106, .35661 (Futago-Jima), 8069, .35660. 
35684, .37769 (Hashima), 8105 (Shiratake), .35743 
(Komenoyama), 36176 (Okinoshima), .36186 (Funa- 
zu), .36714 (Kiuragi). 

S. kit.soni (Newton, 1922) (Psetidoliva). Distribution: 
Late Eocene (Ameki Formation), Nigeria (Eames, 
1957) Material examined: BMNH G42069, 42062, 
42070 to 42076. 

S. kocneni (Ravii, 1939) (Psetidoliva), non Pnrptira ptisilla 
Beyrich. 18.54 (see Ra\n, 1939). Distribution: Pa- 
leocene, Copenhagen, Denmark. 

S. latidtinense (Defrance, 1826) (Bticcintim). = Bticcintim 
setnicostattim Deshayes, 18.35. Distribution: Late 
Paleocene (Thanetian), Pans Basni, France (see 
Cossmann, 1889). Material e.xamined: UCMP loc. 
B-,5.393 (Smceny: MNHN (Pourcy). 

S. letittveini (Bohm, 1926) (Psetidoliva). Distribution: 
Eocene, Namibia. 

S. libtfciim (Quaas, 1902) (Psetidoliva). Distribution: 
Late Cretaceous (Maastrichtian), Eg)pt; EarK Paleo- 
cene (Danian), East Tripoli, Libya 

S. lineattim (Gabb, 1864) {P.setidoliva). = P dilleri Dick- 
erson, 1914 (t\pe of Calorebamn Srjuires. 1989), 
Psetidoliva inornata Dickerson, 1915, P. kirbiji Clark, 
1938, P. markletjensis Clark, 1938, P. tejonensis Dick- 
erson, 1915, P. voltifaefonnis Gabb, 1864 (see text 
under Sidcobttcciniini; for different treatment see 
Squires, 1989). Distribution: EarK' to Late Eocene 
("Meganos" to "Tejon" stages). Pacific North Ameri- 
ca. Material e.xamined: UCMP 3679 (P dilleri). 
11053 (P inoniaitim). 12472 [P markletjensis). 15409 
(P. voltitaefonnis), 15796 (P. lineata). 

S. linosttm (Conrad, 1860) (Psetidoliva). = Psetidoliva mo- 
ercns de Gregorio, 1890 (see Palmer & Brann, 1966). 
Distribution: Middle Eocene (Weches, Stone City, 
am! Cook Mountain Formations), Texas. 

S, maritinuiin (Squires, Zinsmeister & Paredes-Mejia, 



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THE NAUTILUS. Vol. Ill, No. 2 



1989) {Popi'iiocitin), t\pe ot Popiiun-uin Scjuire.s, 
Zinsiiieister & Parede.s-Mejia, 19.S9; = Popfnoetim 
nuiritiiniis hajcu'nsis Squire,s, Zinsiiiei.ster & Parede.s- 
Mejia, 19S9 (see te.vt under Sulrohuccinum). Distri- 
bution: Paleocene (San Francisquito and Sepultura 
Foniiations), Baja California. 

S. niichclini Coquand, 1S62, = Pscudoliva michclini vats. 
longispinata, ntultispinata. and paiicispiiiata. all of 
Salvan, 1954 Distribution: Middle Eocene (Sues- 
sonian). North Africa (.see SaKan, 1954). 

S. minutissimum (SaKan, 1954) {Pscudoliva). Distribu- 
tion: Early Paleocene (Danian: Calcaire de Boua- 
bont), Morocco. 

S. inoabiticum (Blanckenhoni, 1927) (PseitdoHva) . Distri- 
bution: Late Cretaceous (Senonian), Palestine. 

S. inidtinodulosum Vemieij, 1997, = Pseudolivn chnvani 
Glibert, 1973, non Tessier, 1952 (see text under Sul- 
cobuccinum). Distribution: Earlv Paleocene (Dani- 
an: Calcaire de Mons), Belgium. Material exam- 
ined: IRSNB 536S 

S. mntabile douvillei (Olsson, 1928) (PseudoUva). Distri- 
bution: Earlv Eocene (basal Restin Formation), 
PeRi. Material e.vamined: PRI 3681. 

S. m. mntabile (Woods, 1922) (PseudoUva) . Distribution: 
Early Eocene (lower Salina Group: Negritos For- 
mation), Peru (see Woods, 1922; Olsson, 1928; 
Squires ef al, 1989). 

S. m. woodsi (Olsson, 1928) (PseudoUva). Distribution: 
Early Eocene (upper Salina Group: Negritos For- 
mation), Peni (see Olsson, 1928; Squires rt«/., 1989). 
Material examined: PRI 3684. 

S, nanafaUnense (Aldrich, 1931) (PseudoUva). Distribu- 
tion: Earlv Eocene (Nanafalia Formation). Alabama. 

S. nanuin (Gardner, 1945,1 (PseudoUva). Distribution: Pa- 
leocene (lower Midway F"ormation), Nuevo Leon, 
Mexico. 

S, obtusum (Deshaves, 18.3.5) (Bueeiiiunt). Deshaves, 1865 
(see Cossmann, 1889). Distribution: Late Paleocene 
(Thanetian) and Middle Eocene (Lutetian). Paris Ba- 
sin, France. Material examined: MNHN Sapicourt, 
Boisgeloup). 

S. ostranipis ostranipi.s (Harris, 1896) (PseudoUva). Dis- 
tribution: Early Eocene (Bashi Marl Member, 
Hatclietigbee Fomiation), Alabama. 

S. .V. pauper (Harris, 1899) (Pseudoliva). Distribution: 
Early Eocene. Texas. Material examined: TMM 
BEG35590 and 35675 

S. perspeetivum (Conrad, 1860) (PseudoUva). Distribu- 
tion: Late Eocene (Moodys Branch Formation and 
Yazoo Clav), Mississippi. Material examined: GJ\'; 
MGS. 

S. praecursor (Petho, 1906) (PseudoUva). Distribution: 
Late Cretaceous (Senonian), Hnngaiy. 

S'. priinum (Defrance, 1827) iSlnilhioUiria). = Bueeiuuui 
tiara Deshaves, 18.35, PseudoUva pursaneusis Don- 
cieux, 1908, P binodosa Traub. 19.38 (see N'illatte, 
1970; Squires et ai, 1989). Distribution: Late Pa- 
leocene (Thanetian), Europe. Material e.xamined: 
IRSNB (Noaille). 

,S. pusiUuin (Beyrich, 1854) (Purpura), = Pur})ura nodu- 
lo.sa Beyrich, 1854 (see Wrigley, 1941, and text under 
Suh'i>bueeinuiuy Distribution: Late Eocene (Pria- 
bonian: Lattorfian), northwi'steni Europe. 

S. retiisuiu maneoren.se (Olsson. 1928) (Pseudoliva). Dis- 



tribution: Late Eocene (Chira, Mancora, and Heath 
Formations) and ? Earlv Oligocene, Peni. Material 
examined: PRI 3679, .3680 

S. r retnsum (Philippi, 1887) {Gastridium}, = PseudoUva 
parinasensis Woods, 1922 (Vermeij & DeVries, 
1997). Distribution: Earlv Eocene, Algarrobo, 
Chile; Earlv to Middle Eocene (Negritos and Talara 
beds of Olsson, 1928, 19.30), Peru; as PseudoUva sp. 
cf. P. parinasensis. Middle Eocene, Colombia (Clark 
& Durham, 1946). Material examined: SGO PI 
765; UCMP 5020. 

S. r samnnicum (Olsson, 1928) (Pseudoliva). Distribu- 
tion: Middle to Late Eocene (Saman and Talara For- 
mations of Olsson, 1928, 1930), Peru (= Verdun For- 
mation), Peni; Late Eocene (Gem Mainsjie), Cura- 
sao (Jung, 1974). Material e.xamined: PRI 3677, 
3678; UCMP .35021, .35022 (Colombia). 

S. robustum (Briart & Comet, 1870) (Pseudoliva). = Pseu- 
doUva canaliculata Briart & Comet, 1870 (see Gil- 
bert, 1973). Distribution: Earlv Paleocene (Danian: 
Calcaire de Mons), Belgium; ? Earlv Paleocene, Sen- 
egal (Tessier, 1952). Material examined: BMNH 
G8136, 8137, IRSNB 5466. 

S. rogersi (Adegoke, 1977) (Pseudoliva). Distribution: 
Paleocene (Ewekoro Fomiation), Nigeria. 

.S. rosenkrantzi (Traub, 1979) (Pseudoliva). Distribution: 
Late Paleocene (Oichinger Schichten). .\ustna 

S, rude (von Koenen, 1889) (Pseudoliva). Distribution: 
Late Eocene (Priabonian: Lattorfian), Germain' 

S, santander (Gardner, 1945) (Pseudoliva). Distribution: 
Late Paleocene to Early Eocene (Laretlo, Tuscaho- 
ma, Wilcox, Hatchetigbee, and Nanafalia Forma- 
tions), Gulf Coastal Plain of Mexico and the United 
States (see Dockerv, 1980). Material e.xamined: 
MGS (Bells Landing Member. Tuscahoma Fomia- 
tion). 

S. sealina (Heilprin, 1881) (Pseudoliva). Distribution: 
Early Eocene (Nanafalia and Hatchetigbee Foniia- 
tions), Alabama. ? = dechordata White, 1887 (see 
Palmer & Brann, 1966). 

S. .sehweinfurihi (Oppenheini. 1904) (Pseudoliva). Distri- 
bution: Eocene, Cameroon. 

S. sofegginense (Rossi Ronchetti, 1959) (P.'ieudoliva). Dis- 
tribution: Late Cretaceous (Maastrichtian), East 
Tripoli, Libya. 

S. stibcostatum paucicostatum (Rossi Ronchetti, 1959) 
(Pseudoliva) . Distribution: Late Cretaceous (Maas- 
trichtian), East Tripoli. Libya. 

S. .V, subeostatnni (Stoliczka, 1867) (Psendt>lival Distri- 
bution: Earlv Paleocene (Danian: .\rnaloor beds), 
southem India. 

S. tenuieo.statum (Briart & Comet, 1870) (Pseudoliva), = 
Pseudoliva dubia Briart & Comet, 1870 (see Glibert, 
1973). Distribution: EarK Paleocene (Danian: Cal- 
caire de Mons). Belgium. Material examined: 
IRSNB 5475 

S. thielei (Bohm, 1926) (Pseudoliva). Distribution: Eo- 
cene, Namibia. ? = S. kitsoni (see texi under Sul- 
eobueeinum). 

S. tid>ereidifenim ((Joiirail. 1860) (P.seudoliva). Distri- 
bution: Late Paleocene (Gregi2;'s Landing and Bell's 
Landing Members, Tuscahoma Formation; Naiije- 
mov Formation) of Cult and .Atlantic coastal Plains 



G. J. Vernieij, 1998 



Page 83 



(see Squires, 1989) Material examined: MGS; 
UCMP loc. A-1056 (Bells Landincr Meniher), 

S. unicnrintitiiin (Aldricli, ISS6) {Pseudoliva). Distribu- 
tion: Early Paleocene (Matthew's Landing Member, 
Porter.s Creek Formation), Alabama (Squire.s, 1989). 
Material examined: MGS. 

S. vettistum (Conrad, 18.33), t\pe ot Biircinorhis Conrad, 
1865; = Monoceros fusifonnis. M. pi/ndoides, and M. 
sulcatum, all of Lea, 1833 (see Palmer, 1937). Dis- 
tribution: Middle Eocene (Gosport Sand), Alabama. 
Material examined: GJV (Little Sta\e Creek), 
MGS 

S. vicntoritsc (Clark & Durham, 1946) {Pscu(li<liin). Dis- 
tribution: Eocene. Colombia Material examined: 
UCMP 3502.3 

S. zittelli (Petho, 1906) (Pseutiolivd). Distribution: Late 
Cretaceous (Senonian), Hungary 

S. sp. aff. S. rnaritimum (Pseuciolivn). Distribution: Pa- 
leocene (Sonja Member, Agatdal Formation), West 
Greenland (see Squires et ai. 1989; Kollmann & 
Peel, 1983). 

S. sp. (as Pseudoliia .scalina Heilpnn, 1896, non Heilprin, 
1881). Distribution: Earh Paleocene (Chuton 
Member, Porters Creek F"orniation), Alabama (see 
Squires et nl. 1989). 

S. sp. of Olsson, 1934 {Pseiidoliva}, Distribution: Late 
Cretaceous (Maastrichtian: Monte Grande Forma- 
tion), Peru. 

Sidcolivn \"enneij. 1997 

S. modcsta (Olsson, 1928) (Pseudoliia). Distribution: 
Late Eocene (Talara Formation of Olsson, 1930), 
Peni. Material examined: PRI .3668. 

S. monilis (Olsson, 1928) (Pseudoliva), t\pe of genus. Dis- 
tribution: EarK' Eocene (Parinas Sandstone and 
Restin Foniiation), Penr Material examined: PRI 
.3671. 

Testallium Wrmei] & DeX'ries, 1997 

T. cepa (Sowerby, 1846) (Gastridiutu), t\pe of genus; = 
Monoceros labiate Hupe, 1854, Fusus labialis Hupe, 
18.54, Monoceros opimum Hupe, 18.54, Fusus opimus 
Hupe, 1854, Chorus sp. aff. C. blainvillei of \\'atters 
& Fleming. 1972 (see V'emieij & DeVries, 1997), 
Distribution: Early Miocene (Navidad Fonnation 
and Punta Ahuenco beds), Chile; EarK to Early Mid- 
dle Miocene (Chilcatay and Pisco Formations), Peni. 
Material examined: BMNH G26399 

T. escalonia Vermel] & De\ nes, 1997. Distribution: Late 
Pliocene (La Cueya Formation), Chile. Material ex- 
amined: CAS 66800.01-.02; UCMP .39880. 

T voluta (Olsson, 19.32) (Acanthiza). Distribution: Earlv 
Miocene (Heath Formation). Peni (see N'enneij & 
De\'ries, 1997). Material examined: PRI 2.302. 

Triumphis Gray, 1857 

T. distoria (\\'ood, 1.S28) iBuccinuni). tvpe of genus; = 
Columhclln triumphalis Duclos, 1843 (Keen, 1971). 
Distribution: Recent, tropical eastern Pacific. Ma- 
terial examined: GJ\' (Costa Rica. Panama); USNM 
.368541 (Pa\-ta, Pern). 

T. .sp. of Addicott, 1970. Di.stribution: Middle Miocene 
(lower Olcese Sand), Calitoniia 



Zemira H. & A. Adams, 1853 

Z, australis iintecur.soria Ponder & Darragh, 1975. Dis- 
tribution: Early Miocene (LongforcUan and Bates- 
lordian). Mctoria. Australia. 

Z. a. australis (G. B. Sowerby I, 18.33) (Ebunia). type of 
Zemira ; Zemira praeairsoria Tate, 1888; Ebumopsis 
aulae<H-ssa Tiite, 1889 (Ponder & DaiTagh, 1975). 
Distribution: miildle Miocene (Balcombiaii) to Re- 
cent, temperate Australia. Material examined: CAS 
28597 (Victoria); USNM 130901 (Tanner Chamber- 
lain), 202777 and 203906 (Westeniport). 

Z. bodalla Garrard, 1966 Distribution: Recent, southern 
Q)iieensland. 

Z. tessellata Tate, 1893. Distribution: |aii jiic Fonnation 
(Late Oligocene), Victoria Material examined: 
IRSNB (Spring Hill). 

Z. .s^ ot Ponder & Danagli, 1975. Distribution: Glen 
Aire Clay (Late Eocene), Nigeria. Material exam- 
ined: RGM (Bende). 



APPENDIX 2 

Material examined of some species belonging to neo- 
gastropod families other than Pseudolividae. 

Family Olixidae, subfamiK Ancilluiae 

Ban/spira australis (Sowerby, 18(30): GJV' (Leigh, New 

Zealand) 
Ebumn glabrata (Linn;ieus, 17.58): GJV' (near Punto Fijo, 

\'enezuela) 

Family Strepturidae 

Melapium lineatum (Lamarck, 1822): USNM 687845 and 
609711 (Jeffreys Bay. South Afnca) 

Strepsidura turaida (Solander in Brander, 1766): UCMP 
Localities B-5,357 (Cressay Late Eocene), B-5404 
(FleuiY-la-Ri\iere, Middle Eocene), B-.5402 (Her- 
oii\al. Late Eocene), B-6484 (Le\ignen, Late Eo- 
cene), all Paris Basin, France. 

FamiK Buccinidae, siibtamih' Photinae 

Cominella acutinodosa (Reeye, 1846): USNM 691693 and 
691707 (Bandicoot Bay, Barrow Island, Western Aus- 
tralia) 

losepha glandiformis (Reeye, 1847): GJV (several locali- 
ties near Leigh and Dunedin, New Zealand) 

;, ta.smanica Tenison-Woods, 1878: USNM 6.38.596 (Duns- 
borough, Western Australia) 

Nicema amara Woodring, 1964: USNM 24.3665 (lower 
Gatun Formation. Panama) 

N. noctua (OLsson, 1964): USNM 6440 IS and 64.5271 (An- 
gostura Formation, Ecuador, Late Miocene) 

N. solida (Nelson, 1870): YPM 00507 (Cardalitos beds, 
Peru, Late Miocene; see Vermeij & DeVries, 1997) 

N. subrostrata (Wood, 1828): GJV' (San Bias, Mexico) 

N. venada (Olsson, 1964): USNM 644019 (Rio Cayapas. 
Ecuador; Late Miocene) 

Family Melongenidae, subfamily Echinofulgurinae 

Coniulina minax (Solander in Brander, 1766): MNHN 
(Late Eocene, Paris Basin); UCMP Locality B-5.356 
(Crenes, France; Middle Eocene). 



Page 84 



THE NAUTILUS, Vol. Ill, No. 2 



Pmtobustjcon cretaceum (Wade, 1917): USNM 32897 
(Coon Creek, Tennessee; Late Cretaceous: Maas- 
trichtian) 

Families uncertain 

Hijdrotrihulus eleoans Sohl, 19fi4: MGS (Coon Creek, 
Tennessee; Late Cretaceous: Maastrichtian) nodosus 
Wade, 1916: USNM .3291.5 and 1304.56 (Coon Creek, 
Tennessee; Late Cretaceous: Maastrichtian) 

Laccinum lu^ardi (Newton, 1922); ROM (Ameki For- 
mation, Late Eocene, Nigeria) 



Lacinia alveata (Conrad, 18.33): GJ\', UCMP Loc. A-959 
(Gosport Sand. Middle Eocene, Alabama) 

Pttjcho.stjca inomata (Gabb, 1876): USNM 1.3454 and 
1.3455 (Riplev Fomiation, Late Cretaceous (Maas- 
trichtian) ot Tippah Counts', .Mississippi) 

Seymoiirosphaera bulloides Oleinik & Zinsmeister, 1996: 
FU Localities 11.36, 14.30, 1.501 (S. bulloides). 9 (S. 
depressa): 1434 and 14.35 (S. subglobosa): 1.328 and 
1138 (S. elevata) (all names of Oleinik & Zinsmeis- 
ter, 1996), all Paleocene of Seymour Island, Antarc- 
tica. 



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THE NAUTILUS 



Volume 111, Number 3 
May 26, 1998 
ISSN 0028-1344 

A quarterly devoted 
to malacology. 



JUN 4 1998 




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THE €7 NAUTILUS 



Volume 111, Number 3 

Moij 26, 1998 

ISSN 0028-1344 



CONTENTS 






D. L. Geiger 


Recent genera and species of the family Haliotidae 

Rafinesque, 1815 (Gastropoda: Vetigastropoda) 


85 


Cuido Pastorino 
Yuri Kantor 


On the identit\- oi Biicciuum lahijrinthus Gmehn, 1791 
(Gastropoda: Buccinidae) from the Netherlands coast 


117 



JUN 4 1998 



THE NAUTILUS 111(3):S5-116, 199S 



Page 85 



Recent Genera and Species of the Family Haliotidae 
Rafinesque, 1815 (Gastropoda: Vetigastropoda) 



D. L. Geiger 

Department ot Biological Sciences 
Uni\ersit\ ot Soiitheni Calitoniia 
Los Angeles, CA 90089-()371USA 
dgeigerS'scf. use. edu 



ABSTRACT 

A new assessment of all the Recent taxa of the family Halioti- 
dae is presented. The 17 Recent and one fossil genus-le\'el taxa 
are evaluated based on their tvpe species. These t;ua are rec- 
ommended not to he used mitil a coniprehensi\e phvlogenetic 
anaKsis of the fanulv is available. The approximatelv 200 nom- 
inate, species-le\el ta.\a in the Haliotidae are reviewed. Fift\- 
fi\e species witli ten additional subspecies are considered \ alid. 
The following new synonymies are proposed: Haliotis brazicri 
Angas, 1869, + melciilus (Iredale, 1927); H. hari^ravcsi Cox, 
1869, + ethologtts (Iredale, 1927); H. jacnensis Reeve, 1846, 
+ hiiiileiji Ancev, 1881; H mannonita Linnaeus, 1758, -I- gio- 
neensis Gmelin, 1791, -I- rosacea Reeve, 1846; H. pustulata 
Reeve, 1846, -I- joii.sscaiiini Mabille, 1888; H nihif^inosa 
Reeve, 1846. -I- howensis (Iredale, 1929); H nigosa Lamarck. 
1822, + miiltipcrforata Reeve, 1846, -I- reoelata Deshaves, 
186.3; H. striata Lmnaeus, 17.5S, + lanwllosa Lamarck, 1822; 
H. txibemdata Linnaeus, 1758, + incisa Reeve, 1846, -t- ja- 
ponica Reeve, 1846, -t- reticulata Reeve, 1846; H. varia Lin- 
naeus, 1758, 4- harboim Foster 1945, + drinf^ii Reeve, 1846, 
+ aernma Reeve, 1846, Haliotis neglecta Philippi, 1848, is res- 
urrected as a valid species from the Mediterranean. Lectotvpes 
for H inidtiperforata Reeve, 1846. and H revelata Deshaves, 
1863, are selected. For all titxa considered valiil the geographic 
distriliution and sources ot published illustrations are given, 
lutrecjuenth illustrated species are figured. 

Kii/ words: Gastropoda, Hahotidae, taxonomy, syiion\TOies, 
lectotypes, zoogeography. 



INTRODUCTION 

Members of the famih- Haliotidae (abalone) occur in 
most tropical and temperate oceans, particularly in the 
shallow subtidal zone. The first mention of abalone in 
the literature was made in the fourth century B.C. bv 
Aristotle. In the first century A.D. the name oiotia (fittle 
ear) was used bv Plinv. In japan, abalone were men- 
tioned as early as the fourth century A.D. In the medi- 
eval literature of Europe they were noted for the first 
time by Gessner in 1553. The first good illustrations 
were provided bv Buonanni, Lister, Gualtieri, and Ruin- 
phius between 1681 and 1741; these were later cited by 
Linnaeus (1758). The pre-Linnean descriptions of aba- 



lone have been dealt with more extensively in Crofts 
( 1929), Cox ( 1962) and Muller ( 1984a). 

Tiixonomic publications on the family began with Lin- 
naeus (1758), who described the first seven species of 
abalone using his system of binominal nomenclature. His 
work was continued and enlarged bv Gmefin (1791), 
who added 12 taxa. Reeve (1846) described 43 new taxa 
in his monograph, which is one of the most important 
sources for the taxonomv of the familv Hahotidae. In 
the late nineteenth century, three larger monographs 
were published bv Sowerbv (1882), Weinkauff (1883), 
and Pilsbrv (1890), but only a few new taxa were intro- 
duced by these authors. Wagner and Abbott (1978) pro- 
vided a hst of ta.xa incluchng tentative synonymies. Kaich- 
er (1981) illustrated all the species and subspecies she 
considered valid, providing the most comprehensive 
means available to identify the Recent Haliotidae. Pick- 
ei-y (1991) fisted most abalone taxa chronologicallv, in- 
cluding their references. Ubaldi (1993) has started to 
publish a series intended to cover, in alphabetical order, 
all extant haliotid species; to date, four species have been 
treated by Ubaldi. 

The latest valid species described is Haliotis robeiii 
McLean, 1970. Haliotis coccinea canaiiensis Nordsieck, 
1975, is the most recent taxon that has been described. 
All taxa have been based on shells; onlv the neotype of 
H. ituilatcralis Lamarck, 1822, and the types of a new 
species from Brazil are complete specimens with the an- 
imal (Geiger, 1996; Simone, in press). 

The alpha taxonomy of the majority oi species has 
been uncertain, except for most of the important com- 
mercial species. Between 30 (Dauphin ct al., 1989) and 
130 (Cox, 1962) of the more than 200 species-level taxa 
described have been considered valid species. Most au- 
thors have estimated the number of distinct species to 
be approximately 75 (Thiele, 1931; PickeiT, 1980; Kaich- 
er, 1981; Lindberg, 1992). The objective here is to reev- 
aluate this family, to critically review all the pubhshed 
information, and to include additional unpublished ob- 
servations. This groundwork is necessarv in the light of 
a forthcoming phylogenetic analvsis of the entire family, 
because the working unit — the species — should be clear- 



Page 86 



THE NAUTILUS, Vol. Ill, No. 3 



ly understood .so that tlie data matm will not be o]i- 
scured by unresolved taxonomic problem.s. 

The conclusion.s that form the ,sub.stance of this paper 
are presented in three h.sts: Index, Notes, and Valid Spe- 
cies by Faunal Regions. The alphabetical index and the 
vahd species by faunal regions are both cross-referenced 
to the notes. Species illustrated are those that are infre- 
quentlv sho\Mi in other publications. 

MATERIALS AND METHODS 

This work is based on a number ot visits to major mu- 
seums in Europe and the United States, where the avail- 
able type specimens were examined. Museum material 
was supplemented with specimens from private collec- 
tions, including those of Katharine Stewart (Carmel, Cal- 
ifornia), Don Pisor (La Jolla, C'alifornia), Roger Pickery 
(Wilrijk, Belgium), Mark Jones (Auckland, New Zea- 
land), and Benjamin Singer (Rehovot, Israel). Every 
original description was carefully checked in the original 
language. An effort was made to track all relevant sec- 
ondarv literature through later citations and the Zoo/of^- 
ical Record. Some results from ongoing studies of the 
animals are mentioned where they help in the under- 
standing of teixonomic questions; a detailed coverage of 
the characters of the animals is bevond the scope of the 
present contribution and will be pro\ided elsewhere. 
The radular terminology of Geiger (1996) is used. 

The statistical analvsis of the shell morphometrical 
data was performed with STATISTICAL Mac 4.1 
(StatSoft, 1994). For the breakpoint regression, piece- 
wise linear regression with Quasi-Newton estimation 
method and least-square loss function was employed. 
Linear regression on the data sets on either side of the 
breakqx)int \aelded the slope statistics. 

Abbreviations used are: ANSP: Academv of Natural 
Sciences, Philadelphia; BMNH: The Natural Historv 
Museum, London; DMNH: Delaware Museum of Nat- 
ural History, Wilmington, Delaware; HUJ: Hebrew Uni- 
versity, Jerusalem; KBIN; Koniglich Belgische Institut 
for Natuurwetenschappen, Bnissels; LACM: Los Ange- 
les County Museum of Natural History; LSL: Linnean 
Society London; MCZ: Museum of Comparative Zool- 
ogy, Harvard University, Cambridge, Massachusetts; 
MNHN: Museum national d'Histoire naturelle, Paris; 
MIINCi: Museum d'Histoire Naturelle, Geneve; NMW: 
National Museiun Wales, (>ar(hff; SAM: South Austra- 
lian Museum, Adelaide; SBMNH: Santa Barbara Mu- 
.seurn of Natural History, California; USNM: National 
Museum of Natural History, Smithsonian Institution, 
Washington. 

SYSTEMATIC AFFINITIES AND 
CHARACTERISTICS OF THE FAMILY 
HALIOTIDAE 

llaliotidae are part of the prosobranch Vetigastropoda, 
having the nacreous shell, certain characters in the cleav- 
age pattern, a streptoneurous nenous svstem, Kvo bi- 



pectinate ctenidia (gills) served by two heart atria (Dio- 
tocartlia), and possessing ctenidial bursicles (Hickman, 
1988; Haszprunar, 1987; 1993 for review). The row of 
tremata in abalone and the slit of the pleurotomariids 
(Pleurotomariidae) are homologous stnictures, but are 
not chagnostic characters (Delhaes, 1909; Hasz]5runar, 
1993). 

Historicallv, the Haliotidae were considered to be 
closelv related to the Pleurotomariidae and the Scissu- 
rellidae and were united with these families in the su- 
perfamily Pleurotomarioidea. The rhipidoglossan radula 
was thought to be a common character for the Pleuro- 
tomariidae and the Haliotidae. However, the radula in 
the two families differs in many ways. The rows of the 
radular teeth are almost symmetrical in Haliotidae, but 
in the Pleurotomariidae they are distinctly asymmetrical. 
The rachidian t(joth is well-formed in the Haliotidae, but 
reduced in the Pleurotomariidae. The fine outer mar- 
ginal teeth in the Haliotidae show denticulate cusps (Wu 
and Huang; 1989; Herbert, 1990; Geiger, 1996; Stewart 
and Geiger, in press). In the Pleurotomariidae, however, 
a fan of articulated bristles is fo\md (Hickman, 1984; 
Harasewych and Askew, 1993; Anseeuw and Goto, 
1996). A comparison of the Pleurotomariidae and the 
Haliotidae to the Scissurellidae is not appropriate, be- 
cause the latter are subjected to different evolutionary 
constraints due to their small size and their detrital diet 
(Fretter and Graham, 1976; Herbert, 1986), which is 
reflected in their radular structure. The radulae of ju- 
venile abalone were examined by Tong (198.5), and are 
similar to those of small Trochidae (Hickman and Mc- 
Lean, 1990), and Scissurelhdae (Marshall, 1993). The 
independence of radular morj:)hology from feeding ecol- 
ogy has to be (juestioned due to the extensive morjiho- 
logical plasticitv of the radula in response to the feeding 
ecolog)' of the respective animals. The radulae of the 
Pleurotomariidae and the Haliotidae will not help to re- 
solve their phvlogenetic relationship, because the former 
is that of a specialized spongivore, the latter that of a 
strict macroalgal herbivore. The coding of paedo- and 
peramoiphic stnictures adds further problems, as in the 
case of the radular characters of the Scissurelhdae and 
the Haliotidae. Either stage-specific stnictures are con- 
sidered, which overstate the degree of differentiation. 
The rachidian in mature animals is serrate in Scissurel- 
hdae, but bears a smooth cutting edge in the Haliotidae. 
Alteniati\ely, heterochronic processes are taken into ac- 
count, creating characters with inapplicalile character 
states. In this case the serrate rachithan tooth of animals 
<5 mm unites the Scissurelhdae and Haliotidae, but the 
rachidian characters for animiJs > 10 mm are inappli- 
cable to the Scissurellidae. The use of the radula to re- 
solve family level relationships within the Vetigastropoda 
is, therefore, questionable (see iilso Haszpninar, 1993). 

The nearly .symmetrical body plan oi Haliotis has been 
(ited as being plesiomoiphic, which situates the group 
vei-y close to the root of the flexogloss;ite prosobranchs 
(Fleure, 1904; Salvini-Plawen and Haszpninar, 1987). 
This \iew is supported In the primitive sperm ultrastnic- 



D. L. Geiger, 1998 



Page S' 




30 35 40 45 
shell length (mm) 



Figure 1. Plot ot sliell length \ersus number ot open tremata in H. tuberculata from numerous Mediterranean populations (n = 
433). Note the positive correlation between the two parameters, for which a logarithmic curve has been fitted. Note the intersection 
with the X-axis around 2 mm shell length, the size at which the first perforation is foniied. The first specimens with fi\e open 
tremata are found at 7 mm, those with six open tremata at 15 mm, those with seven open tremata at 24 mm. 



ture (Levvds ct a]., 19S0; Healy, 19S8; 1990; Healy and 
Harasewvch, 1992). However, se\'eral left parts of the 
paired body structures, e.g., the gonad and the kidnev, 
are reduced or modified in Haliotidae (Crofts, 1929; Ha- 
szpnniar, 198Sa). The nervous svstem is close to the 
primitive condition of Plciirotomaha, but in Haliotidae 
the additional structures of the epipodium and the os- 
phradium are inner\'ated bv pleural and xisceral ele- 
ments, respectiveH'. The Haliotidae and the Trochidae 
have svnapomorphic osphradi;d characters (Haszprunar, 
1985; 1993), but die Trochidae are clearlv separated 
from the ScissureUidae, Pleurotomariidae and Haliotidae 
by the lack of the right ctenidium and associated organs 
(Salvini-Plawen and Haszpnmar, 1987). 

The three famihes within the former superfamiK' 
Pleurotomarioidea are now placed in their nominal su- 
perfamilies, i.e.. the Haliotidae are now in the Haliotioi- 
dea, on the same le\el as the Pleurotomarioidea, Scis- 
surelloidea and the rem;iining Vetigastropoda (Haszpni- 
nar, 1988b: Heal\- and Harasewych, 1992). 

PROBLEMS PERTAINING TO THE TAXONOMY 
OF THE FAMILY HALIOTIDAE 

Tremata as a taxonomic character and teratological 
type specimens: Members of the famih Hahotidae 
can be easUv recognized h\ the depressed shell and the 
tremata, the row of holes used for respiration, release of 
gametes, and defecation (Ino, 1952; Tissot, 1992; Vol- 
tzow and Colhn. 1995). One mav potentially confuse 
them with some members of the StomateUinae, a trochid 
subfamiK- (Hickman and McLean, 1990; Picker>-, 1995). 
Some taxa with imperforate shells described in the genus 
Haliotis are actually StomateUinae (e.g.. H. impertusa 
Burrow, 1815), and are not dealt with here. StomateOids 
that resemble haliotids (Genu, Sfoinatella and Microtis) 
are rather small (< 40 mm), have no right ctenidium, a 
flat shell, no tremata, and no spiral sculpture. The\ 



might suggest imperforate specimens of juvenile H a.s- 
iniiia Linnaeus, 1758, but the latter have several distinct 
spiral ridges (see Kaicher, 1981), which are no longer 
formed as the shell grows larger than 3.5 to 4 cm. Spec- 
imens of the trochid genus Granata have been errone- 
ously identified as imperforate H. cijclobates Peron, 1816 
(Geiger, pers. obs.). 

In most descriptions of abalone the number of open 
tremata is indicated, erroneously suggesting that the 
number is diagnostic for identification at the species lev- 
el. However, the number of open tremata changes dur- 
ing the growth of the .shell (Hemphill, 1907; Sinclair, 
1963). The lanal shell has no tremata at all; the first one 
is formed at a size of approximately 1-3 mm (Crofts, 
1929; Muravama, 1935; Bonnot 1940; Ino, 1952; Shibui, 
1971; Mu ci ai, 1976; Bevelander, 1987). Figure 1 shows 
the size dependent change of the number of open holes 
for the European H. tuberculata Linnaeus. 1758. Linear 
break'jjoint regression (r-=0.69) estimated the break- 
point at 5.5 open holes, corresponding to a size of ap- 
proximately 33 mm, where the slope changes from 3.8 
(SE = 0.53) to 0.0017 (SE = 0.56). The number of open 
tremata varies greatly within a species and between pop- 
ulations (Hemphill, 1907; Schepman, 1909; Sinclair, 
1963; Talmadge, I960; Geiger and Groves, submitted). 
Therefore, the notion that the number of open tremata 
or total number of tremata is fixed within am' given spe- 
cies is misleading. Taxa erected solely because of differ- 
ences in the number of open tremata have to be reject- 
ed, as in the case of H. multipciforata Reeve. 1846 (note 
32), or H. crachcrodii Leach, 1814, with its subspecies 
honita Orcutt, 1900, and califonticnsi.s Swainson, 1822. 
Specimens from some populations on Guadalupe Island 
off Baja California show the morphologv' of H. cracher- 
(idii calif ornieusi.s. These specimens ha\e more and 
smaller tremata. which are also more closel)' spaced than 
in specimens from the mainland. Specimens from pop- 
ulations having the califonucnsis morphologv that have 



Page 88 



THE NAUTILUS, Vol. 111. No. 3 




Figures 2-5. Infrequentlv illustrated Ualiotis. 2. Haliotis cf. diicrsicolor Reeve. 1846. IH nini. MNHN [not numbered]. No 
localitv- data. An imperforate specimen of Haliotis. 3. H. brazieri — hargravesi. 31 mm. K. A. Stewart collection. Solitary- Island, 
Coffs Harbour. New South Wales, Australia. This specimen begins growth with the smooth morpholog\- of H brazieri. but midway 
changes to the spiral ridges typical of H. har^ravesi. 4-5. H brazieri Angas, 1869. 29 mm. MHNG [not numbered]. Australia. 



been cultured on the m;iinland of centnil California 
changed the size and spacing of the tremata to those of 
b^pical H. crachcrodii crachcrodii (B. Owen, pers. 
comm.); this indicates that these characters are under 
environmental control. 

Imperforate abalone shells have been found (Smith, 
1893; Manjuand, 1906; Hemphill, 1907; Dall, 1919; 
Leighton, I960; Geiger, pers. obs.; Figure 2), but are 
rather rare. Even rarer are specimens with a double row 
of tremata (Smith, 1888; Hamada, 1982). A somewhat 
larger number of specimens witli a continuous sUt in- 
stead of the row of tremata are known {H. laevigata. 
Gray, 1856; H. asinina, H. crachcrodii, H. parva Lin- 
naeus, 1758. H. planata Sowerby, 1882, H. nifesccns 
Swainson. 1822, H tuhcrciilata: C.eiger, pers. obs.). New 
tremata are formed at the anterior margin of the shell, 
and posteriorly the mantle eventually seals them when 
they are no longer functional. Occasionallv one to several 



tremata are closed out of the sequence just described 
(Geiger, 1991: fig. 8; H. paixa. H. pulchcrrima Gmelin, 
1791: Geiger, pers. obs. ). Only in H. clcgam Philippi, 
1844, is this phenomenon found regularlv and becomes 
the rule in large specimens (see Wilson, 1993: pi. 3, figs. 
2A. 2B). 

Haliotis impciforata Dall. 1919, H lii.sas Fmlav 1927, 
and H. holzncri Hemphill, 1907, are based on imperfo- 
rate type specimens. It is clear that these are teratolog- 
ical shells not warranting taxonomic recognition (note 
66). 

Three tspe specimens show further deformities. Hal- 
iotis .sicboldii Reeve, 1846, (note 64) has been described 
from a distorted shell, where the row of tremata is lo- 
cated much closer to the periphen of the shell. The 
same applies also to H. whitchou.sci (Colnian, 1959) 
(note 55), although the distortion is not as pronounced 
as in H. sicboldii. This t\pe of a deformation is also 



D. L. Geiger, 1998 



Page 89 



known from some specimens of H. cracheroclii (LACM 
23452; SBMNH 13522; USNM 199890). Haliotis dic- 
goensis Orcutt, 1900, (note 65) has an extremely thick 
and stout shell, a growth form that had been induced b\ 
boring organisms, most hkelv sabellid poKchaetes. 

Hybrids: The occurrence of hybrids may be challeng- 
ing to the biological species concept, but can be better 
understood from the standpoint of the evolutionar\' spe- 
cies concept. As long as the hvo species that h\bridize 
keep their identit)', they remain intact: otherv\-ise retic- 
ulate speciation takes place (cf. Wiley, 1981). When dis- 
cussing hybrids, it is assumed at the outset that the two 
parent specimens belong to Uvo discrete species, which 
nia>- not be the case. We may also have a case of un- 
realized intraspecific variabiUU' witliin a single species. 

H\bridization among the Cahfomian species is well 
determined and studied on the basis of the shell, the 
epipodium, and bv using immunological techniques, H\- 
brid specimens with intermediate shell and epipodial 
characters occurred with a frequency of approximately 
1;500 (Owen ct al.. 1971). These authors also found ev- 
idence for back crosses of hybrids with some of the pa- 
rental species, demonstrating that the Fl hybrids are 
fertile. These results were later corroborated by labo- 
ratorv rearings (Leighton, 1988). The identity of the rec- 
ognized species was confirmed with DNA sequence data 
(Lee and Vacquier, 1992; 1995). Although hybrids in- 
volving H. cracherodii were successfully reared in the 
laboratory (Leighton, 1988), these are noticeably absent 
in the natural environment. This can be accounted for 
bv two factors First, H cracherodii has an entirely 
smooth and uniformly black shell, whereas all the other 
Cahfomia species show some sculpture and may have a 
color pattern in the shell. As it is much more difficult to 
generate a new character (sculpture, color pattern), than 
to modih' an existing one, the simple condition in H. 
cracherodii prevails in hybrid specimens (overshaddow- 
ing effect). Second, H. cracherodii is the only Califor- 
nian abalone found in the intertidal region. The species 
is ecologically separated from the remaining species. 
Within the remaining sLx Cidifomian species, 13 out of 
the 15 possible combinations have been documented 
(Talmadge, 1977b; Leighton, 1988). The hvo missing hy- 
brids both involve H. fulgens Philippi, 1845, which has 
been shown in the phvlogenetic analysis of DNA se- 
quence data to be slightly more distantly related to the 
other species found in Cahfomia (Lee and Vacquier, 
1995). Prezvgotic barriers (analogous to pre-mating bar- 
riers in broadcast spawners), such as spawning season 
and vertical chstribution of the species, have to be taken 
into account. Unfortunately, no information on the fer- 
tility of F2 and backcross specimens is aviiilable, al- 
though many profound changes affecting the fertilits' 
may take place after the Fl generation (see King, 1993 
for review). 

In a second case, two Australian species H laevigata 
and H. rubra Leach, 1914, form hybrids that occur in 
nature (Anonymous, 1973; R. Fallu, pers. comm.). These 



hvo species are also well-known and have been shown 
to be distinct species (Brown, 1993; Lee and Vacquier, 
1995). In this case, hybrids have been identified on the 
basis of enzvme electrophoresis data. The hybrids occur 
in varying frequency between 0% and 2% in several pop- 
ulations studied. Good evidence has been presented for 
the presence of backcrosses of hybrids with either pa- 
rental species. The hybrids are not believed to represent 
a starting point of reticulate speciation because of the 
different habitat requirements of the two species, which 
is strongly reflected particularly in the tliickness of the 
shell (Shepherd, 1973; Brown, 'l995). 

Hybrids are reported from two sympatric populations 
of the hvo subspecies of H. discus Reeve, 1846, H. dis- 
cus discus and H. discus haniiai Ino, 1952 (Fujino et al, 
1980; Sasaki et ai, 1980). Although a case of svmpatric 
subspecies may seem a contradiction in terms, it can be 
seen as a necessary step of gradual, sympatric speciation 
(see Futuvma, 1986). As the two subspecies are very 
difficult to distinguish, enzvme electrophoresis was used 
to identib.' them with poKmorphic loci of several en- 
zymes. Later the same data and method were used to 
identifv hybrids behveen the two taxa considered. It is 
not clear whether the variabilitv of one biological species 
was rather extensive, or whether true hybrids were 
found; no data on the identification of the specimens 
studied using alternative methods had been mentioned. 

Yet another case is the least clear and has not been 
discussed in the literature to date. It involves the follow- 
ing taxa from southeastern Australia: H. hrazich Angas, 
1869, H. cthoJogus (Iredale, 1927), H. hargravcsi Cox, 
1869, and H. melcuhis (Iredale, 1927), all uncommon to 
rare. I regard H. melcuhis as a svnonvm of H. brazieri, 
and H. ethologus a synonyin of H. hargravcsi (notes 62, 
66 and 72). HaUotis hargravcsi shows extensive intra- 
specific variation in relation to number and elevation of 
spiral ribs present on the dorsal surface of the shell. 
Haliotis brazicii (Figures 4-5) has no spiral sculpture at 
all, only an uneven but smooth surface. Intergrading 
specimens, i.e.. potential hybrids, show slight spiral 
grooves, which, however, are much less distinct than any 
form within the range of H. hargravcsi. In some speci- 
mens the early whorls lack spiral cords; these will appear 
abruptly later during shell growth (Figure 3). Such pat- 
terns suggest a highly variable species with a few con- 
chological forms. The above obsers'ations do not support 
taxonomic separation of these forms. 

M. Jones (pers. comm.) noted that H. brazieri and H. 
hargravcsi are separated bv their vertical distribution. 
Haliotis brazieri occurs from the lower reef surface to 
40 m and H. hargravcsi from 40 m downward to the 
slope of the reef, where thev can be found under conil 
heads and boulders. Coleman (198:86) confirmed the 
relatively deep water habitat ("15 to 40 m") of H. har- 
gravcsi. Hybrids are most often found around 40 m, the 
tlepth vv'here the vertical ranges of the hvo species over- 
lap. Specimens of H. brazieri and H hargravcsi have 
been found under the same boulder. The epipochum of 



Page 90 



THE NAUTILUS, Vol. Ill, No, 3 



Table 1. Genus-le\'el taxa in ttie faiiiil\- Ilaliotiilae. od: original designatii 



inonohpv. sd: suli.seqnent designation. 



Taxon 



\'alid t\pe species 



Original type species (if different) 



Eiilialiotis W'enz, 19.38 

Ell rot is Habe and Kosuge, 1964 

Exohiiliotis Cotton and Godfrey, 193.3 

Haliotis Linnaeus, 17.58 

Mtirintinriti Iredale, 1927 

Miohaliotis Itoigavva and Toniida, 1982 

Neolwliotis Cotton and Godfrey, 1933 

Nordotifi Habe and Kosuge, 1964 

Notohaliotis Cotton and Godfrey, 1933 

Ovinotis Cotton, 1943 

Padollm Montfort, ISIO 

Paua Fleming, 1952 

Sanhaliotis Iredale, 1929 

Schismotis Gray, 1856 

Sulcuhis Adams and Adams, 1854 

Teiiwtis Adams and Adams, 1854 

Tinotis Fischer, 1885 

Usahaliotis Habe and Kosuge, 1964 



inUlne Linnaeus, 1758 [od] 

luhcmilata Linnaeus, 1758 [ni] 

a/clobatcs Peron and Lesueur, 1816 [m] 

asinina Linnaeus, 1758 [sd: Montfort, 1810] 

brazieri Angas, 1869 

nmabilis Itoigawa and Tomida, 1982 [m] 

scalaris Leach, 1814 [od] 

oij^nntea Gnielin, 1791 [od] 

nibro Leach, 1814 

ovinn Gmelin, 1791 [od] 

partn Linnaeus, 1758 

iris Gmelin, 1791 [m] 

vnria Linnaeus, 1758 [od] 

laevigata Donovan, 1808 

tuberculata Linnaeus, 1758 

asinina Linnaeus, 17.58 [m] 

asinina Linnaeus, 1758 [obj. syn. of Haliotis] 

cracherodii Leach, 1814 [m] 



nulcnins Iredale, 1927 [sd: Wenz, 1938] 

naevosa Martyii, 1784 [od] 
nibicundus Montfort, 1810 [m] 



excisa Gray, 1856 [not available] 

incisa Reeve, 1846 [sd: Cossmann, 1918] 



the hybrid specimens showed intermediate characteris- 
tics (M. Jones, pers. comm.). 

It is well known that abalone show gregarious spawn- 
ing behavior (Murayama, 1935; Shepherd, 1986). Hy- 
brids may appear under the circumstances described 
above. Vacquier and co-workers (Vacquier et ai, 1990; 
Lee and Vacquier, 1992; 1995; Vacquier and Lee, 1993; 
Lee ct al, 1995) have demonstrated that although the 
protein lysin from the head of the acrosomal vesicle in 
the head of the sperm strongly promotes intraspecific 
fertihzation, it does not fully prevent interspecific fertil- 
ization. Although no good data on the frequency of the 
Austrahan hybrids are av;iilable, it is evidently rather low. 
It is open to discussion whether we observe here a case 
of an erratic fertilization pattern of tyvo distinct species, 
or whether two recently diverged species still occasion- 
ally form hybrids, or whether we stantl at the cradle of 
a reticulate speciaticjn event. 

GENUS-LEVEL TAXA 

A list of all supraspecific taxa of the family HaUotidae is 
given in Table I. Comments on a few taxa are given 
here. 

Haliotis, sensu stricto: The subse()uently designated 
t\pe species (Montfort, 1810) of the genus Haliotis, H. 
asinina, has been accepted by the majority of authors 
{e.g.. Iredale, 1910; Kennard W al, 1931; Wenz, 1938; 
Talmadge, 1963a; Ubaldi, 1985; 1993). Other species 
have been cited as type species, but those citations are 
erroneous {e.g., Cossmann, 1918: H tuhcrculata. Thiele, 
1931; Cotton, 1943; Co.x, 1962: H. inidae Linnaeus, 
1758. Children/if/e Kennard rt ai, 1931: H. iris Gmelin, 
I79I). The type designation by Montfort (I8I0) was un- 
fortunate because he chos(> as t\pe species the most 
atypical species within the family. The two presumed 
type .specimens of H. asinina are deposited in the LSL, 



and correspond with what is currently known as this spe- 
cies. The hpe localitv has been designated as Amboina 
{= Ambon), Indonesia (Talmadge, 1963a). Cotton (1943: 
176) cbscussed the status of H. asinina as t\pe species 
and following Adams and Adams (1853-1858) stated 
"that asinina is the genotype of the Haliotis of Montfort 
1810 and not of the tnie Haliotis Linnaeus 1758," a 
statement in contratliction with the original te.\t. Mont- 
fort (1810:115) only introduced Padollus as a new genus: 
"... nous avons cm pouvoir en former un genre parti- 
cuher" (. . we have thought to be able to form from it 
a distinct genus). Haliotis asinina is onlv designated as 
type of the genus Haliotis on page 120: "Sous la denom- 
ination doreille de mer, tons les anciens conchyliologues 
ax'oient deja reconnu ce genre .... Nous avons choisi 
pour t)pe, an heu de I'haHotide commun (haliotis vul- 
garis, hahotis tuberculata [sic]), celui auquel on a donne 
le nom d'oreille d'ane" (Under the name of ear of the 
sea, all the old conchologists have already known this 
genus. . . . We have chosen as t\pe, instead of the com- 
mon abalone {Haliotis vulgaris, Haliotis tubcradata), the 
one, for which one has given the name of donkey's ear 
[H. asinina]). Montfort clearly designated the hpe spe- 
cies of Linnaeus Haliotis, and not a monospecific genus 
for H. asinina, because two other taxa are included un- 
der Haliotis. The fact that a common name is used in 
the text of the designation (see ICZN Article 12c) dt)es 
not invalidate it. Monfort (1810:119) used the following 
titles for the description of the species: "Espccc .seirant 
dc ti/pe ail genre. Haliotide ureiUe d'ane. Haliotis asini- 
nus." (Species sening as t\pe of the genus. Haliotid 
Donkey's ear Haliotis asinintis). First, the type species 
is clearly designated, and second, the association be- 
tween common name and scientific name is unequivoc;J. 
The name Haliotis stems from the two Greek words 
lialios (the sea) and (Uis. otis (the ear). The gender of 
the genus is feminine because Linnaeus u,sed tJie femi- 



D. L. Geiger, 1998 



Page 91 



nine ending for the specie.s-le\el taxa (leri\ed from an 
adjective, particularly for H asiiiina. the type species of 
the genus by subsequent designation of Montfort (1810). 

Other genus-level taxa, and their type species: One 

objecti\e SMionvrn (a ta\on based on the same t\pe spe- 
cies) of Hdlioti.s has been introduced, H a.sinina: Tcin- 
otis Adams and Adams, 1854. Note that The Genera of 
Recent Mollusca by Adams and Adams (1853-1858) has 
as a publication date of 1858 on the inside cover, but in 
Volume 2 (p. 661) the actual publication dates of the 
various parts were indicated. For part 14 on Haliotidae 
the date is June 1854; this latter is the correct date for 
Adams and Adams' abalone taxa. 

The genus PadoUus Montfort, 1810, with the type spe- 
cies bv monotxpv, P nihiciindus Montfort, 1810 {non H. 
nibicunda Rotling, 1798: see note 20), is problematic. 
Haliotis scalaris (Leach, 1814) has been listed as the 
type species oi PadoUus (e.g.. Knight ct ai. I960; Wag- 
ner and Abbott, 1978; Lindberg, 1992:16), but P nd)i- 
cundiis Montfort is a sMion\in of H paixa and not of 
H. scalaris (see note 20); therefore, the vaUd t\pe spe- 
cies for PadoUus is H. parva. The fact that the type spe- 
cies in the original description of PadoUus is no longer 
invalid does not invahdate the description (ICZN Rec- 
ommendation 67B). Haliotis nd)icunda (Montfort) can 
be unequix'ocallv identified and the senior syiionvm is 
the correct, vahd tvpe species. 

Neohaliotis Cotton and Godfrey, 1933, has been syii- 
onymized with PadoUus on the basis of ha%ing the same 
type species: H. scalaris (see above: Pickery, 1991). 
However, as I have shown that H. nd>icundus Montfort 
is not a synonym of H. scalaris but of H pana. the 
synonvinv oi Neohaliotis with PadoUus has to be reject- 
ed. 

The t\pe species of Sulcuhis Adams and Adams, 1854, 
is H. incisa Reeve, 1846, which is a s\nion\iii of H. tuh- 
erculata (see also note 16). As H tidwnidata has also 
been chosen as the type species of Eurotis Habe and 
Kosuge, 1964, the latter is now a subjective, junior svn- 
onym of Sulculus. 

Most genus-level taxa had their type species designat- 
ed by the original author. These designations made in 
the nineteenth centun' may not meet modem conven- 
tions for designation of a t\pe species in respect to the 
specificity of the language used. However, 1 (unlike 
Fleming, 1952) interj^ret "explicith' designated" (ICZN 
Article 67b) somewhat more generously. 

Adams and Adams ( 1853-1858) gave the genus name, 
followed bv its diagnosis and the mention of one partic- 
ular species as "Ex." (example = type species). This ex- 
ample cannot be confused with the species considered 
to belong in that genus, which were fisted further below 
in smaller print and in two columns. This form had been 
utifized for all the genera {Haliotis Linnaeus, Teim>tis H. 
and A. Adams, PadoUus Montfort), but not for the sub- 
genus Sulcidtis, where no example was given. I consider 
this constniction as an explicit designation (ICZN Article 
67b), which cannot be confused with the exception men- 



tioned under ICZN Article 67cl ("mention of a species 
as an example of a genus or subgenus") in conjunction 
with the example that follows in the Code. The narrow 
reacfing of the word "example" .shall not obscure the 
clear intentions of the authors. Fleming (1952) cfid not 
accept the designation for Teiuotis and attributed the 
subsequent designation to Cossmann (1918). 

Marinatiris was described by Iredale (1927) without 
any perceptible intention to designate either M. inelcu- 
his or M ethologus as the type species; the genus was 
introduced in an extremely casual form. Fleming (1952: 
229) made claims for the designation ("Ty])e (here des- 
ignated) '), but had overlooked the clear designation by 
Wenz (1938:172). 

Schisinotis Gray, 1856, has been given in some fists 
(Pickery-, 1991; Ubaldi, 1993). Gray\fid not intend to 
introduce a new name for the specimens he cfiscussed 
as monstrosities of H. albicans Quoy and Gaimard, 1834 
(= H. laevigata), but only indicated a suitable, hypo- 
thetical name. The name is a nomen nudum and is not 
available (see also note 70). 

Haleotis Binkhorst, 1861, has been cited as an objec- 
tive synionvm of Haliotis, i.e., is treated as an available 
emendation of Haliotis (e.g.. Knight ct ai, 1960). An 
emendation according to ICZN 33b(i) and 33b(iu) must 
be 'demonstrablv intentional," otherwise it is considered 
as an incorrect subsequent spelfing (ICZN 33c), which 
is not a\iulable. As there was no demonstrable intent of 
the author to emend the original spelfing, the name 
must be considered as unavailable. Tinotis Fischer, 1885, 
however, is available as the author has shown a clear 
intent to emend the name (Fischer, 1885:845): "Tinotis 
H. et A. Adams, cm. (Teinotis)." It is an objective syn- 
onym of Haliotis. 

Use of genus-level taxa: Some authors (e.g., Tal- 
madge, 1963a; Habe and Kosuge, 1964; Hara and Fujio, 
1992) have used several genera in the family Hafiotidae. 
Pickery (1991) provided a fist and all references for the 
Recent taxa, which have been ranked as genera or sub- 
genera (Table 1). As pointed out recently (Geiger, 1996), 
I consider the usage of these genera to be unjustified at 
this time for the following reasons: 

1) In the descriptions of the one fossil (Itoigawa and 
Tomida, 1982) and the 17 Recent supraspecific taxa only 
the t\pe species had been assigned, occasionalK- with 
selected species listed. Of the 200 species-level taxa only 
approximately 83 have ever been assigned to any su- 
praspecific taxon, and 22 of those to more than one 
group, demonstrating the problematic supraspecific tax- 
onomy of the hafiotids. The descriptions of these genus- 
group taxa are entirely typological and no author has 
attempted to pro\ide serious differential diagnoses. 

2) Only tvvo stucfies have utilized s\stematic characters 
other than shell moipholog\ to detemiine the relation- 
ships of h;ifiotid species. Brown (1993) stutfied 17 spe- 
cies using enzyme electrophoresis, and Lee and Vac- 
quier (1995) used cDNA sequences of the sperm acro- 
.somal protein lysin (see Vacquier and Lee, 1993 for re- 



Page 92 



THE NAUTILUS, Vol. 111. No. 3 



view) of 22 haliotid.s. The nominal .supraspecific taxa and 
the hmited number of associated species are not in ac- 
cordance with the groups hypothesized by these more 
recent studies. However, the t\pe species ni H allot is, H. 
asinina (see above for discussion), was not included in 
either study, making any sound taxonomic decisions im- 
possible. 

3) The only inferable consensus in the literature (Mc- 
Lean, 1966:151-153; Undberg, 1992) as well among ab- 
alone researchers (Workshop "Evolutionary Biologv- and 
Genetics of Abalone" during the Second International 
Sviiiposium on Abalone Biology, Fisheries and Culture, 
Febniars 1994, Hobart, Tasmania) was not to use any 
genus-level taxa other than Haliotis. Yet there was an 
equally strong consensus among the workshop partici- 
pants that the diversity within the family may well justify 
the recognition of several genus-level taxa. However, 
monophvletic groups will have to be determined from a 
phylogenetic study encompassing the majority of haliotid 
species. 

SPECIES-LEVEL TAXA 

Use of subspecies: Subspecies are defined as allopa- 
tric populations with a fixed character (Futuyma, 1986). 
Thev do not yet represent discrete, evolutionan,- lin- 
eages. Interbreeding at the periphery of these popula- 
tions is not necessarily a sign of the erroneous applica- 
tion of the subspecific classification, but will show that 
the populations are not yet independent species or dis- 
crete lineages. Only in very few shallow-water, broad- 
cast-spawning gastropods have subspecies been de- 
scribed, but in HaUotis these have been invoked a num- 
ber of times (Lindberg, 1992), i.e., in the following 
groups: H. di.^cii.s. H. mariac Wood, 1S28, H p\tst\data 
Reeve, 1846, H. ntbra, H. scalaiis, H. fiiberculata, H. 
varia, H. virginea Gmelin, 1791, and the populations of 
Califomian haUotids occurring on Guadalupe Island off 
Baja California, Mexico. In most ca.ses the subspecific 
division is found along a temperature gracUent; i.e., sub- 
species are described from chfferent latitudes and not 
from different longitudes. Most continental coast fines 
nm in a north-south direction, which mav be used to 
explain the above observation. On the other hand, the 
Indo-Malayan Archipelago offers rich opportunities for 
the developent of isolated populations, but no subspe- 
cies have been proposed for species with a wide longi- 
tudinal range such as H. asinina (Thailand-Fiji), H. 
clathrata Reeve (East Africa-Samoa), H. ovina (Mal- 
dives-Tonga), or H. planata (Thailand-Fiji). The excep- 
tions to the latitudinal pattern are Kvo southern Au.stra- 
Uaii species, H. .scalaiis and H. nihra. 

In H. (li.'icus, the caryotypes differ between the two 
subspecies (Nakamura, 1985), although their populations 
interbreed at their common boundary (reviewed in Fu- 
jino, 1992). In this case the two taxa are more likely to 
represent subspecies than ecomorjihs. 

The two formerly recognized subspecies of H. tiiber- 
atlata — H. tiihcrrulata fiihcrnilata and H. fuhrrnihita 



laniellosa Lamarck, 1822 — have identical carv'otvpes 
(Colombera and Tagfiaferri, 1983), and have been re- 
cently shown to have identical sequences of the lysin 
protein (Lee and Vacquier, 1995; see also note 5). 

In New Zealand, four geographically separated sub- 
species of//, virginea Gmefin are reported (note 60): H. 
virginea virginea, H. virginea cmpata Gould, 1847, //. 
virginea moiioria Powell, 1938, and H. virginea huttoni 
Filhol, 1880 (Kaicher, 1981; Ubaldi, 1986). These sub- 
species show gradual changes in sexeral moqjhological 
characters used to distinguish them due to environmen- 
tal parameters associated with geographical location 
(Talmadge, 1957a). M. Jones (pers. comm. ) has found 
some other forms on the very remote islands off New 
Zealand that seem to be more stable and distinct than 
the subspecies mentioned above. In these two cases the 
described subspecies seem to be ecomorphs not jusH- 
fving taxonomic separation. However, as no hard data is 
available for the subspecies of H. virginea and, because 
they are widely used, thev are here retained. 

None of the Califomian subspecies is retained be- 
cause in those cases for which data is available (//. crach- 
erodii califoniiensis, H. cornigaia dicgocnsis: see above 
and note 65), they have been shown to be mere eco- 
morphs or teratological specimens. One former species 
has been allocated at the subspecies rank (H. kamtschat- 
kana assimilis D;dl, 1878: see McLean, 1966; Owen et 
al, 1971; note 67), 

Two pairs of subspecies are found in southern Austra- 
fia. These are longitudinally separated: the eastern H. r. 
nihra with the western H. rubra conicopora Peron, 1816 
(see note 53), and the eastern H. scalaiis einmac Reeve, 
1846, with the western H. s. .scalaiis (see note 56). In- 
formation other than shell morphology (Shepherd, 1973; 
Brown, 1993) suggests one polymorphic species in both 
instances. However, the shells can be cfistinguished rea- 
sonably well, and the geographic cfistribution of the 
moiphs is more or less disjunct. The a\';ulable data sug- 
gests subspecific status for the respective populations. 

Due to the limited information currenth axiiilable, the 
last three cases are provisionally studied as follows: H. 
inariae dentata Jonas, 1846, is easily recognized by the 
deep hirrows resulting in the denticulated anterior mar- 
gin of the shell, which are missing in the nominate sub- 
species. The biogeographical data on the species is scant. 
No assessment of geographic variation is possible. I re- 
tain the two subspecies of //. mariac. 

HaUotis pustulata cnicnta Ree\e, 1846, has a reddish 
coloration, a \ei"S' Hat shell, and is found particularly in 
the Red Sea. HaUotis pustulata pustulata on the other 
hand is more sculptured, usually dark green to mud col- 
ored, and is fo\md along the east African mamland. 
There are no striking differences bet^veen the animals, 
and molecular data on the two morphs is not \et avail- 
able. I tentatively retain the two subspecies. However, 
as their relation to H nigo.sa Lamarck, 1822, is currently 
uin-esolved (see note 12), they may eventually be treated 
as subspecies of H. mgosa, or may be united in a single 
species without further division into subspecies. 



D. L. Geitjer. 1998 



Pase 93 



Table 2. Differences between tlie \alid taxa of this studv with their status in Wagner and Abbott (1978) and iHustrations in Kaicher 
(1981), Subspecies rank is indicated h\ indentation nnder the respecti\e species. The following taxa were regarded eis \a!id bv 
Wagner and Abbott (1978) bnt are here varionslv SMion\niized (see Index for details): caitarien.sis, dringii. ^ibha. hanlet/i. Iiowensis. 
japonica. kraiissi. tnhcradata laincllosa. nwlcuhis. muUiperforata, snlco.sa. vixlirata. ichitehousei. Forms listed In' Kaicher (1981) 
have been ignored unless specifically indicated. The following taxa shouni by Kaicher (1981) are either variously synonyinized here, 
or she illustrated specimens that cannot be identified (see Index tor details): bistriata, cornigata oiceni, dissona. elevata. gemma, 
multiperforata. sepicidata, ichitehousei. 



This stnd\ 



Wagner and Abbott (1978) 



Kaicher (1981) 



hrazieri Angas, 1869 
crachcrodii Leach, 1814 
clathrata Reeve, 1846 
divcrsicoli'r Ree\e, 1846 
dohniiana Dunker, 186.3 
planata Sowerbw 1882 
liargravesi Cox, 1869 
jacnensis Reeve. 1846 
kamtschatkana Jonas, 1845 

assimilis Dall, 1878 
marine dentata Jonas, 1846 
madaka (Habe, 1977) 
mannorntn Linnaeus, 1758 
neglccta Philippi, 1848 
ovina Gmelin, 1791 
pusttilatei Reeve, 1846 

cnienta Reeve, 1846 
roei Gray, 1826 
ntbiginosa Reeve, 1846 
nibra Leach, 1814 

conicopora Peron, 1816 
nigosa Lamarck, 1822 
scalaris emmae Peron, 1816 
speciosa Ree\e, 1846 
squamosa Gray, 1826 
tuberculata Linnaeus, 1758 

coccinea Reeve, 1846 
uniJateralis Lamarck, 1822 
larin Linnaeus, 1758 

virginea Gmelin, 1791 
crispata Gould, 1847 
huttoni Filhol, 1880 
tnoriorin Powell, 19.38 



brazieri 

craclwrodii 

as rubra clathrtita 

not mentioned 

"no information" 

planata 

hargravesi 

jacnensis 

kamtschatkana 

assimilis 

not mentioned 

not mentioned 

undetermined species 

not mentioned 

ovina 

pustulata 

not mentioned 

roei 

symonym of varia 

rubra 

conicopora 

syiiouNin of tidwrculata 

cmmac 

synonym of fulgens 

squamosa 

tuberculata 

coccinea 

unilatcralis 

varia 



virginea 

as form of virginea 
as fomi of virginea 
as fomi of virainea 



also as melculus 

including all ssp. 

also as crebrisculpta auct. and venusta 

also as aquatilis and all ssp. 

dohniiana 

also as graijana 

also as ethologus 

also as hanleiji 

also as aulaea 

assimilis 
mariae dentata 
not mentioned 
as guineensis 
not mentioned 
shows cijclobates 
also as revelata 
cnienta 
also as sulcosa 

as varia rubiginosa, Iiowensis 
also as ancile 
conicopora 

as pustulata form alternata 
emmae 
speciosa 

cf diversicolor shown 
also as lamellosa 
also as zealandica 
varia shown 
also as unilatcralis. dringii and ssp, papulata. 

pustulifera. viridis 
also as gibba 
crispata 
huttoni 
morioria 



For H. varia. seven subspecies are regularly found in 
the literature, but none are recognized here. As the 
name suggests, this species is e.xtremelv xarialile. I have 
been unable to find any geographic pattern, and inter- 
mediate specimens abound. These ta.xa are clear e\a- 
dence of the typological thinking of the nineteenth cen- 
tury. 

Index: In this section I give mv opinion of the taxo- 
nomic status of every taxon ot Recent Hidiotidae. Spell- 
ing mistakes, enoneous dates, and incorrect ta\on-au- 
thor combinations have not been included, unless uide- 
spread confusion exists. Controversial opinions and new 
assessments of the taxonomic status ot various taxa are 
indicated in the "Notes" that follow. The notes are ar- 
ranged according to their appearance in the section "Val- 
id Species by Faunal Region" below, i.e.. are grouped bv 
valid species. 



The tAvo most recent, but brief, taxonomic assess- 
ments oi the family Hahotidae were proxided liy Wagner 
and Abbott (1978) and Kaicher (1981). All differences 
between the present study and the opinion of these au- 
thors are listed in Table 2. 

An analysis of all the species-level taxa reveals that 
only 27% of described taxa are still considered valid spe- 
cies. The status of the sub.species (5%) is very much 
debated, as the unit of a subspecies is somewhat vaguely 
defined as a geographically limited population with a 
certiiin character, raising the percentage oi all \alid taxa 
to slightly less than one-third (see also discussion below). 
Taxa that had originally lieen described as either forms 
or varieties are excluded from modern taxonomy, and 
represent what we generally call ecomorphs. I do not 
use Latin form names in an informal fashion (contra 
Reid, 1996), as all aviiilable evidence for Haliotidae sug- 



Page 94 



THE NAUTILUS, Vol. HI, No. 3 




Figures 6-9. Infrequentiv illustrated Haliotis. 6-7. Haliotis multiperforata Reeve, 1846. 63 mm. BMNH. Mus. Cuming. Lec- 
totype here selected. 8-9. H rcvclata Deshayes, 1863. 58 mm. MNHN. Bovirhon. Lectotvpe here selected. 



gests that such taxa do not constitute discrete gene 
pools. The invalid taxa — sviionyms (46%), homonyms 
(5%), nomina duhia (6%), nomina inula (2%), eco- 
morphs (6%), unavailable names (3%) — account for over 
two-thirds of all pulilished species-level t;«a. 

The status of some taxa is still unclear; some mav nev- 
er be fully resolved due the loss of \^f^e material, in 
other cases there is currently too little material available 
to make a definitive assessment. Unresolved cases are 
mentioned here. The type of H. cunuliculata Fischer, 
1907, was lost prior to 1872 (Ivanov and Kantor, 1991) 
and its .synonvniy with H. paina is uncertain (note 19). 
The following taxa can only be tentatively synonyniized, 
because the tvpe material remains to be located and the 
original description and/or illustration do not allow a 
clear assessment: crenata Swainson, 1822, glabra Swain- 
son, 1822, sciitulum Reeve, 1846 (notes 47, 48), sepi- 
culata Reeve, 1846, and sinuata Perry, 1811 (note 22). 
Seven taxa cannot be identified and are treated as nom- 
ina dubia: adriatica Nardo, 1847, bistriata Gmefin, 1791 
(notes 5, 14), irnperforata Gmelin, 1791 {not Dall, 1919) 
(note 66), intcrntpta Valenciennes, 1831 (note 73), par- 
ina Valenciennes, 1831 (note 73), plicata Ciinelin, 1791, 
and rotundata Perrv', 1811 (note 22). The taxa mandata 
Kuster, 1840 (note 18), maculosa Kiister, 1840 (note 18), 



modcsta auct. (note 75), secen\enda Monterosato, and 
schroctch Menke (note 76) are treated as nomina nuda, 
because they could not be traced to the original source. 
The deposited type specimen of H. victoiiac Brazier is, 
in the absence of an original description, a nonwn nu- 
dum (note 77). H<dioti.s ncglccta Philippi, 1848, is ten- 
tatively resurrected but the material a\;iilable is \en' Lim- 
ited and restricted to shells only (notes 5, 8, 9). Haliotis 
exigua Dunker, 1863, is tentativelv retiiined as a valid 
species (note 63). 

Lectotvpes are here selected for hvo taxa: H. multi- 
perforata Reeve, 1846 (note 32), and H. rcvclata De- 
shayes, 1863 (note 12). The figured t)pe specimen of 
unknowai provenance of H multipciforata. which is here 
designated as lectotvpe (Figures 6-7), is clearly a spec- 
imen of H. ntgosa Lamarck (note 12), however, with 
rather weak spiral sculpture. The two other specimens 
in the lot are H. tubcrculata. The number of open tre- 
mata of these specimens is somewhat higher than the 
usual, but not bevond the expected and documented 
range of the latter species (data not shown). Conse- 
(juentlv a taxonomic separation is not justified. 

Haliotis rcvclata Deshaves, 1863, was described from 
lie de Bourbon (= Reunion Lsland). The MNHN holds 
three lots of tvpe material of this species with a total of 



D. L. Geiger, 1998 



Page 95 



six specimens. One lot with a single specimen is labeled 
as "ex auteur, " another of three specimens is labeled as 
"syntypes, " and a third lot with two specimens is called 
"type." From the labeling of the specimens, it is not 
clear which one may be the holotype; therefore, all six 
specimens are presumed to be .syntypes. Deshayes' orig- 
inal illustrations (plate 36, figs. 1 and 2), were meticu- 
lousK drauai, but cannot be matched to anv ot the shells 
in the MNHN. In fact, the illustrations do not resemble 
any of the species found on Reunion Island, possibly due 
to excessive artistic freedom. The rather long description 
lacks much necessaiT detail. Onlv two quantitatixe in- 
dications are of some help. The length of the largest 
specimen (61 mm) and the cited six open perforations 
are applicable to one specimen in the third lot, but this 
specimen has irregular growth on the coliunella that is 
clearly not illustrated or mentioned. The illustration 
shows only a weak growth Une at the level of the pen- 
ultimate hole, which is also found in the second speci- 
mens of the third lot. This specimen is here designated 
lectotvpe (Figures 8-9), the remainder become paralec- 
totypes. Haliotis revelata is identified as H. nigosa La- 
marck, a species common at the type locahty. The status 
of H. nigosa Lamarck itself is not resolved (see also note 
12). 

Haliotis fatiii is an unpubhshed name by the late H. 
Rehder (USNM) given to specimens that somewhat re- 
semble H vaiia Linnaeus, 1758. Ubaldi (1993) hsted H. 
fatiii with the authority' "Rheder [sic]. 1981 ?". This hst- 
ing is a nomcn nudum (see also note 71). 

In the index that follows, the taxa with their status 
and/or the currently recognized valid species name are 
given. The taxa in boldface are currently recognized as 
valid species and subspecies with the original genus giv- 
en if different from Haliotis s.s. The numbers in brackets 
refer to the notes that follow. Abbrexaations: juv.: juve- 
nile shell; s.L: sensu lato; ssp.: subspecies; syn.: synonym; 
var.: \ariet\', form; ?; uncertiiin statement (status, iden- 
tification). 

adriatica Nardo, 1847; nonien dubium 

albicans Quoy and Gaimard, 18.34: laevigata Dono\an, 1808 

alfredensis Bartsch, 1915; speciosa Reeve, 1846 

alicna (Iredale, 1929) [in Sanhaliotis]: varia \ar. paptdnta 

Reeve, 1846 
aleata Roding. 1798; aiistralis Gnielin, 1791 
altcmnta .Sowerbv, 1882: n/f^o.sn Lamarck, 1822 [.31] 
ancde Ree\e, 1846: jm; nibra Leach, 1814 [52] 
aquatilis Reeve, 1846: diversicolor Reeve, 1846 
asitiina Linnaeus, 1758; valid [4] 
asinum Donovan, 1808: asinina Linnaeus, 17.58 
assimiUs Dall, 1878: ssp. of kamtschatkana Jonas, 1845 [67] 
a\tricfa Reeve, 1846: var. of varia Linnaeus, 1758 
aulaca Bartsch, 1940: kamtschatkana assimilis Dall, 1878 [67] 
sp. nov. Simone, in press: \ahd [1] 
aiistralis Gmelin, 1791: \alid 
bar!>oini Foster, 1946: varin Linnaeus, 1758 [1] 
bistriata Gnielin, 1791: noiucn dubium. tubcrculata Linnaeus, 

1758, s.l. [5, 14] 
bistriata Costa, 1829; homonym, van, s\ai. of tuberculata var. 

lamellosa Lamarck. 1822 



bisundata Monterosato: tuberculata var lamellosa Lamarck, 

1822 [15] 
bonita Orcutt, 1900: crachcrodii \ar, californiensis Swaiiison, 

1822 
brazieri Angas, 1869; \aiid [.34, .35] 
caelata Roding. 1798: ovina Gmelin, 1758 
califoniiana Valenciennes, 1831: nifcsceus Swainson, 1822 
californiensis Swainson, 1822: var., s\ii. of cracherodii Leach, 

1814 
canaliculata Fischer, 1807: paixa Lnmaeus, 1758 ? [19] 
canaliadata Lamarck, 1822: honionvm, parva Linnaeus, 1758 
canadensis Nordsieck, 1975: tuberculata coccinea Reeve, 1846 
capensis Dnnker, 1844: midae Linnaeus, 1758 
carinata Swainson. 1822: parva Linnaeus, 1758 
cingulata Roding, 1798: parva Linnaeus, 17.58 
clathrntn Lichtenstein, 1794: elegans Philippi, 1844 [25] 
clathrata Ree\'e, 1846: valid (homonym) [24, 25] 
coccinea Ree\e, 1846: ssp. of tuberculata Linnaeus, 17.58 [17, 

18] 
coccoradiata Reeve, 1846: \';ilid [49] 
concinna Reeve, 1846; varia Linnaeus, 1758 
conicopora Peron, 1816: ssp. of rubra Leach, 1814 [.5.3] 
coreanica Weinkauff 1883: nomen nudum [68] 
cornigata Wood, 1828: valid [26[ 
costata Swainson, 1822; aiistralis Gmelin, 1791 
cracherodii Leach, 1814: valid 
crebrisculpta Sowerby, 1914: valid [25, 36] 
crcnata Swainson, 1822; scalaris Leach, 1814 ? 
crispata Gould, 1847; ssp. of virginea Gmelin, 1791 [60] 
cntenta Ree\e, 1846: ssp. of pustulata Ree\e, 1846 [10] 
cunninahaini Gray, 1826: rubra conicopora Peron, 1816 [27, 

53] 
cyclobates Peron, 1816: valid 
dalli Henderson, 1915: vahd [2] 

decussata Philippi, 1850; marmorata Linnaeus, 1758 [7] 
dentata Jonas, 1846: ssp., var. of nmriae Wood, 1828 
diegoensis Orcutt, 1900: var of corrugata Wood, 1828 [65] 
discus Ree\e, 1846: \alid [62] 

dissona (Iredale, 1929) [m Sanhaliotis]: vahd [36, .39] 
diversicolor Reeve, 1846: valid [37] 
dohmiana Dunker, 1863: valid [40] 
(lrinfj.ii Ree\e, 1846; varia Linnaeus, 1758 [45] 
diibia Lamarck, 1822: nomen dubium [69] 
echinata Sowerbv, 1882: jacnensis Reeve, 1846 [31, 41[ 
elatior Pilsbiv. 1890: \'ar. of midae Linnaeus, 1758 
elegans Philippi, 1844: \alid [23, 50] 
elcvata Sowerby, 1882; squamata Reeve, 1846 [31, 57] 
emmae Reeve, 1846: ssp. of scalaris Leach, 1814 [56] 
ethologus (Iredale, 1927): hargravesi Cox, 1869 [.34] 
excavata Lamarck, 1822: ci/clobates Peron. 1816 
excisa Gray, 1856: unavailable, laevigata Dono\an, 1808 [70] 
exigua Dmiker, 1877: valid [63] 
expansa Talmadge, 19.54; cracherodii Leach, 1814 
fatui Rehder, 1981: unavailable [71] 
ficifonnis Menke, 1844: spadicea Doiio\an, 1808 
fidgens Philippi, 1845: valid 
funebris Reeve, 1846: squamata Reeve, 1846 [58] 
gemma Reeve, 1846: varia Linnaeus, 1758 [46] 
gibba Philippi, 1846: virginea Gmelin, 1791 [61] 
gigantea Chemnitz, 1788: unavailable, gigantea Gmehn, 1791 

[59. 64] 
gigantea Gmehn, 1791: valid [64[ 
gigas Roding, 1798: gigantea Gmelin, 1791 
glabra Chemnitz, 1788: unavailable, glabra Gmehn, 1791 [59] 



Page 96 



THE NAUTILUS, Vol. Ill, No. 3 



glabra Gnielin, 1791: valid 

afahra Swainson, 1822: homonviii, laevigata Donovan, 1808 ? 

granti Pritchard and Gatliff, 1902: rubra conicopora Peron, 

1816 [53] 
granulata Rotling. 1798: varia Linnaens. 17.58 
grai/ana Sowerb)', 1882: plaiiata Sowerb)-, 1882 [31] 
gniiicri Philippi, 1848: \ar., ,svii. of diiersicolor Reeve, 1846 
guadalupensis Talmadge, 1964: \ar. offtilgens Philippi, 1845 
guinecnsis Gmelin, 1791: mannorata Linnaeus, 1758 
hanlei/ana Sowerby, 1882: notion dubiitm [31, 72] 
luinlrip .-^ncev, 18S1. jacnensis Reeve. 1846 [41] 
hannai Ino, 1953: ssp. of discus Gnielin, 1791 
hargravesi Co.x. 1869: valid [34, .35] 
hattorii Bartseh, 1940: \ar. of nifcscens Swainson, 1822 
holzneri Hemphill, 1907: \ar. of crachcrodii Leach, 1814 [66] 
howensis (Iredaie, 1929) [in Snnhnliotis]: ndnginosa Reeve, 

1846 [43] 
huttoni Filhol. 1880: ssp. of virginea Gmelin, 1791 [60] 
iinpciforata Gmelin, 1791: nomen diibium [66] 
imperforata Dall, 1919: honionvm, crachcrodii Leach, 1814 

[66] 
improbula Iredale, 1924: s\ii., \ar. of rid)ra Leach, 1814 
incisa Reeve, 1846: tuberculata Linnaeus, 1758 [16] 
intemipta Valenciennes, 1831: nonwn dubium [73] 
iridis Karsten, 1789: unavailable, iris Gmelin, 1791 [74] 
iris GmeUn, 1791: valid [.54] 
jacnensis Reeve, 1846: valid [41] 

janiis Reeve, 1846: tiibcrcidata coccinca Reeve, 1846 [17] 
japonica Reeve, 1846: tuberculata Linnaeus, 1758 [16] 
jousseaumi Mabille, 1888: pustulata Reeve, 1846 [11] 
kamtschatkana Jonas, 1845: valid [62, 67] 
kraussi Turton, 1932: parvia Linnaeus, 1758 
laevigata Donovan, 1808: valid 
lamellosa Lamarck, 1822: var. of tuberculata Linnaeus, 1758 

[5, 13] 
latilabris Philippi, 1848: oiina Gmelin, 1791 [8] 
lauta Ree\e, 1S46: seiniplicata Menke, 1843 [49] 
lucida Requien, 1848: tuberculata \ar. lamellosa Lamarck, 1822 
lusus Finlay, 1927: var of crachcrodii Leach, 1814 [66] 
maculafa Kiister, 1840: nomen dubium [18] 
maculosa Kiister, 1840: nomen dubium [18] 
madaka (Habe, 1977) [in Nordotis]: valid [64] 
mariae Wood, 1828: \alid [26] 
mannorata Linnaeus, 17.58: valid [4, 5, 6] 
mannorata Ree\e, 1846: honioiiMn, virginea Gmelin, 1791 
mclculus (Iredale, 1927) [in Marinauris]: brazieri Angas, 1869 

[35] 
midae Linnaeus, 1758: valid [04] 

modesta auct.: nomen nudum ?, midae Linnaeus, 1758 [75] 
morioria Powell, 19.38: ssp. of virginea Gnielin, 1791 [60] 
inullipeijornta Reeve, 1846: nigosa Lamarck, 1822 [.32] 
naevosa Phihppi, 1844: rubra Leach, 1814 [54] 
nebulata Reeve, 1846: nigosa Lamarck, 1822 [33] 
neglecta Philippi, 1848: valid ? [5, 8, 9] 
nodosa Philippi, 1845: comigafa Wood, 1828 
ovina Gmelin, 1791: vahd 

oweni Talmadge, 1966: var of corrugata Wood, 1828 
papidata Reeve, 1846: var of varia Linnaeus, 1758 
parma \'alenciennes, 1831: nomen iluliium [73] 
parva Linnaeus, 1758: valid [4] 

parva Risso, 1826: homonviii. tubemdala \ar lamellosa La- 
marck, 1822 
pellucida von Salis. 179.3: tuberculata \'ar lanvUosa Lamarck, 
1822 



perfusa Ree\e, 1846: rugosa Reeve, 1846 [.33] 

picta Rikling, 1798: glabra GmeUn, 1791 

planata Sowerby, 1882: valid [42] 

planilirata Reeve, 1846:/(//gcn.s Philippi, 1845 [47] 

plicata Karsten, 1789: unavailable, australis Gniehn, 1791 [74] 

plicata Gnielin, 1791: nomen dubium 

ponderosa Adams, 1848: nifescens Swainson, 1822 

pourtaleHti Dall, 1881: \alid [1, 2, 3] 

pulcherrima Gnielin, 1791: valid 

pustulata Reeve, 1846: vahd [12] 

pustulifera Pillsbrv, 1890: varia Linnaeus, 1758 

queketti Smith, 1910: valid 

reticulata Reeve, 1846: tubemilata Linnaeus, 1758 [16] 

revea Bartseh, \'^4(): fulgens. nomen nudum 

revelata Deshayes, 1863: rugosa Reeve, 1846 [12] 

roberii McLean, 1970: valid ]2] 

roedingi Menke, 1844: squamosa Grav, 1826 [29] 

roei Gray 1826: valid [27] 

rosacea Reeve, 1846: mannorata Linnaeus, 1758 [6] 

rosea Orcutt, 1900: crachcrodii Leach, 1814 



rotundata Perrv, 1811: 



en dubiu 



nibicunda Roding, 1798: pana Linnaeus, 1758 [20] 
nibicnnda (Montfort, 1810) [in Padollus]: homoimii, pana 

Linnaeus, 1758 [20] 
rubiginosa Ree\'e, 1846: \alid [43] 
rubra Leach. 1814: vahd [25, 51] 
rufescens Swainson, 1822: \'alid 
rugosa Lamarck, 1822: valid [12] 

rugosa Reeve, 1846: homonyiii, tuberculata Linnaeus, 1758 
nigosoplicata Chemnitz, 1788: una\ailable, australis Gnielin, 

1791 ].59] 
nigosoplicata Reeve, 1846: australis GmeUn. 1791 [.59] 
sanguinea Hanley, 1840: spadicea Donovan. 1808 [21] 
scalaris (Leach,' 1814) [in Padollus]: valid [.56] 
scabricostata Menke, 1843: roei Gray, 1826 
schroeteri Menke: nomen dubium [76] 
scutulum Reeve, 1846: varia Linnaeus, 1758 ? [47, 48] 
secernenda Monterosato: tuberculata \ar lanwllosa Lamarck, 

1822 
semiplicata Menke, 1843: \alid 
seinistriata Ree\'e, 1846: varia Linnaeus, 17.58 
sepiculata Ree\e, 1846: diversicolor Reeve, 1846 ?, tuberculata 

Linnaeus, 1758 ? 
sicboldii Reeve, 1846: gigantea GiiieHn, 1791 164] 
sinuata Penv, 1811: spadicea Dono\an, 1808 ? [22] 
smithsoni Bartseh, 1940: kamtschatkana assimilis Dall, 1878 
sorenseni Bartseh, 1940: valid 
spadicea Donovan, 1808: valid [21] 
speciosa Reeve, 1846: \'alid 
splendi-ns Reeve, IH46: fulgens Philippi. 1845 
splendidula Williamson, 1893: crachcrodii Leach, 1814 
squamata Reeve, 1846: \alid 
squamosa Gray, 1826: valid [27, 28] 
stomatiaefonnis Reeve, 1846: squamata Ree\e, 1846 [39] 
striata Linnaeus, 1758: tuberndata \ar lamellosa Lamarck, 

1822 [4. 13] 
strigata Weinkanff, 1883: mannorata Lmnaeus, 1758 
subvirginea Weinkanff, 1883: virginea Gnielin, 1791 
sulcosa Philijipi. 1845: roei Gray, 1826 
supeiiexta Lischke, 1870: var, svii. of diversicolor }{eeve. 1846 

[38] 
tayloriana Reeve, 1846: var, syii. of diversicolor Reeve, 1846 
trieostalis Lamarck, 1822 : scalaris Leach, 1814 
tricostata Wood, 1828: scalaris Leach, 1814 



D. L. Gei^er. 1998 



Page 97 



tuberculata Linnaeus, 1758: valid [4. 5, 1.3] 

tiibifcra Lamarck, 1822: gigantea Gmelin, 1791 

tun eh Bartsch, 1940: var. offtilgcns Philippi, 1845 

uniUtteralis Lamarck, 1822: valid [30] 

varia Linnaeus, 1758: valid [4] 

varia Risso: homon\Tn, tubcmdata \ar. lainellosa Lamarck, 

1822 [4, 44] 
venusta Adams and Reeve, 1848: clathrata Reeve, 1846 [24] 
victoriae Brazier: nomen nudum ?, rubra Leach, 1814 [77] 
virginea Gmelin, 1791: \alid [60] 

virgiiwa Reeve, 1846: homon\ni. mannorata Linnaeus, 1758 
viridis Reeve, 1846: varia Linnaeus, 1758 
vixlirata (Cotton, 1943): rubra conicopora Peron, 1816 [53] 
vulgaris da Costa, 1778: tuberculata Linnaeus, 1758 
walallensis Steams, 1898: valid 
wliitehousei (Colman, 1959) [in Sanhaliotis]: rubra Leach, 

1814 [55] 
zealandica Ree\e, 1846: coccinea Reeve, 1846 [17] 
ziczac Reeve, 1846: glabra Gmehn, 1791 



NOTES: 

01 Haliotis barbouii Foster, 1945, has been a verv con- 
troversial species. It was described from a single, 
beach shell from the coast of Brazil. It has been 
hvpothesized that is is either a distinct species, or a 
mislocated specimen of either H. potirtalesii Dall, 
IS81, from the Caribbean (Rios, 197.5) or the only 
PolvTiesian species, H. pulchcrrima Gmehn, 1791 
(Cox, 1962; Rios, 1985, 1994). Haliotis barbouii has 
not been included in the discussion of Brazilian Hal- 
iotidae bv Simone (in press). Klappenbach (1968) 
mentioned a li\ing specimen taken off Brazil, refut- 
ing the claim the species represents speciments left 
by tourists (cf. Cox, 1962), Specimens of H. pour- 
talcsii have been indicated from tiie Gulf of Mexico 
to as far south as Venezuela and Brazil (Henderson, 
1915; Foster, 1946; Agua\o and Jaunie, 1947; Harn; 
1966; Guice, 1968; Klappenbach, 1968; Sarasiia, 
1968; Nijssen-Meyer, 1969; Titgen and Bright, 1985; 
Ode, 1986: Martinez and Tuiz, 1994). the more 
southern reports of H. pouiialcsii including Klap- 
penbach's (1968) live specimens can be attributed to 
confusion \\ith a new species described (Simone, in 
press). 

Inspection of the tvpe of H, barbouri (MCZ 152469) 
revealed its true identitv as a somewhat aberrant 
form off/, varia Linnaeus, 1758. At a size of 22 mm 
it is a rather small specimen for the species. In gen- 
eral, small specimens of abalone tend to be rounder 
than larger ones (cf. Stewart and Geiger, in press: 
fig. 4). Hence, based on the overall shape of the 
shell one ma\- be led to regard the tvpe of H. bar- 
bouri as closelv related to H. pulchcrrima. Howe\er, 
the sculpture differs greatlv Most significantK, in H. 
pulchcrrima a narrow spiral band adjacent to the 
row of tremata and devoid of oblique radial ele- 
ments is found near the suture. A similar band is 
preseiU in H. jacncnsis Reeve, 1846 (cf. Figure 16, 
note 4l). In H. varia and the type ot H. barbouri 



this bare space is absent. The tvpe of H. barbouri 
somewhat evokes the t\pe specimens of H. gemma 
Reeve, 1846 (= H. varia: see note 46) in terms of 
.size, rotundity, and sculptural elements. 

02. Haliotis (/a//; Henderson. 1915, and H robcrii Mc- 
Lean, 1970, are two small, insular species found off- 
shore from Central America at the Galapagos Is- 
lands {H. dalli) and Cocos Island {H. roberii). The 
two species are fairly similar but can be distin- 
guished as indicated by McLean (1970). Radulae of 
both species share a very narrow lateral tooth 1 and 
concentric rings on the cephahc tentacles. These 
characters are shared only with H. pourialcsii Dall, 
1881, and the new species to be described from Bra- 
zil (Simone, in press), but is not seen in any of the 
other 30 abalone species studied so far (Geiger, 
pers. obs. ). 

03. The neotvpe of H. pourialcsii Dall, 1881, is now 
broken in man\- pieces. The original illustration of 
the neot\pe (Henderson, 1915: pis. 4.5—46) showed 
an intact shell. 

04. When Linnaeus (1758) introduced the first seven 
hahotids (H. asinina, H. mannorata, H. midac, H. 
paixa. H. striata, H. tuberculata, H. varia). the con- 
cept of tvpe specimens was not vet established. The 
LSL holds specimens in the collection, and I agree 
with Talmadge (1977a) that most correspond well 
with the current concept of the respective species. 
Talmadge (1977a) noted a single specimen of H. 
tuberculata. whereas I found sLx specimens with a 
note h\ S. P. Dance from 1963, also referring to six 
specimens. No specimens are currently designated 
as types. As indicated by Talmadge (1977a), there is 
no specimen of H. parva in the LSL. Designation 
of a neotvpe is unnecessarv because the species does 
not present any nomenclatural problems (ICZN Ar- 
ticle 75b). 

05. The H. tuberculata group contains the following 
main taxa: H. tuberculata Linnaeus, 1758, lamcUosa 
Lamarck, 1822, coccinea Reeve, 1846, bistriata 
Gmehn, 1791, and possibly also H. mannorata Lin- 
naeus, 1758 (c/ Talmadge, 1963a) (Figures 18-19) 
and H. ncglccta Phihppi^ 1848 (Figures 20-21). Of 
the well-known taxa in the group H. tuberculata was 
the earliest to be named; the group name is attrib- 
uted to this species. All the species are found in 
Europe and northwest Africa (Mediterranean, Brit- 
tany to Sao Tome and Gabon, Canary Islands, 
Azores). cDNA analysis of the protein Ivsin (Lee and 
Vacquier, 1995) has showni that tuberculata and la- 
mcUosa from the Mediterranean are the same spe- 
cies. The tuberculata and the lamellosa forms are 
usually found within one population (Geiger, pers. 
obs. ). The taxon lamellosa, therefore, refers to a 
variety or ecomorph of H. tuberculata. Haliotis coc- 
cinea was shown to be closelv related to H. tuber- 
culata (Lee and Vacquier, 1995). It differs form H. 
tuberculata in onl\' four positions of the 132 amino 
acid sequence whereas most species differ in ap- 



Page 98 



THE NAUTILUS, Vol. Ill, No. 3 



proxiniatelv 15-20 amino aciil positions. Despite the 
small difference in amino acitl secjneiue, the shells 
are readily distinguishable and H. coccinea is found 
only at the Canar\- Islands. Two conditions warrant- 
ing the subspecific status of H. tiihcrculota coccinea 
are present: distinct character and specific geo- 
graphic location. From shell moqihologv observa- 
tions, H. bistiiata seems to be a variation of H. tub- 
ereiilata scnsu lata (see note 14). Haliotis mannor- 
atu Linnaeus, however, seems to be distinct in terms 
of shell morphology- (Figures 18-19), but neither an- 
atomical characters, nor biochemical data are av;iil- 
able. Its status in relation to H. tiiherculata remains 
unresolved. 

06. The specimens labeled H. iiiiiniunata Linnaeus, 
1758, are conspecific with what is best known as H. 
rosacea Reeve, 1846. This latter has been synony- 
mized with H. guineensis Gmelin, 1791 (Talmadge, 
1963b; Ubaldi, 1987) (Figures 18-19). Despite the 
possibihty of exchange of material (K. Way, pers. 
comm.), three hnes of evidence suggest that the 
specimens are likely to represent the species as de- 
scribed bv Linnaeus: 1) Sowerby (1882) already in- 
dicated the synonymy between H. mannorata Lin- 
naeus, H. rosacea, and H. guineensis (explanation to 
plate 11, figs. 88, 89); 2) some old specimens of this 
species labeled H. mannorata Linnaeus have been 
found in collections [e.g.. DMNH 011036); 3) S. P. 
Dance stated on a label from 1963 that "from sev- 
eral undocumented shells I have selected one that 
matches specimens in Brit. Mus. (Nat. Hist.) labeled 
'H. rosacea Reeve. This is almost certainly the ex- 
ample mentioned by Hanley as present." Therefore, 
the correct name for the continental west African 
species is H. marmorata Linnaeus. Talmadge 
(1977a), in his discussion of Linnean haliotids, er- 
roneously svnonvinized H. mannorata Reeve (non 
Linnaeus) with H. virginea Reeve (non Gmelin). 
Haliotis mannorata Reeve is clearly H. virginea 
Gmehn from New Zealand and not H. mannorata 
Linnaeus (= H. virginea Reeve) from west Afiica. 

07. Haliotis decussata Philippi, 1850, was described in 
Phihppi's (1847-1851) third volume, with the date 
of the volume given as 1851. However, the pages 
with the description oi Haliotis are dated April 1850, 
which is the correct date. 

OS. Philippi referred to figure 4 Un his H. latilahris Phi- 
hppi, 1848, and figure 5 for H. neglccta Philippi, 
1848; this is obviously an error and one should con- 
sult figures 5 and 4 for these species, respectively. 
Pickery (1991) indicated 1851 as pubfication date, 
but the species had been already described by Phi- 
lippi (1848:16) prior to that date. 

09. Philippis (1847-1851) illustration of//, neglecta Phi- 
Uppi, 1848, is similar to green and brown mottled 
H. scjuamata Reeve, 1846, which led Sowerby (1882: 
27) to synonymize these two taxa: ". . . H. neglecta 
of Philippi ... is undoubtedly identical with our 
present species [H. .stomatiaejormis]" (see also note 



39). However, the tvpe localitv of H. neglecta is Si- 
cily in the Metliterranean Sea, whereas H. scjuamata 
is an Australian species. Ubaldi (1987) indicated this 
species as being distinct, occurring on islands close 
to Sicily, Malta and Lampedusa, but without illus- 
trating any specimens. Some specimens that corre- 
spond very well to the figured specimen have been 
located: one in MNHN (Sicily and Palermo), four in 
HUJ (6313a, b: Lampedusa and Giardini), one in 
DMNH (097371: near Malta Island), and one in my 
personal collection (Malta: Figures 20-21). Haliotis 
tiiherculata Linnaeus, 1758, with the form lamellosa 
Lamarck, 1822, is the only native, well known, and 
highly variable species in the Mediterranean. Hali- 
otis neglecta might just represent aberrant speci- 
mens of H. tubcrculata (cf. Weber, 1928). However, 
the specimens 1 have seen are very distinct and un- 
likely to be only a variation of H. tiiberculata. I con- 
sider H. neglecta to be a valid species, but it would 
be desirable to compare the epipodium, the radula, 
and DNA sequence data. 

Interestingly, a similar case is found in the Conidae. 
Conns ventricosus Gmehn, 1791, is a species li\ing 
throughout the Meditenanean. In the Sicily Chan- 
nel area, C. vaijssicri Pallary, 1906, is present (Villa, 
1985), but this species has also been regarded as a 
mere form of C. ventricosus (Poppe and Goto, 
1991). 

10. Haliotis pti.stulata cntenta Reeve, 1846, migrated 
from the Red Sea into the Mediterranean Sea 
through the Suez Canal. It has been found along 
the Israeli coast (Talmadge, 1971; Fainzilber, 1984) 
and in Lybia (Giannuzzi-Savelli et ai. 1994). One 
alcohol-preserved specimen without the shell from 
Greece shows a more or less identical epipodium 
when compared to specimens of H. pustnlata cnien- 
ta from the Red Sea. This latter is rather chfferent 
from H. tubcrculata Linnaeus, 1758, from the Med- 
iterranean. It is, therefore, possible that this species 
has advanced further into the eastern Mediterra- 
nean. The few reports in the fiterature of this mi- 
grant species in contrast to others — e.g., Stromhus 
cleconis (Roding, 1798) (see e.g., Fischer, 1993; 
Lindner, 1993) — may be due to the fact that the 
shells of the Red Sea species often are not strikingly 
different from the native H. tubcrculata. 

11. The geographic provenance of H. jousseaumi Ma- 
bille, 1888, was not given in the original description 
(Mabille, 1888), but the label of the specimen held 
in the MNHN cites the Red Sea ("Mer Rouge"). 
The name has only been used once thereafter by 
Tidmadge ( 1956), who correctly s\nionymized it with 
// pustnlata Reeve, 1846. 

12. A lectotvpe for H. rcvelata Deshayes, 1863, is here 
selected (see main body of te.xt. Figures 8-9). The 
specimens are conspecific with what is better known 
as H. nigosa Lamarck, 1822. Herbert (1990) spec- 
ulat(»d about the .sviionvmy between H. pustnlata 
Reeve, 1846, and H. nigosa Lamarck, something 



D. L. Geiger. 1998 



Page 99 



that hatl alreacK' been hinted bv Wagner and Alibott 
(197S) in their sviionvinization ot H. altcniata Sow- 
erby, 1882, with H. pustulata. I have not seen any 
material that has intermediate characters, and ques- 
tion the SMion\iiiv between the two ta\a. The t;ixa 
cnienta Reeve, 1846, and pustulata may eventually 
be treated as a subspecies of H. nigosa Lamarck, 
pending further clarification based on their respec- 
tive anatomies, but I tentativelv consider them as 
here a distinct species (cf. Herbert, 1990; Geiger, 
1996). 

13. Haliotis striata Linnaeus, 1758, belongs in the H 
tuhcrculata Linnaeus, 1758 group (see note 5). I 
agree with Talmadge (1977a) that it corresponds 
with what is well kuowii as H. tuhcrculata var. la- 
mellosa Lamarck, 1822 (Figures 28-29). If H tuh- 
crculata is a svnonvm of H. lamcllosa. then H striata 
is also a sviionym of H. tuhcrculata. Haliotis tuhcr- 
culata is preferred over H. striata as the v;ilid name 
for the species because it is the established name. 
However, if taxonomic distinction between H. tuh- 
crculata and H. lamcllosa is favored, then H. striata 
has priorit)' over H. lamcllosa. Suppression of H. 
striata would be desirable, as pre\iousIy suggested 
by Pilsbiy( 1890:87). 

14. Weber (1928) mentioned the extensive variability of 
H histriata Gmelin, 1791, from Tenerife, Canars- 
Islands, some of which correspond with its originiil 
description, some that are typical H. tuhcrculata 
Linnaeus, 1758, and concluded that the two taxa are 
most hkel\- conspecific. He did not mention H. coc- 
cinea Reeve, 1846, and possiblv was synouMuizing 
too much under one taxon (see also note 9). The 
taxon histriata is mostly encountered in the old ht- 
erature, and a few specimens from old collections 
labeled H. histriata could be found in the BMNH 
and ANSR Most of the specimens that show some 
resemblance to old illustrations (e.g.. Reeve, 1846: 
pi. 11, fig. 33) originated from the Canary Islands 
and the Azores. The specimens are characterized by 
strong radi;J grooves and a flared aperture. A sep- 
aration of H. histriata from other members of the 
H. tuhcrculata group (see note 5) cannot be ac- 
cepted. The localities, illustrations, and specimens 
labeled as H. histriata do not show any coherent 
pattern. Haliotis histriata should be regarded as a 
growth form of either H. tuhcrculata coccinca for 
specimens from the Canai"v Islands and the Azores, 
or of H tuhcrculata or H marmorata in the case of 
specimens from the East Atlantic m;iinland. As H. 
histriata most Ukelv refers to three different species- 
level taxa, I prefer to treat it as a iiomcn duhium. 
which further provides stabilits to the nomenclature, 
because the populations from the Canary Islands are 
well knov\ai under the epithet coccinca. 

15. Haliotis hisundata Monterosato is hsted as a variety 
of//, tuhcrculata Linnaeus, 1758, in Priolo (1948) 
and Ghisotti (1964), but cannot be traced. 

16. Haliotis incisa Reeve, 1846, H. japonica Reeve, 



1846, and H. reticulata Reeve, 1846, are all reported 
from Japan. This type locality is (|uestionable and 
the type specimens look exactlv hke the Mediterra- 
nean H. tuhcrculata Linnaeus, 1758, as also indicat- 
ed bv Reeve (1846) and discussed in Dunker (1882), 
Weinkauff (1883), McLean (1966), and Kaicher 
(1981: card no. 2882). Weinkauff (1883:59) never- 
theless reported for H. japonica similar specimens 
"without any doubt" from Japan, but not giving the 
locality record more credibility. In modem times no 
similar specimens have been reported from the well- 
stuched area of Japan. Haliotis japonica and H. tuh- 
crculata var. lamcllosa Lamarck, 1822, ha\e also 
been .svnonvmized erroneously (Pilsbry, 1890:87) 
with H. aquatilis Reeve, 1846, this latter being a 
synonyin of H. diversicolor Reeve, 1846. 

17. The epithet coccinca Reeve, 1846, is assigned as a 
subspecies of//, tuhcrculata Linnaeus, 1758. This is 
here established after examination of cDNA se- 
quencing data (Lee and Vacquier, 1995), distinct 
shell characters, and its isolated geographic occur- 
rence at the Canary Islands (see also note 5) Hali- 
otis janus Reeve, 1846, is a color form of //. tuh- 
crculata coccinca (Talmadge, 1958). 

18. Haliotis maculata Kiister, 1840, and H. macido.sa 
Kiister, 1840, (spelhng variations?) are mentioned 
by Sowerby (1882:36,' pi. 9) and Weinkauff (1883: 
83) as senior svnonpns of H. coccinca Reeve, 1846. 
The name is based on figure 137 in Martini and 
Chemnitz (1769) (non-binominal), which shows verv 
clearly H. tuhcrculata coccinca (see note 17). The 
original source could not be traced, and Kiister's taxa 
were not fisted in Sherbom (1922; 1932). According 
to Weinkauff, Kiister's name has priority over 
Reeves, an opinion not followed here, because the 
citation of Kiister's work could not be located. 

19. The type specimen of//, canaliculata Fischer, 1807, 
was lost prior to 1872 (Ivanov and Kantor, 1991). 
Fischer (1807) indicated the presence of a deep, spi- 
riil canal and a brownish shell with many spiral cords 
in this small species. These characters point toward 
H. parva Linnaeus, 1758, a species to which he cfid 
not refer. The svnionvmv is tentative. 

20. Haliotis ndncunda (Montfort, 1810) has also been 
attributed erroneously to Gray (1826) (see also note 
27). Gray clearly referred to Montfort. Montfort 
used this species as tvpe species for his genus Pa- 
dollus and did not refer to any other author, which 
makes it fikelv to be a new tiixon introduced bv him 
and not //. nihicunda Roding, 1798 (synionym of//. 
parva Linnaeus, 1758). Haliotis nd)icunda (Mont- 
fort) was inflected to the mascufine by Montfort for 
his new genus Padollus (see Knight ct al.. I960; 
Picken,-, 1991; also Schremp, 1981 1125). The type 
locahtv is the "African coasts." The key feature of 
the species is the spiral ridge on the shell; a char- 
acter seen only in //. parva among the African spe- 
cies (Iredale, 1927:334). Padollus nd)icundus Mont- 
fort has also been synonyiTiyzed with //. scalaris 



Page 100 



THE NAUTILUS, Vol. Ill, No. 3 



(Leach, 1814) (see also note 56), a species that, how- 
ever, occurs onlv in Australian waters. Because H. 
nibicunda Rocling and P nihicundus Montfort refer 
to the same species, they must lielong to the same 
genus, for which the species have to take the correct 
generic ending of the adjectival species name, caus- 
ing P. nibictindus Montfort either way to become a 
secondary- homonvm. Note that although PadoUus is 
based on an invalid name, this does not make Pa- 
doUus itself in\'ahd. 

21. Holioti.s sanguiuca Hanley, 1840, was reported also 
to have been described in 1808, in the same year as 
the senior synonvm H. spadicca Donovan, 1808. 
MuUer (1986) discussed the date of publication of 
H. sanguinea at length and concluded that it was 
actuallv described in 1840. 

22. The descriptions of H. rotundata Perry, ISll, and 
H. sinuata Perry, 1811, given by Perry (1811: pi. 52) 
are very brief and general, and his figures are rather 
styUstic. Pilsbrv (1890) listed both as unidentified 
species. The undulation of the apertural margin as 
well as the general shape of the shell of H. sinuata 
is reminiscent of H. spadicca Donovan, 1808. 

23. Haliotis clathrata Lichtenstein, 1794, has so far 
been considered a nomcn duhium (Wagner and Ab- 
bott, 1978). This causes H. clathrata Reeve, 1846, 
to be an invalid, junior, primary homonym. Haliotis 
clathrata Lichtenstein is discussed by Geiger in 
press and identified as H. clcgans Phihppi, 1844. 
Geiger and Stewart (under consideration) have pe- 
titioned the International Commission on Zoological 
Nomenclature to suppress H. clathrata Lichtenstein 
to presei-ve H. elegans and H. clathrata Reeve. This 
position, pending decision by the Commission, is 
provisionally adopted here (ICZN Article SO). 

24. Haliotis venusta Adams and Reeve, 1848, has so far 
been considered chstinct from H. clathrata Reeve, 
1846, due to the absence of radial lamellae on the 
type specimens. Several characters can be obsen'ed 
on both series of type specimens of H. clathrata 
Reeve and H. venusta (1 holotype, 2 paratypes each 
in BMNH): 1) a pattern of appro.\imately 4-.5 faint 
growth lines followed by one stronger one; 2) tre- 
mata only slightly oval, but rather large in compar- 
ison to the shell; 3) all shells of orange and white 
color, despite some fading; 4) in both series, some 
specimens with spire fully visible in ventral position, 
some only partially; 5) numerous spiral cords; 6) 
usually three to four of these cords stronger than 
others; stronger cords regularly spaced between su- 
ture and row of tremata; 7) the Indo-Pacific type 
locahties for both ta.xa: Baclayon, Island of Bohol, 
PhiUppines for H. clathrata Reeve, and Eastern Seas 
for Haliotis venusta. Haliotis clathrata Reeve is dis- 
tinguished from H. venusta bv the discrete, numer- 
ous radial lamellae in the respective holotvpes. The 
lamellae of //. clathrata Reeve are formed along the 
stronger growth lines, i.e., every four to five faint 
growtli lines The lamellae of // ctatlirafa Reeve 



represent elevated, strong growth lines in H. ven- 
usta, hence the two structures are homologous. Ra- 
chal lamellae are known to appear randomly in pop- 
ulations of a single species oi Haliotis. The best doc- 
umented case is that of H. tuberculata Linnaeus, 
1758, from the Mediterranean, where the lamellate 
form is well-known as H. lamcllosa Lamarck, 1822, 
H. tuberculata lainello.sa, or H. tuberculata form la- 
mcllo.sa (Ubakh, 1987; Poppe and Goto, 1991). In a 
large series of over 400 specimens all intermediate 
forms could be found within any given population 
(Geiger, unpubl. data). The two taxa have been 
shown to have an identical caryohpe (Colombera 
and Taghaferri, 1983) and an identical cDNA se- 
quence of the acrosomal sperm protein lysin (Lee 
and Vacquier, 1995). Although the material for H. 
clathrata Ree\'e and H. venusta is not as abundantly 
available as for the Mediterranean species, I am 
convinced that a similar case can be made for the 
Indo-Pacific taxa in question. I, therefore, svnony- 
mize H. venusta under H. clathrata Reeve. 

25. Haliotis crcbri.sculpta Sowerby, 1914, has been s\-n- 
onyinized with Haliotis clathrata Reeve, 1846 (Ku- 
roda and Habe, 1952). It has also been Usted as a 
subspecies of H rubra Leach, 1814 (Talmadge, 
1957b). Both opinions are rejected; the validity- of 
the species is discussed in Stewart and Geiger (in 
press) and the reader is referred to this work for an 
in-depth treatment (see also note 36). 

26. Haliotis corrugata Wood, 1828, has also been attrib- 
uted to Gray. However, Wood (1828) published a 
figure with the name of H. comigata in the supple- 
ment t(j his Index Testaceologicus. Gray apparently 
supplied some of the material, on which the figures 
were based (p. iiv of Supplement), but Wood actu- 
ally pubhshed the figure; he is consequently the au- 
thor of the taxon. 

27. Serious confusion about the date of publication of 
Gray's (1826) work is found in the literature. The 
publication dates indicated are 1826 and 1827. The 
work was edited in two volumes, where volume 1 is 
dated 1827, but one copy of volume 2 that I have 
seen is dated 1826 and another copy is dated 1827. 
Generidly, 1826 is accepted as the correct date, be- 
cause it is the earliest confirmed date of pubfication. 

28. Haliotis squamosa Gray, 1826, was erroneously de- 
scribed from Australia, but actiudly occurs in a re- 
stricted area in southern Madagascar The species 
was rediscovered in Madagascar bv Dautzenberg 
(1932) and again by Stewart" (1984).' 

29. Haliotis roedingi Menke, 1844, was described by 
Menke (1844:97) in "2 neue Haliotis Arten, bes- 
chrieben vom Herausgeber" (2 new species of Hal- 
iotis, described by the editor). According to Menke, 
the species was alreadv kiiown and named by Chem- 
nitz; Menke gave the name "Haliotis Roechngi. 
Chemn." It is clear that the species was knowii to 
Chemnitz, but was described by Menke. Based on 
the description as well as the localits' ((/ Pilsbry, 



D. L. Geiger. 199S 



Page 101 



1890: 112, note 28). I agree vvith Stewart (1984) that 
Menke, 1844, is the author and that H. rocdingi is 
a synonym of H. .squamosa Gray, 1826. 

30. The taxonomic status of Haliotis uiiilatcralis La- 
marck, 1822, has been much disputed (Figures 32- 
33). Geiger (1996) designated a neotvpe and dis- 
cussed the taxon in detail. Menke (1830:88) listed 
this species. Mention of the species in Casto de 
Elera (1896) in his catalog of shells from the Phil- 
ippines is (juestionable. This latter author also men- 
tioned species kmown at the time not to occur on 
the Philippine Islands. The best example is H. mar- 
iae Wood, 1828, a species endemic to Oman. 

31. Sowerbv's abalone taxa from the Thcsaunis Conclu/- 
lioniin ha\'e been dated 1883 (Abbott and Dance, 
1983) or 1887 (Pickery, 1991). The individual vol- 
umes were issued in a number of parts {cf. British 
Museum (Natural Historv), 1915). Volume 5 was 
published bet\veen 1882 and 1887. The hahotids ap- 
peared in the first part of Volume 5 in 1882, which 
is the correct date for these taxa. It is not a printing 
date, as most taxa are listed in the Zoological Record 
of 1882 (Martens, 1882). 

32. A lectotype for H. midtipciforata Reeve, 1846, is 
here selected (see main bodv of text, Figures 6-7). 
The synonymy with H. spadicea Donovan, 1808 (see 
Wagner and Abbott, 1978) is certainly wrong. 

33. Haliotis pciiiisa Bartsch, 1915, and H. nchulata Tur- 
ton, 1932, are indicated in Barnard (1963) as syn- 
onyms of H. spadicea Donovan, 1808. I assume that 
Barnard intended to indicate H. pciiiisa Reeve, 
1846, sensH Bartsch (1915) and H. nebulata Reeve, 
1846, scnsu Turton (1932). The tvpes of H. peiiusa 
and H. nchulata are clearly referable to H. rugosa 
Lamarck, 1822 (see also note 12). 

34. Haliotis hargravesi Cox, 1869, and H. ethologus 
(Iredale, 1927) are uncommon to rare in southern 
Queensland and northern New South Wales. White- 
head (1981:5) specifically noted for H. hargravesi 
that "knowm localities are as for H ethologus. " The 
only distinguishing characters between these two 
taxa is the number of spiral ribs and their elevation. 
However, these characters seem to be rather vari- 
able, pointing to extensive intraspecific variabilits', 
and indicating presence of a single, polymorj^hic 
species. The hvo taxa represent slight \ariations 
within the morphological range when a large enough 
sample is examined. 

35. The tvpe ot H. inelculus (Iredale, 1927) is severely 
chipped, has verv elevated tremata and hardh' an\- 
spiral ribs: it suggests H. brazieii Angas, 1869. Wil- 
son (1993) figured a specimen with many thin spiral 
threads as H mclcuhis. I regard it as a specimen in 
the H hargravesi Cox, 1869 — brazieii continuum 
(see also note 34 and Hybrids section above). 

36. For H. crebrisculpta Sowerby, 1914, three syntypes 
are known in the BMNH, the NMW, and in the 
USNM. The specimen in the BMNH is very similar 
to die figured specimen in Sowerby (1914); the 



specimens in the NMW and the USNM represent 
H. clathrata Reeve, 1846 (see notes 24, 25). The 
specimen in the BMNH has been designated as the 
lectotype (Stewart and Geiger, in press). Haliotis 
dis.soua (Iredale, 1929) is here considered a vahd 
species (cf. note 39; Figures 10-11), although 
Whitehead (1981) listed it as a synonym of H. cre- 
hrisculpta. Haliotis dis.wna has also been s)'nony- 
mized by Wagner and Abbott (1978) with H. div- 
crsicolor Reeve, 1846, and its forms (Figures 12- 
13), an opinion I strongly oppose. The characters 
common to H. dissona and H. divcrsicolor (more or 
less smooth shell with spiral ridges) are due to the 
txpe specimen of H dissona being badl\' worn and 
rather small. The distinct elongated shape of the 
shell of H. dissona with rather deep spiral ridges and 
grooves in fresh specimens sets it apart from H. div- 
crsicolor The illustration of Iredale (1929) exagger- 
ated the sculpture of the shell to a great extent; it 
may be considered a reconstructive drawing of a 
fresh shell from a worn specimen. 

37. Haliotis divcrsicolor Reeve, 1846, is well known 
from the temperate northwestern Pacific, from cen- 
tral to southern Japan (Lindberg, 1992) and some- 
what further south. The species has now been found 
in a small number oi independent lots Irom Bah and 
New Caledonia, but onlv from locaUzed upwelfing 
areas (Figures 12-13). UpweUing areas are thought 
to exist on many of the surrounding islands that may 
provide habitat for this temperate species (S. A. 
Shepherd, pers. comm.). The shells are more elon- 
gated in general shape than tvpical ones, and are as 
highk' arched as H. squamata. Additionally, they are 
mostly of uniform, dark sepia coloration, which may 
be interpreted as a case of melanism. Melanistic 
shells are well kiiown in some gastropods such as in 
the Cvpraeidae, and New Caledonia is known for a 
higher frequency of such dark colored specimens. 
Whether these tropical populations of H. divcrsico- 
lor represent refugia or have only been colonized in 
geologically recent time is not known. A taxonomic 
separation seems inappropriate. 

38. Two spelling variations are found in the Uterature: 
H. supci-tcxta Lischke, 1870, and H. supratexta. The 
original speUing was H. supertexta. 

39. Talmadge (1962) treated H stomatiacformis Reeve, 
1846, as a subspecies of H. varia Linnaeus, 1758. I 
disagree vvith his conclusion after inspection of the 
t\pe materiid: H. .itomatiaefonnis is a typical H. 
.squamata Reeve, 1846 (see note 57). Specimens 
normally identified as H. .stomatiaefonnis are usually 
referable to H. di.ssona (Iredale, 1929) {cf. note 36). 

40. Haliotis dohrniana Dunker, 1863, is a httle known 
species with thstinct affinities to H. varia Linnaeus, 
1758, as already indicated bv Dunker (1870:7). The 
epipodium shows distinct characteristics (Geiger, 
pers. obs.). 

41. Haliotis hanlei/i Ancev, 1881, and H. jacnensis 
Reeve, 1846, have been considered to be distinct 



Page 102 



THE NAUTILUS, Vol. Ill, No. 3 




Figures 10-15. IiilrecjueiitK illustrated llaliotis. 10-11. Haliotis dissona (Iretlale, 1929). 33 mm. R. Picker,' collection. New- 
Caledonia. 12-13. H. diversicolor Reeve, 1846. 30 mm. ANSP 319655. Bali Beach Hotel. 14-15. H. exigua Dunker, 1877. 21 mm. 
NMW 1955. 158. 2133. Japan. 



species (e.g., Talmadge, 1963a; Kaicher, 1981). The 
distinguishing characters were thought to he the 
general shape of the shell, and the extent to which 
the shell bears cords. These characters are highly 
variable and the two taxa represent slight variations 
within the uiorj')hological spectnnn of one biological 
.species. The geographical distribution of the two 
taxa is congruent. The smooth, dorsal part of the 
shell proximal to the row of tremata is a synapo- 
morphy with H. ptilchcniina (Jmelin, 1791 (see note 
1), and a diagnostic- character of the species (Figures 
16-17). 
42. Sowerby inchcated Caipeuter as authority l(jr //. 



planata Sowerby, 1882. Weinkauff (1883:76) could 
not find the source; he indicated "tibi?" (Latin for 
where?'). Todav Sowerbv is usualK- credited with 
the authorship of the taxon. Haliotis planata is oc- 
casionalK' confused with H. varia Linnaeus, 1758. I 
agree with Talmadge (1963a) that chstinctive epi- 
podial ni()q-)liologies separate the two species. 
43. Haliotis nihi^nosa Reeve, 1846, has most often 
been used as a .synonym, form, or subspecies of H. 
varia Linnaeus, 1758 {e.g., Kaicher, 1981). Haliotis 
/i()irc((.v(.v (Iredale, 1929) was described and is en- 
demic to Lord Howe Island. Old specimens labeled 
as H. niliii'iuo.sa from Lord Howe Island were lo- 



D. L. Geiger, 1998 



Page 103 




Figures 16-21. InlifijuentK illustrated llaUoti.s. 16-17. Haliotisjacnensis Ree\e, 1846. 12 iiiin. R. Pickery collection. No location 
data. 18-19. H inannonita Linnaeus, 1758. 63 mm. LSL. Man- africanus. 20-21. H. neglecta Philippi, 1848. 29 mm. CoUectiou 
Geiger AAB 51a. Malta Island, 



cated in the BMNH (K. Stewart, pers. comm. ) and 
in the HUJ (12557). The types of the two species 
are ven* similar, and I agree with K. Stewart that the 
two names refer to the same species. Haliofis ni- 
bi^inosa has priority over H. houcnsi.s, despite the 
latter being better knowni (Figures 26-27). 

44. The tvpe locahh' of H. vaiia Linnaeus, 175(S, is Phil- 
ippine Islands as designated by Iredale (1910). 

45. It is unclear whether or not H. diinj^ii Reeve, 1846, 
represents a distinct species or a form of H. vaiia 



Linnaeus, 1758. As H varia is such a xariable spe- 
cies, I am inchned to synonymize H. dringii under 
H. varia. Many specimens identified in collections 
as H dringii are actuallv H. jacncnsis Reeve, 1846 
(Figures 16-17). 
46. Haliotis gemma Reeve, 1846, is quite certainly a col- 
or form of H. varia Linnaeus, 1758, as seen from 
the series of four type specimens in the BMNH, 
with one specimen marked "tvpe." The svTion\iTii- 
zation is not entirely certain as the shells are rather 



Page 104 



THE NAUTILUS, Vol. Ill, No. 3 




Figures 22-27. liifrequentK illustratt'd Ihiliolis. 22-23. Haliotis phmata Scmvrln, 1SS2. 22. .39 mm. NMW 19.5,5. 158. 2124. 
Pliilippine.s. 23. .3.3 mm. NM\\' 19.55. 158. 2125. Guadalcanal. 24-25. H (pirkctti Smith, 1910. .33 mm. NMW 1955. 158. 2129. 
Pondoland. 26-27. H. nibimnosa Reeve, 1846. 25 mm. USNM 791422. Lord Howe Island. 



.small; juvenile ahalone are notoriou.slv difficult to 
identify, and H. varia i.s an extremely variable spe- 
cies. The selection of the specimen labeled "tvpe" 
is (juestionahle because the measurements of the 
shells and the indications in the description of the 



species do not match. The illustration of H. gemma 
is "magnified double" and the illustration is 19. 5 
mm long; hence, the corresponding specimen 
should be ;i little less than 10 nun long. The speci- 
nu'Ti labled "t\pe" measures 20. 5 mm, which is 



D. L. Gei^er, 1998 



Page 105 



twice the size of the originid specimen. I believe that 
the reference to "magnified double" in the original 
description was incorrect. Kaicher (1981) illustrated 
another, unidentifiable specimen as H <icm»ui. 

47. I found no t\pe specimens for H. planilirata Reeve, 
1846, or H. saitulum Reeve, 1846, in the BMNH. 
Talmadge (1964) indicated a t\pe of H planilirata 
in the BMNH. 

48. From the description "wa\'ed, here ami there larg- 
er," tlie coloration 'oli\'e brown . . . dotted and spot- 
ted with green," and the irregular spiral ridges with 
occasional thickenings to be seen in the figure 64 of 
Reeve (1846), I tentati\elv identify H. scutuluin 
Reeve, 1846, as H. laria Linnaeus. 1758 (cf. note 
47). 

49. Haliotis laiita Reeve, 1846, was tentatively placed in 
the s\non\ni\- of H. coccoradiafa Reeve, 1846, by 
Talmadge (1960). Alter inspection of the t\pe spec- 
imens of both species in the BMNH, it is clear that 
the two taxa are not svtiou vinous. 1 agree with other 
authors that H lauta represents H. scmiplicata 
Menke, 1843. 

50. The authorship of H clegans has also been indicated 
as "Koch in Philippi" [e.g.. Wells and Bryce, 1985: 
34). The date printed on the description page of H 
elcgans is June 1844. This is the correct date. Phi- 
lippi (1842-1845) indicated Koch as a reference; 
however, it is apparent that PhiUppi actually wrote 
the description and that only a short note was taken 
directh' from Koch. Accordingly, I consider Phihppi, 
1844, the author. 

51. The taxon as described by Leach (1814: pi. 23) is 
H. ruber. The correct inflected spelling for this ad- 
jectival species epithet, however, is H. nibra. Both 
speUings are found in the fiterature. 

52. Halioti.s ancile Reeve, 1846, has been shown to be 
a juvenile H. nibro Leach, 1814 (Geiger, 1996; 
Stewart and Geiger, in press), and not a green form 
of H. pmtulata Reeve, 1846 (cf. Talmadge, 1956), 
or H. japonica Reeve, 1846 (cf. Wagner and Abbott, 
1978)" (note 16). 

53. Haliotis conicopora Peron, 1816, and H nibra 
Leach, 1814, have been showii to be closeh' related 
on the basis of allozvme frequency data (Brown, 
1993: 430): "Haliotis conicopora clustered with the 
three populations of H. nibra, with a maximum D 
of 0. 018. Similar genetic distances were found be- 
tween all conspecific populations studied (0. 003 in 
H. roei and 0. 014 m H laevigata)." Further on 
pages 430-431: "Haliotis conicopora appears to rep- 
resent an allopatric population (Western Austrahan) 
of H. rubra, as originalK' suggested bv Shepherd 
(1975) and noted bv Browai and Murray (1992a)." 
However, the two ta\a can reacUlv be distinguished 
by the shell thickness (dependent upon wave energy 
of the habitat), coloration (a potential function of 
food availabihty), and the presence of strong spiral 
cords (somewhat variable in both taxa) on a bumpy 
dorsal shell surface in H. nibra. Additionally, H. ni- 



bra is found in southeastern Australia, whereas H. 
conicorpora occurs in southwestern Australia. The 
genetic data suggest conspecifitv, but the shell and 
the disjunct geographical distribution inthcate dis- 
tinct taxa. Thus, there is some justification for sub- 
species recognition of conicopora under H. nibra. 
although this remains to be fully resolved. 

54. Halioti.s iris is credited either to Martyn (1784: cf. 
Sinchiir, 1963), or to Gmehn (1791:' r/ Pickery-, 
1991; Lmdberg, 1992). Haliotis pulchenima has 
mostly been attributed to Gmelin except bv Cf)tton 
(1943), but H. nacvosa exclusively to Martvii. AH 
three taxa are figured and named in Linnean binom- 
inal nomenclature bv Martvn. However, the work of 
Martxn has been invahdated by ICZN opinion 456 
(ICZN, 1957). Martyn s taxa are, therefore, not avail- 
able and are now crechted to the author who there- 
after mentioned the .species for the first time, which 
is Gmehn (1791) for H. iris, and Philippi (1842- 
1845:147) for H. naevosa. PhiMppi's Volume I is 
dated 1845, but the page with the description of H. 
naevosa is dated "Juli 1844;" therefore, the latter is 
the correct date for H. naevo.sa. 

55. Only a single specimen, i.e., the holotype, of H. 
whitehousei (Colman, 1959) is known. It is consid- 
ered by P. Colman (pers. comm. ), who described 
the taxon, as a teratological specimen of H. nibra 
Leach, 1814. Despite regular collecting at the type 
locahty, no second specimen has ever been found 
(P. Colman, pers. comm.). 

56. Intermechate specimens of H. scalaris (Leach, 1814) 
and H. emmae Reeve, 1846, are well known from 
western South Australia, indicating a close relation- 
ship between the two taxa (Shepherd, 1973; Wilson, 
1993). The variabihtv seems to be caused bv the 
interplay of en\ironmental factors such as wave en- 
erg)-, depth, and temperature (S. A. Shepherd, pers. 
comm). Subspecific status of the taxon emmac under 
H. scalaris is inchcated, as the morphological gradi- 
ent behveen the two taxa occurs onl\- in a small geo- 
graphical area, with the larger part of the species 
distribution showing only one of the morphologies. 

57. The hpe specimen of H. clevata Sowerby, 1882, has 
not been found. The specimen was not figured in 
the standard dorsal and \-entral position, but was tilt- 
ed. However, it is e\ident from the dorsal sculpture 
and the eccentric spire that a small specimen of H. 
.sqitamata Reeve, 1846, was illustrated. According to 
Sowerby (1882:27), it differs from H. stomatiaefor- 
mis Reeve, 1846 (syn. of H. squainata: see note 39), 
by having strong, scaly ridges, a character rather typ- 
ical for juvenile specimens of H. scpiamata (Talmad- 
ge, 1955). 

58. Haliotis funebris Reeve, 1846, was svnonvmized by 
Hedley (1914) with H. diversicolor Reeve, 1846, and 
H. tayloriana Reeve, 1846. My inspection of the 
Upe materi;il in the BMNH shows that H. funebris 
is clearlv a svnonvm of H squamata Reeve. 1846 
(see Pilsbn,-, 1890:92). 



Page 106 



THE NAUTILUS, Vol. Ill, No. 3 



59. Udliotis '^iaaniea Chemnitz, 1788, H jjabra Chem- 
nitz, 1788, and H. nigosoplicata Chemnitz, 1788, 
are names that derive from the two first words of 
the original descriptions and tliat. therefore, cannot 
be inteqireted as binominal names: the work and 
tlie authorship of these taxa are not av;ulable (ICZN, 
1954). The names were validated bv Gmehn (1791: 
H. gigantea, H. ojahra) and Reeve (1846: H. nigo- 
soplicato). 

60. Haliotis virginca Gmelin, 1791, has been divided 
into four, geographically separated subspecies: H. 
virginca virginea. H. virginca crispata Gould, 1847, 
H. virgiiw luitfoiti Filhol, 1880, and H. virginca mo- 
no liaFowell 1938 (Kaicher, 1981; Ubaldi, 1986). 
Conflicting opinions are expressed as to whether 
these are valid subspecies (Powell, 1979; Ubaldi, 
1986), or whether they would rather represent vari- 
ations caused bv differences in water temperature 
(Talmadge, 1957a). No good data are available to 
support either of the hypotheses; I retain usage of 
subspecific taxa. 

61. H. gihha Philippi, 1846, is given as a svnonvm of H 
virginca Gmehn, 1791, by Suter (1913), supported 
by the figures in Reeve (1846) and Weinkauff 
(1883). 

62. Old specimens of H. discus liannai Ino, 1952, from 
Japan are usuallv identifieil on the original label as 
the NortJi American H kamtschatkana Jonas, 1845, 
to which it is strikingly similar. Haliotis discus han- 
nai and H. k. kamtschatkana differ at the species 
level (Owen et al. 1971; Brown, 1993; Lee and Vac- 
quier, 1995). 

63. Haliotis cxigua Dunker, 1863, is usuallv encountered 
in material from the RMikvii Archipelago (Figures 
14—15). It has morphological affinities to both H. 
varia Linnaeus, 1758, and H. divcrsicolor Reeve, 
1846. Pilsbry (1895) regarded H. cxigua as a juvenile 
of H. divcrsicolor The flat shells are rather nonde- 
script, have an irregular sculpture, and are usually 
of a dark, muddy coloration. Whether these speci- 
mens deserve recognition at the species level is un- 
clear. Inspection of preserved material would cer- 
tiiinlv help to resolve the status of this species. 

64. The type specimen of H. sicholdii Reeve, 1846, is an 
aberrant specimen, as indicated e.g., by Dunker (1882: 
148): "Haec species mihi est \alde dubia, . . . credam 
eam nU representare nisi confonnationem H;iliotis gi- 
ganteae monstruosam." (This species is verv doulitful 
to me, ... I believe it does represent nodiing but a 
grotesque form of Haliotis gigantea). The types of H. 
sicholdii and H. gigantea Ginebn, 1791. refer to the 
same species (Habe, 1983). The taxon sicholdii had 
long been used for a fuitlier species eventually named 
H. niadaka (Habe, 1977). The two species can be dis- 
tinguished as follows. In H. gigantea a line drawn 
dirough the apex of the shell and tlic last perforation 
results in very une(juiil areas of tlic shell, wliereas the 
same line drawn in H. madaka divides the surface area 
of the shell approximatciv in hail. 



Haliotis gigantea Menke was mentioned in Wein- 
kauff (1883:25) as pubhshed in Menke (1843), but 
Sherbom (1922) considered 1830 to be the year of 
publication of this taxon. Menke (1830:87; 1843:31) 
referred in both cases to H. gigantea Chemnitz, 
1788, an unavailable name first mentioned thereaf- 
ter by Gmehn (1791:3691), who listed this species 
from Austraha ("Hahitat rarissima ad novam Holan- 
diam" [lives very rarely in Australia]). However, the 
type specimen of H. gigantea Gmelin, 1791, figured 
in Habe (1983) clearly shows the well-known north- 
west Pacific species. Due to the erroneous tvpe lo- 
cality, H. gigantea has been incorrectly synonyinized 
with the common, commercially exploited H. rubra 
Leach, 1814, from southern Australia (e.g., Menke, 
1843). 

65. The type specimen of H. dicgoensis Orcutt, 1900, is 
a monstrosity induced bv boring (jrganisms. most 
hkely sabellid polvchaetes (see also Oakes and 
Fields, 1993). 

66. Adult specimens with missing tremata have been re- 
ported occasionallv in the literature. These repre- 
sent deformations (see Leighton, 1960), rather than 
vahd taxa Haliotis imperforata Gmelin, 1791, mav 
belong in this category or may be a misidentified, 
non-haUotid species. In the case of H. impciforata 
Dall, 1919 (non Gmehn, 1791), H. lusus Finlay, 
1927, was proposed as a nomcn novum. Finlav 
(1927), however, had overlooked the name H. crach- 
crodii liolzneii Hemphill, 1907, which has priority. 

67. Haliotis assinulis Dall, 1878, is a subspecies of H. 
kamtschatkana Jonas, 1845, as discussed bv McLean 
(1966). 

68. Haliotis coreanica Weinkauff, 1883, is mentioned in 
Habe and Kosuge (1964) and Habe (1983) as a .syn- 
onym of H. gigantea Gmehn, 1791. This sviionvniy 
is based, however, on a misunderstanding of the 
German text of Weinkauff by these two authors. 
Weinkauff (1883:27-28) wrote'(translated from Ger- 
man): "From Mr. Paetel I received on a loan basis 
a species, which was labeled H. coreanica A. Adams, 
which, however, could not be separated from H. dis- 
cus, which would be considered a further variety'. 
However, I cannot find where this species is de- 
scribed." Although the second and the third state- 
ments contradict one another to a certain extent, my 
inteipretation of the text is that Weinkauff consid- 
ered the specimen labeled H. coreanica A. Adams 
to be the same as his H. gigantea Var. \ = H. discus 
Reeve, 1846. This particular specimen was not fig- 
ured by Weinkauff, niling out the possibility that an 
illustration could serve the purpose of a valid de- 
scription. The name is a nomcn nudum. 

69. The type specimen of H. duhia Lamarck, 1822, is 
not in MHNG or the MNHN. Most of Lamarck's 
types are otheivvise deposited in tlie MHNG {cf. 
Mennod and Binder, 1963). 

70. Haliotis cxcisa Gray, 1856, is not listed in Sowerby 



D. L. Geiger. 1998 



Paee 107 



(1882) or Weinkauff (1883). but is indicated in Ah- 
bott and Dance (1983) and in Picken' (1991) with- 
out indication of page number under Gray (1826) 
and not Gra\ (1856). Sclii.sinoti.'i cxciso was men- 
tioned in Grav (1856:148), Init as a h\pothetic;il 
name for the teratological specimens with sHt tre- 
mata he had at hand: "When I first saw the shell, I 
was inclined to regard it as a monstrositv; but when 
I considered the unif()rmit\' ... in the specimens 
. . . , I thought that it nught be the t\pe of a new- 
form, for which Schismotis cxcisa would be a good 
name. ... I am inclined to befieve that the sht in 
the specimens is . . caused bv the eroded and ev- 
idently diseased state ot the specimens." Clearlv, no 
new taxa had been described and the genus and spe- 
cies are unavailable. 

71. Haliofi.s fafiii Rheder [sic]. 1981, is mentioned in 
Ubaldi {1993:11 3-1). The date is most likely based 
on Kaicher's (1981: card no. 2902) statement: "This 
subspecies [of H. vaiio Linnaeus, 1758] is currently 
under study by Dr Harald Rehder (USNM) and will 
probabK" be described in detiiil before the end of 
the vear (1981)." The name was mentioned a second 
time in the same year (Anon. , 1981). However, the 
species description was never pubhshed. The pro- 
posed tvpe specimen for the species from Tonga has 
been deposited in the USNM. The name H fattii is 
currentK' not a\ailable. 

72. Haliotis hanleijana Ancey, 1881, has some affinities 
with H. clathrata Reeve, 1846 (see also notes 24, 
25). However, this identification is highK tentatixe. 

73. Haliotis califonuana Valenciennes, 1831, H intcr- 
nipta Valenciennes, 1831, and H. parma Valenci- 
ennes, 1831, were described from "America." The 
author stated that it was unknov\ai whether the spec- 
imens were obtained from the east or west coast. 
The specimens were 36 mm and larger, making it 
unhkelv that they could have been the small H. 
pourtalcsii Dall, 1881, H. dalli Henderson, 1915, or 
H robcrii VIcLean. 1970. Haliotis parma has mark- 
ings from the shell nmscle in the shell (Valenci- 
ennes, 1831), which additionally suggests that it rep- 
resents one of the large Cahfomia species. Iff/ cal- 
ifomiana, H. internipta. or H panna should lie s\ni- 
onyiiis of either H ful^cns Philippi, 1845, H. 
hamtschakana assimilis Dall 1878, H walallcnsis 
Steams, 1898, or H. sorcnscni Barsch, 1940, then 
Valenciennes' name would ha\e priority; the other 
Califomian species (H. crachcrodii. Leach, 1814, H 
cornifiata. Wood, 1828, H nifcsccns Swainson, 
1822) were described before 1831. Haliotis inter- 
nipta has been synonymized with H cracherodii. 
and H caJiforniana with H nifcsccns (Caqienter 
1864:521 /ir7r McLean, 1966:156, 159); the sviionv- 
mv of H. panna is unresolved and the ta\(jn is treat- 
ed here as a nomcn dubiiim. If any of Valenciennes' 
taxa should eventually be shown to be senior s)n- 
onvms, then an application to the ICZN ff)r the sup- 
pression of Valenciennes" taxa would be advisable. 



74. Karsten (1789) published two names for Haliotis, H. 
iiidis Karsten, 1789, and H. plicata Karsten, 1789. 
These two taxa are synonyms of H. iris Gmelin, 
1791, and H. australis Gmelin, 1791, respectively. 
Karsten's t;L\a would be the senior s\iion\'ms, but 
Rosenberg (1996) has formulated a ca.se to suppress 
Karsten's work that is currentK- pending with the 
ICZN. His argument, based on' ICZN Article 80, is 
accepted here. 

75. Haliotis modcsta auct. was mentioned by Menke 
(1845:194) as a juvenile specimen of H. capensis 
Dunker, 1844 (= H. midac Linnaeus, 1758). The 
original source of H. modcsta is unknown to me. 
Menke (1845), as the onh- author, also put H scmi- 
plicata Menke, 1843, into s\iion\-m\' with H. capen- 
sis. I do not accept his opinion for the following 
reasons. H. scmiplicata occurs exclusivelv in Austra- 
han waters, whereas H. midac is endemic to South 
Africa. Comparisons of the shells of adult H. scmi- 
plicata and juvenile H. midac, which are of the same 
size, reveal the following characters. Haliotis scmi- 
plicata has distinct spiral cords, whereas in H midac 
they are onlv weakly developed. Spiral undulations 
form nodes in the middle of the shell of H. scmi- 
plicata. which give rise to a sUght, but fairly distinct, 
spiral ridge, whereas H. midac tends to forms a spi- 
ral depression appro-dmateh- one third from the su- 
ture. The shape of H scmiplicata is much more 
elongated than the rotund shell of H midac from a 
size of approximately 2-3 cm f)nw-ards. 

76. Haliotis scliroctcii Menke is mentioned by Wein- 
kauff (1883:83) as a taxon of doubtful status. Wein- 
kauff indicated a potential synononymy with H. scu- 
tnhim Reeve, 1846 (= H. varia Linnaeus, 1758 ?, 
(/ note 48). Weinkauff did not know of the original 
publication, and it is not fisted in Sherborn (1922; 
1932). I regard it as a nomcn dubium. 

77. A specimen labeled as tvpe of H. victoriac Brazier 
is kiiowTi from the SAM (R. Pickers-, pers. comm. ); 
however, no formal description has been located so 
far, which renders the name unavailable (ICZN Ar- 
ticle 12c). The specimen can be identified as a H. 
nibra Leach, 1814, Any description of H. victoriac 
is \-er\ Ukelv to have been published after 1814, as 
the onlv abalone species had been described b\- Bra- 
zier in 1878. In case a published description of H 
victoriac should be found, the taxon would then onh- 
become a further juiiior .synonym of H. nibra. 

VALID SPECIES BY FAUNAL REGIONS: 

The broad geographical distribution is indicated below- 
for each taxon considered valid here. In the case of a 
minor overlap the species is fisted onlv in the major 
province. The fomiat of the entries is as follows; taxon. 
+ sviionvms. (Figures herein). Geographic cfistribution 
(illustrations) [Notes], Nomina dubia. nomina ntida and 
unaviiilable names are not included under the s\nony- 
mies. Tentative sviionvins are indicated bv- a question 



Page 108 



THE NAUTILUS. Vol. Ill, No. 3 



mark after the ta.\on, and the author of a sv-nonvm is 
onl\- indicated in case ot homonymy (see Inde.x to Spe- 
cies-Level Taxa for details). 

The distributions of the species are mainlv according 
to Macnae and Kalk (1958), Kira (1962), Habe (1964), 
McLean (1978), Muller (1984b), Ubaltli (1986), Dharma 
(1988), Herbert (1990), Wilson (1993), Geiger (1996), 
Stewart and Geiger (in press), and various collection re- 
cords in museums and private collections. Species oc- 
curring in more than one province are marked with an 
asterix and are cross-referenced; their distribution is in- 
dicated for the area vvithin the respective pro\ince. 

The hst of illustrations is not exliaustive, but focuses 
on the more recent pubhcations. The following numer- 
ical code has been used: 1, Abbott (1954); 2, Kira (1962); 
3, Habe (1964); 4, Keen (1971); 5, Hinton (1972); 6, 
Dance (1974); 7, Hinton (1978); 8, Powell (1979); 9, 
Eisenberg (1981); 10, Kiiicher (1981); 11, Bosch and 
Bosch (1982); 12, Kilburn and Rippey (1982); 13, Abbott 
and Dance (1983); 14, Sharabati (1984); 15, Stewart 
(1984); 16, Wells and Br\ce (1985); 17, Springsteen and 
Leobrera (1986); 18, Richards (1987); 19, Dharma 
(1988); 20, Drivas and Jay (1988); 21, Salvat et al. (1988); 
22, Poppe and Goto (1991); 23, Barash and Danin 
(1992); 24, Wilson (1993); 25, Giannuzzi-Savelli et al. 
(1994); 26, Geiger (1996); 27, Stewart and Geiger (in 
press); 28, Simone (in press). 

Species that have seldom been figured {H. dissona 
(Iredale, 1927), H. exigua, H. jacnensis Reeve, 1846, H. 
mannorata Linnaeus, 1758, H. ncglccta, H. planata. H 
ijuchctti Smith, 1914, H. nibigiuosa Reeve, 1846, H. spc- 
ciosa Reeve, 1846; H. unilatcralis) are here illustrated in 
Figures 10-33 along with some noteworthy specimens 
{H. diversicolor Reeve, 1846, from Bah, H. striata Lin- 
naeus, 1758). 

Caribbean 

sp. now Simone. in press. Venezuela to central Brazil (28) [note 

1]. 
pourtalesii Dall, 1881. S Florida to Surinam (10, 13) [notes 1, 
2,3]. 

European and Senegalese 

mannorata Linnaeus, 1758. -I- decussata, guineensis. rosacea, 
strigata, virginea Reeve {non Gmelin). (Figures 18-19). 
Central W Africa (10 [as H guinccnsis], 13 [as H ro.sa- 
cea]) [notes 4, 5, 6, 7]. 

neglecta Philippi. 1848 (Fipires 20-21). Isl. S of ItaK [notes 
5, 8, 9|. 

° jiiistulata cnienta Reeve. 1846. Israel and Lvbia; see also east 
African pnnince (10 [as H cnienta], 14, 23, 25) [notes 
10, 11, 12]. 

tiiherctilata tuherciilata Linnaeus, 1758. + bistriata Gmelin, 
bistriata Costa, bisundata, inrisa. jantis, japonica, lamel- 
losa. lucida. pana Risso (nott Linnaeus), pelliicida. reti- 
culata. nig(Ka Reeve (non Lamarck), .'iccemenila. striata, 
varia Risso (non Linnaeus), vtilgaris. Mediterranean, Brit- 
tany to Morocco (10 [as H lanicllosa]. 13, 22) [notes 4. 5, 
13.' 14 15, 16]. 

tuberadata coccinea Reeve, 1846. + cauiiriensis. zealandica 
Canary Lsl,, Azores (10, 22 [hotli as //, coccinea]) [notes 
5, 13. 17, 18]. 



South African 

midac Lnmaeus, 1758. -I- capensis, elatior St. Helena Bav to 
W Transkei (6. 10. 12. 13, 18) [note 4]. 

parva Linnaeus, 1758. + canaliculata Fischer ?. cantdiculata 
Lamarck, carinata, cingulata, kraussi, nibiainda Roding, 
nibiainda (Montfort). False Bav to East Lf)ndon (10, 12, 

13, 18) [notes 4, 19, 20[. 

queketti Smith, 1910 (Figures 2-4—25). Transkei to S Mozam- 
bique (10). 

spadicea Donovan. 1808. + ficifonnis. sangtiinea. simiata ? 
Partridge Point, Cape Penmsula to N Natal (9, 10, 12, 13, 
18) [notes 21, 22]. 

spcciosa Reeve, 1846. + alfredensis. (Figures 30-31). Port Al- 
fred to W Transkei (10 [also as H. speciosa form alfreden- 
sis]. 13, 18). 

Eastern African, Red Sea and Persian Gulf 

" clathrata Ree\e, 1846. + vcnusta. Ken\a, Madagascar, Mas- 
carene Isl.. Rodrigues Isl. , Aldabra, Seychelles, Chagos 
Arch., Maldi\'es; see also tropical Pacific pro\ince (5 and 
19 [both as H. crebriscidpta]. 10 and 27 [also as H. ven- 
iista]) [note 24, 25]. 

mariae mariae Wood, 1828. Oman (10, 11) [note 26]. 

mariac dentata Jonas, 1846. Oman (10 [as H. mariae form den- 
tata]) [note 26[. 

pustulata pustidata Reeve, 1846. + jousseaumi. N South .Africa 
to Persian Gulf (10) [notes 10. II, 12]. 

° pustidata cnienta Ree\e, 1846. Particularly Red Sea; see also 
European and Senegalese pro\ince (10 [as H cnienta], 

14, 23, 25) [notes W. 12]. 

.squamosa Gray, 1826. + roedingi. S Madagascar (15, 27) [notes 
27, 28, 29]. 

unilatcralis Lamarck. 1822 (Figures 32-;33). Central E Africa 
to Red Sea. Aldabra, Madagascar, Mascarene Isl. (14 [as 
Sanhaliotis pu.stidata]. 20 [as H varia], 26) [note 30], 

nigosa Lamarck. 1822. + altcrnata, mtiltiperforata, nebulata, 
pertusa. revelata. Mascarene Isl. Madaga.scar, central E 
Africa (6 and 20 [both as H pustulata], 10 [;is H. pustulata 
fonn alteniata]) [notes 11, 31, 32, 33]. 

Indian Ocean and Tropical West Pacific 

asinina Linnaeus, 1758. + asinum. S-most Japan to Sydney, 

Andaman Isl. to New Caledonia (2, 5. 6, 7, 10 [jm'enile 

and adult], 13 16, 17, 19, 24) [note 4]. 
brazieri Angas, 1869. + melculus. (Figures 4-.5). S Queensland 

to Jenis Bay, New South Wales "^(7, 10, 13, 24) [notes 34, 

35[. 
" clathrata Reeve, 1846. + venusta. Andamans to American 

Samoa, S Japan to Sydney; see also east African province 

(5 and 19 [bodi as H. crebriscidpta], 10 and 27 [also as 

H venusta]) [note 24, 25]. 
crebriscidpta SowerbN', 1914. New Caledonia (10, 27) [notes 

25, 36]. 
° diversicolor Reeve, 1846. + aquatilis. gnmcri. supertexta. 

tnijloriana. (Figures 12-13). China, Taiwan, Bali; see also 

northwestern Pacific pro\ince (2 [as H. supertexta], 3, 10 

[also as fonns gnineri. taijloriana. supratexta, and H. 

aquatilis]) [notes 37, 38]. 
di.s.sona (Iredale, 1929). (Figures lO-lD. Queensland. New 

Caledonia, Tonga (10) [note 36. .39[. 
dohmiana Dunker 1863. New Caledonia, Tonga (10, 21. 27) 

[note 40], 
glabra Gmelin, 1791. + picta, ziczac. Philippines, MaUiku, 

Lesser Sunda Isl, (9, 10, 13, 17, 19 [as H, planata]). 
hargravesi Cox, 1869. -I- ethologus. S Queensland to N New 



D. L. Geiger, 199S 



Page 109 




Figures 28-33. Intn-ijiu'iitK illustrated Iliiliotis 2.S-29. Udliotis stnatti Liiiiuieiis, 175S. 56 ruin Out- of tlie six spt-cinifus in 
LSL, Mare eiiropaeus. 30-31. H. speciosa Reeve, 1846. 31 mm. NMW 1955 158, 2126. Algoa Ba\. 32-33. H. uniltiteralis Lamarck, 
1822. 32. 23mm. Collection S. Singer. Red Sea, Gulf of Aquaba, 33. 24 mm. Collection Geiger AAB 48a. Elat, Red Sea. 



South Wales (7, 13. 10 and 24 [both also as H ethologus]) 
[notes 34, 35]. 

jacnensis Reeve, 1846. + echinata, hanleiji. (Figures 16-17). 
S-most Japan (S of Amami Isl.), Phihppines, New Cale- 
donia, Marianas, Micronesia (3, 10 [also as H haiilri/i]) 
[notes 31, 41]. 

ovinn Gmelin, 1791. + cnclatn. latilahris. Maklixes to Tiia- 
motus, SW Japan. Philippines, \ietnam, Queensland to 



Western Australia (2, 5, 7, 9, 13, 16. 17. 19, 21, 24) [note 
S]. 

° planata Sowerbv, 1882. + graijana. (Figures 22-23). Phihp- 
pines to Fiji, Northern Territories; see also northwestern 
Pacific province (10 [also as H oraijana]) [notes 31, 42]. 

ptilchcrritua Gmelin. 1791: Tiiamotus, Henderson Isl. (9, 10, 
13). 

ndbi^inosa Reeve, 1846. + Itowensis. (Figures 26-27). Lord 



Page 110 



THE NAUTILUS. Vol. ill. No. 3 



Howe Isl. (10 [a.s H. variii nihiginosa and H. lioicensis], 
7 and 13 [both a,s H. howcnxis]) [note 43]. 
varia Linnaeus. 1758. + alwna. iistricta, barbouri, concinna. 
clringii, gemnm. grantilata. paptilata. pustulifera. saitu- 
Itim, semistriata, viridis. Sri Lanka to Tonga, S Japan, Phil- 
ippines, central W Western Australia to Sydney (2, 5, 7, 
9. 10 [also as ssp. astricta, papulata, pustulifera, viridis. 
and H. unilntcrnlis]. 13. 16. 19) [notes 1, 4, 44, 45, 46, 
47, 48], 

Temperate Australian 

coccoradiata Reeve. 1846. New South Wales to E Victoria (7, 

9. 10. 24) [note 49]. 
cijclobates Peron. 1816. + excavata. South Australia to central 

S Western Austraha (9, 10 [also as H ovina], 13, 16, 24), 
elegans Philippi, 1844. + cinthrata Lichtenstein {non Reeve). 

Western Australia (7, 9, 10, 13, 16, 24) [notes 23, 50]. 
laevigata Donovan, 1808. + albicans, glabra Swainson? {non 

Gmelin). Victoria to SW Western Australia (6, 7, 10, 13 

16, 24). 
roei Grav, 1826. + scabricostata, sidcosa. Victoria to central W 

Western Australia (6, 7, 10 [also as H sulcosa], 13, 16, 24) 

[note 27], 
rubra rubra Leach, 1814. + ancile. improbula. naevosa. iihi- 

tehousei. New South Wales to South Australia, Tasmania 

(7 and 13 [both as H. ntber], 10 [also as ruber, r. impro- 

btdum, ancile]) [notes 25, 27, 51, 52, 53, 54, 55], 
rubra conicopora Peron. 1816. + cunninghami. granti. vixlir- 

ata. Victoria to Freeniantle SW Western Australia. (7, 16. 

24 [all as H conicopora], 10 [also as H. c. vixlirata]) [note 

53]. 
scalaris scalaris (Leach, 1814). + crcnata ?, tricostatis, tricos- 

tata. W South Australia to central W Western Australia (7, 

9, 10, 13, 16, 24) [notes 24. 56]. 

.scalaris emmae Reeve, 1846. Victoria to W South Australia (7, 

10, 13, 24 [all as emmae]) [note 56]. 

semiplicata Menke, 1843. + lanta. SW Western Australia (7, 
13, 24) [note 49]. 

squamata Reeve, 1846. + elcvata. funebris. stomatiaefonnis. 
Central W Western Australia to Northern Territories, Bah 
(7, 9, 10. 16, 19, 24) [notes 31, .39, 57, 58], 

New Zealand 

australis Gniehn, 1791. + aleata. costata, nigosoplicata. New 

Zealand (8. 10, 13) [note 59]. 
iris Gmehn, 1791. New Zealand (6, 8, 10, 13) [note 54]. 
virginea virginea GmeHn, 1791. + gibba, marmorata Reeve 

(non Linnaeus), subvirginea. S South Isl. and Stewart Isl., 

N South Isl. to S North Lsl. (6. 8. 10) [notes 60, 61], 
virginea crispata Gould, 1847. NE North Isl. (8, 10) [note 60]. 
virginea huttoni Filhol. 1880. Auckland Isl.. Chapman Isl. (8, 

10) [note 60], 
virginea morioria Powell, 19.38. Chatham lsl. (8, 10) [note 60]. 

Northwestern Pacific 

° divcrsicolor Reeve, 1846. + aquatilis, gnineri, .^uperiexta. 

taijloriana. (Figures 12-13). S Hokkaido; see also tropical 

Pacific province (2 [as H. .•iupertexta]. 3, 10 [also as forms 

gmneri, taijloriana. sitpratexta. and H aquatilis]) [notes 

37, .38]. 
discus discus Ree\e. 1846. Honshu. Ksuslni. Shikoku (10) 

[note 62]. 
disctis hannai Ino, 1953. Korea. Hokkaido. NE Honshu (2. 10, 

13) [note 62]. 
exigua Diniker 1877 (P^ignres 14-15). S Japan (Okinawa) (10) 

[note 63]. 



gigantea Gmelin, 1791 + gigas. sieboldii. tubifern. Honshu (2, 

10) [note 64]. 
madaka (Habe, 1977). S and central Honshu. Kviishu (2. 10 

[both as sieboldii]) [note 64]. 
" planata Sowerby, 1882. + grayana. (Figures 22-23). S of 

Yakiishima; see also tropical Pacific province (10 [also as 

H. grayana]) [notes .31, 42]. 

Northeastern Pacific 

corrugata Wood, 1828. + diegoensis, nodosa, oweni. Central 
to central Baja Cafifomia (1, 6, 9, 10 [also as ssp. oweni], 
13) [notes 26, 65, 66]. 

cracherodii Leach, 1814. + bonita. californiensis. expansa, 
holzneri, imperforata Dall {non Gmelin). lustis, rosea, 
splendidida. N Calitoniia to central Baja California (1, 6, 
9, 10 [as H. cracherodii californiensis and H. c. cracher- 
odii], 13) [note 66]. 

fulgens Philippi, 1845. + guadalupensis, planilirata,splendens, 
turveri. Central California to central Baja Cahfomia (1, 6, 
10 [also as ssp. guadalupensis], 13) [note 47]. 

kamtschatkann kamtschatkana Jonas, 1845. Alaska to Point 
Conception, S California (1, 10, 13) [notes 62, 67]. 

kamtschatkana assimilis Dall, 1878. + aulaea. smithsoni. Cen- 
tral to S California (1, 9, 10, 13 [all as H assimilis]) [notes 
62, 67]. 

rufescens Swainson, 1822. -I- calijomiana. hattorii. ponderosa. 
N California to Central Baja California (1, 6, 13). 

sorenseni Bartsch, 1940, Point Conception to central Baja Cal- 
ifornia (10). 

walallensis Steams, 1898. S Washington to S Calitoniia (10, 
13). 

Tropical Eastern Pacific 

dalli Henderson. 1915, Galapagos Isl, Gorgona Isl. (4, 10, 13) 

[note 2]. 
roberti McLean, 1970. Cocos Isl. (4, 10) [note 2]. 

ZOOGEOGRAPHY: 

The .specie.s in South Africa, New Zealand and the north- 
eastern Pacific are all endemic, and with the exception 
of one questionable report (Macpherson, 19.5.3: the 
South African H. spadicca Donovan, 1808, in Australia) 
no witle spread species have been foimd in these areas. 
The East African and the Indo-Pacific provinces are also 
more or less isolated from each other; Geiger (1996) 
discussed the apparent faunal barrier around India. Only 
H rlathrata Reeve, 1846, occurs in both areas (Stewart 
and Geiger, in press). Indications particularly t)i H. varia 
in the East African province, and H. pustulata in the 
central Indo-Pacific are erroneous (Geiger, 1996; pers. 
obs). In the Australian province the separation of trop- 
ical and temperate species is not verv' shaq); consider- 
able overlap has to be noted tor a few species. Only a 
limited munber of species are widespread over several 
faunal provinces, i.e., H. asinina. H. clathrata Reeve, H. 
(uina and H varia. A much more detailed, specimen 
based account of the distribution of all abalone species 
will be provided elsewhere. 

Three biogeographical models have been introduced 
and are reviewed in (Tciger and Groves (submitted). Tal- 
madge (1963a) proposed the Pacific Rim hvpothcsis with 
nuiltiple centers of radiations ;dong the Pacific islands 



D. L. Geiger, 1998 



Page 111 



from Japan to Australia and New Zealand. Lindberg 
(1992) pointed to the highest Lli\ersit\- oi alxilones benig 
found in the central Indo-Pacific, which may (or niav 
not) indicate the origin ot the group there. Based on 
published chromosomal data Geiger and Groves (sub- 
mitted) suggested a potential origin of abalone in the 
Tethvs Sea, which is ;ilso discussed b\' Lee and Vacquier 
(1995). 

ACKNOWLEDGMENTS 

This review^ would not have been possible without the 
critical remarks bv several colleagues, who also made 
their knowledge and data generously available to me: 
Phil Colman (Sydnev, Australia), Youn-Ho Lee (Pasa- 
dena, California), Mark Jones (Auckland, New Zealand), 
Roger Pickerv' (Wilrijk, Belgium), Benjamin Singer (Re- 
hovot, Israel), Scoresby A. Shepherd (AdeUiide, Austra- 
ha), Katharine Stewart (Carmel, California), Victor Vac- 
quier (San Diego, California), Rick Fallu (Canberra, 
Australia), and Buzz Owen (Gukila, CaBfomia). James 
H. McLean (Los Angeles, California), Veronica Miller 
(Los Angeles, C;difomia), Scoresby A. Shepherd, Jose H. 
Leal (Sanibel, Florida), and two anonyinous rexiewers 
read the manuscript, including earlier drafts, and made 
valuable comments. 

I would like to thank all the staff at museums, who 
helped to make stays at their institutions successiul, 
helped with library research, and made specimens avail- 
able for study: Henk Mienis (HUJ), Kathie Way and Ju- 
ha Freeman (BMNH), Ahson Trew (NMW), Yves Finet 
and Claude Vaucher (MHNG), Roger Pickew (KBIN), 
Raye Germon, Jerry Harasewych, and Alan Kabat 
(USNM), James McLean and Lindsey Groves (LACM). 
Paul Scott and Henry Chanev (SBMNH), Terrence Gos- 
liner (CASIZ), David Undberg (UCMP), Paula Mikkel- 
sen (DMNH and AMNH), Gan- Rosenberg and Ned 
Gilmore (ANSP), Peter Jung (NHB), Philippe Bouchet, 
Pierre Lozouet and Philippe Maestrati (MNHN), Mar- 
garet Gosteh (NMBE), K. Boss (MCZ) Dominik 
Brantschen and Verena Geiger (Bern, Switzerland) kintl- 
ly made adchtional material aviiilable. The staff at Han- 
cock Library at USC helped to locate many obscure ref- 
erences. The visits of DMNH and ANSP were made 
possible through a DuPont Merck Scholarschip in Mal- 
acologv', and a Jessup Fellowship, respectively. Research 
resulting in some of the new anatomical data reported 
here was supported liv grants from the Hawaiian Mal- 
acological Society', the Western Society for Malacology, 
and the Lerner Gray Fund for Marine Research 
(AMNH). I express my gratitude to all these persons and 
institutions. 

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THE NAUTILUS 111(3);1 17-118, 1998 



Paee 11" 



On the Identity of Bucciniim Jahijhnthus Gmelin, 1791 
(Gastropoda: Buccinidae) from the Netherlands Coast 



Guido Pastorino 

Depto. Paleozoologi'a Invertebrados 

Museo de La Plata 

Paseo del Bos(jue s/n 

1900 La Plata, Bueno.s Aires 

ARGENTINA 

npa.storCs'c'riba.edu.ar 



Yuri Kantor 

A.N. Se\ertzo\- Institute of Problems of 

Evolution 
Russian Academy of Sciences 
Leiiinskij Prospect, 33 
Moscow 117071 RUSSIA 
vxiri@invert.se\in. msk.ru 



Biicciiuim laht/riiithus was described and illustrated bv 
Martini (1777! Vol. 3, Pi. 120, fig. 1103) who based the 
new taxon on a single specimen from Scheweningen, 
Holland. Martini's work, however, was rejected for no- 
menclatorial purj^oses bv the International Commission 
on Zoological Nomenclature (1954). Gmelin (1791), hav- 
ing cited both the name and Martinis illustration, is con- 
sidered to be the author of B. hhijrinthus. The specimen 
illustrated bv Martini, deposited at the Zoologisk Mu- 
seum, Kobenhaviis Universitet, Denmark (ZMUC), is 
the holot\pe oi B. lahiirinthus. 

In his monograph of the family Nassidae, Trs'on 
(1882) listed Bucciiiuin lahijrinthus as a probable senior 
synonvm for Biiccinanopf: coclilidiuiu Chemnitz (= Biic- 
cinanops cochlidiuin (Dillw)n, 1817)) from the Patago- 
nian coast, calling into question the Dutch type localitv. 
In a revision of the Biillia group, Allmon (1990) followed 
Trvon in reporting B labi/riuthus as a possible older 
name tor Biiccinanops cochlidiinn. 

In the course of preparing revisions of several famihes 
of southern South American gastropods, the senior au- 
thor has had the opportunity' to examine the tspe ma- 
terial of many Patagonian species housed in European 
museums, and to compare these types with fresh-col- 
lected material. Examination of the hpe of Biiccinuiu 
labi/iinthus revealed that it was neither conspecific nor 
congeneric with Biiccinanops cochlidinm. but more like- 
ly represented a grosslv deformed specimen of a com- 
mon local species. 

The John Gwvn Jeffrevs Collection, housed in the Na- 
tional Museum of Natural Historv, Smithsonian Institu- 
tion, since the 1880s (Waren, 1980), contains a large 
number of teratological specimens of common Europe- 
an species, among them several severelv deformed Btic- 
ciniun iindatnni from England. Comparisons of the ho- 
lot)pe of Bnccinum lahi/iinthus Gmehn with some of 
the specimens from Jeffrey's Collection revealed striking 
similarities in shell shape (Figs. 1-8). Such deformities 
are likely a result of trauma to the mantle edge and the 
outer hp of the shell by a predator. The adapical regions 
of the spires resemble those of undamaged specimens 
oi Bnccinum undainm, while the onset of abnormal mor- 



phology is demarcated bv an obvious, repaired break in 
the shell. 

The holotype of Bnccinum labyrinthus has such a 
shell scar on the upper portion of its spire (Fig. 3, ar- 
row), after which the distortion of the whorls begun and 
the characteristic shoulder cord was formed. 

The originallv reported hpe locality of Bnccinum la- 
hiirinthus falls within the geographic range of Bnccinum 
nndatum. which extends from Labrador to the eastern 
Barents Sea (Golikov, 1980) and is close to the area 
where the deformed Jeffrevs' specimens were collected. 

We therefore place Bnccinum labi/rinthus Gmelin, 
1791 in the svnomnnv of Buccinum undatum Linnaeus, 
1758. Buccinum lalnpinthus is not a senior svnonvin of 
Buccinanops cochlidinm (Dillwvn, 1817), as suggested 
by Tiyon (1882) and Allmon (1990). 

ACKNOWLEDGMENTS 

Special thanks are due to Tom Schiotte, Zoologisk Mu- 
seum, Kpbenhavnis Universitet, who kindly provided 
photographs of the hpe of Bnccinum labi/hnthus. and 
to M. G. Harasewvch, National Museum of Natural His- 
torv, Smithsonian Institution, for his hospitality during 
our work at the USNM. 

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International Commission on Zoological Nomenclature. 1954. 
Direction 1 — Opinion 184. Opinions and Declarations 



Page 118 



THE NAUTILUS, Vol. Ill, No. 3 




Figures 1-8. Burriiiuin undatiiin Llniuu'ii.s, 175.S. 1-3. HoloUpe ot Biicciiuim Idlii/niilliiis Gnielm, 1791, ZMUC uniiunibered, 
Sclieweniiigen, Holland, length .56 mm. 4-5. USNM 191932, Jeffrey.s collection, W'hi.stable.s Flats, England, length 59 nnn. 6-7. 
USNM 192081, Jeffreys Collection, no locality data, length 64,2 mm.' 8. USNM 1920S2, Jeffreys Collection, no localit\- data, length 
56.3 mm. Arrows indicate repaireil hreak anil onset of deforinitv. 



rendered hv the International Commission on Zoological 
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Jeffreys, with the location ot the t\pe material. Concho- 
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lication No. 1, 60 pp., 8 plates. 



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TH Et^NAUTI LUS 



Volwne 111, Number 4 

May 26, 1998 

ISSN 0028-1344 

CONTENTS 

L. R. Saul Eight aporrhaid gastropod species from tlie Cretaceous of 

the Pacific Slope of Nortli America and clarification of the 
tMpe species of Pciissoptcra 119 

Shawn J. Wilcox Salinity tolerance of the freshwater bi\al\ e Drcisscna 

Thomas H. Dietz poh/moqiha (Pallas, 1771) (Bivalvia: Dreissenidae) 143 

Notices 149 




THE NAUTILUS 111(4):119-142, 1998 



Page 119 



Eight Aporrhaid Gastropod Species from the Cretaceous of the 
Pacific Slope of North America and Clarification of the Type 
Species of Perissoptera 



L. R. Saul 

Natural Histon Museum of Los 

Angeles Couiih' 
900 Exposition Bo\ile\ard 
Los Angeles, CA 90007 USA 
lulasaul@aol.c()ni 



ABSTRACT 

Eight aporrhaiil species iroin Pacific Slope deposits ot late Ear- 
K' Cretaceous through earlv Late Cretaceous age are assigned 
among three genera, Aporrltais. Lntialn, and Alaiiiiu'lla n. gen. 
Except for two species of Lntialn, Lntinln sp. ot Albian age 
from Oregon and Lntinln hcliaca n. sp. of Cenomanian age 
from Fresno and Shasta counties, Calitoniia, the species are of 
Turonian age. Pacific Slope Lntiala was most species-diverse 
during the Turonian and has not been found in >oimger Pacific 
Slope deposits. The three Turonian Latinia are Lntinln cnlifor- 
nica (Gabh) from Sisldvou and Sliasta comities, Cahtonna, Ln- 
tinln sigiiM n. sp. from the Redding area, Shasta (>ount^■, and 
"Ainrin" nodosn Packard, 1922, from the the Redding area, 
Shasta Countv, and the Santa Ana Moimtains, Orange Count\', 
California. Aporrhnis drnchuki n. sp. is from the Santa Ana 
Mountains, as are two species included in Alnrimrlla n. gen., 
Alarimella nnne n. sp. and the reassigned species, Alnrimelln 
veto (Packard, 1922). Alnrimelln resembles Rimclln of the 
Strombidae, but possesses an ex]")aniled (juter lip The total 
number of Tironian aporrliaids knowm trom the Santa Ana 
Mountains, southern Cahtomia is increased to six species and 
includes the first record of Aporrlinis on the Pacific Coast ot 
North Amenca. Selection of a t>pe species tor Perissoptern 
from among Cretaceous species listed bv Tate, 1865 ensures 
that Arrhoaes is not supplanted h\ Perissoptern 

Kei/ uorcl.s: Aporrhaidae, Aporrhnis. Perissoptern. Latinia, 
Alariinella. Cretaceous, Turonian, Calitoniia, 



INTRODUCTION 

This paper atlds two new species to the four species of 
Aporrhaidae pre\ioiislv recorded from beds of Turonian 
age in the Santa Ana Mountiiins, Orange Countv', Cah- 
fomia. Anchitra iHclicaulax?) tiicosa Saul and Popenoe, 
1993; Pijktcs diaphron Popenoe, 1983; Alaria nodosa 
Packard, 1922 which was considered to be a synonym of 
Arrhogcs (Latiala) califomicits (Gabb, 1864) bv Popen- 
oe, 1983; and Aporrhais vctiis Packard, 1922, which is, 
herein, assigned to the new genus Alarimella. are the 
previously described species. The new species are Apor- 



rhais drachuhi and Alarimella aitae. Recognition ot tliese 
species in the Tmonian of the Santa Ana Mountains in- 
creases the known diversity of aporrhaiids in those fau- 
nas and enkirges the paleogeographic distribution of 
Aporrhais to include the West Coast of North America 
during the Turonian. 

Re-e.xamination ot "Alaria" nodosa and Latiala ealijor- 
niea inchcates that the larger, tliicker shelled "Alaria" no- 
dosa has fewer axial rilis on the spire and should not be 
included in L. ealifontiea. Latiala ealifornica is not rec- 
ognized from the Santa Ana Mountains but is present in 
the Sisldvou Moimtains of Oregon and California and in 
the Redcbng area, Shasta County, CiJilomia. Latiala no- 
dosa (Packard, 1922) is a common to abimdant fossil in 
the Turonian ( )f the Santa Ana Mountains and also is pres- 
ent in beds of similar age in the Redding area, Shasta 
Counh', CtJifomia. The third Latiala species, L sigma n. 
sp. is describetl from the Turoiuan of the Reckling area, 
Shasta County. A fourth species, L. hcliaca n. sp., is based 
upon specimens ot Cenomanian age from Sunflower Val- 
ley, Fresno Countv', and specimens formerlv identified as 
Arrltojies ealifoniiciis from the Ono area, Shasta Countv; 
C;ilifbmia. A fifth species of Latiala is recognized from 
specimens of Albian age from Grave Creek, Jackson 
Coimh', Oregon. Naming of this new species awaits better 
preseiA'etl materiiJ. Talile 1 hsts the cbscussed species by 
age and indicates their geographic tlistribution. Areas 
from which these eight species of aporrhaiids have been 
collected are indicated on Figine 1. 

Several recent papers lia\'e added to the number of 
aporrhaids described from the Cretaceous of the Pacific 
Slope. Popenoe (1983) defined the genus Ptjktes and tlis- 
ciissed species raTiging in age from Turonian to Maas- 
trichtian. Saul and Popenoe (1993) discussed tvvo species 
o{" Anchura (Hclicatilax) of Turonian age. Elder and Saul 
(1996) discussed 10 species o( Anchura with ranges of 
1.5 to 4 m.v. within the Late Cretaceous (Coniacian 
through Maastrichtian interx'al). 

Popenoe (1983) proposed tvvo subfamihes of Apor- 



Page 120 










THE NAUTILUS, 


Vol. 


111. No. 4 


Tabic 1. .\c;e 


and 


s;eo<;r, 


uphic distriliiitldii iif discussed apoirhaid species. 




















California 








Age 


Species S 


C 




N 




Oregon 


Turoiiian 

Cenomanian 

Albian 






Alariinrlla aiuw n. sp. X 

Alarimella veta (Packard, 1922) X 

Aporrhais drarhuki n. sp. X 

Latiala californica (Gabb, 1864) 

Latiala nodosa (Packard, 1922) X 

Latiala siania n. sp. 

Latiala heliaca n. sp. 

Latiala heliaca n. sp. 

Latiala sp. 


? 
X 




X 
X 
X 
X 
X 




X 
X 



S = southern; C = central; N = northern. Juveniles at various horizons in the Great Valley Sequence on west side of the Great 
Valley, central California, resemble those of L californica. L nodosa, and L. heliaca. 



rhaidae, Aporrhainae and Arrhoginae. Among character- 
istics of Aporrhainae mentioned by Popenoe are a prom- 
inent bent digitation on the vving at the shoulder, the 
cUgitation bent mediallv with its chstal end posteriorly 
directed, additional labral cUgitations developed in some 
genera, and labral sinuses well developed. Characteris- 
tics of Arrhoginae include predominentlv axiallv directed 
sculpture, wing with an entire margin except for a prom- 
inent posteriorly directed digitation arising from the pos- 
teriolateral border of the outer lip, and, commonly, a 
straighter rostrum. Aporrhais drachnki exhibits charac- 
teristics of Aporrhaiinae. Latiala sp., Latiala Iwliaca, L. 
californica, L. sigma, and L. nodosa e.xliibit characteris- 
tics of Arrhoginae. Zinsmeister and Griffin (1995 added 
the subfamily Struthiopterinae for aporrhaids from the 
southern rim (VVeddellian Province) of the Pacific 
Ocean. Roy (1994) suggested that Sfnitliioptcra might 
be an Arrhogcs Gabb, 1868, but Stnithioptcra and other 
Struthiopterinae have two spiral carinae and Arrhoges 
lacks carinae. Rov (1994) used 25 morphologic charac- 
teristics and divided the Aporrhaiidae into two groups of 
genera. Ml with multidigitate apertural margins and M2 
with simpler apertural margins. He includes Aporrhais 
in Ml, and Arrhoges, of which Latiala Sohl, 1960, was 
proposed as a subgenus, is in M2. 

Alarinwlla does not fit readily into the above subfam- 
ilies. Its elongate shape, short anterior rostnmi, posterior 
canal extending up the spire, and its sculpture resemble 
those of Riinclla and other Ri nwl I a -\i\^i.^ gastropods 
(C]lark and Palmer, 1923). Alarimclla also resembles Ca- 
lyptraphonis binodifcnis PerriUiat and Vega, 1997, in its 
sculpture, but it lacks the callus coating and curled over 
posterior canal of C. binodifcnis and has an expanded 
outer lip shaped somewhat like that n[ Drcpaiiochcihis. 
The Rimclla-\ike gastropods and Cahiptraphonis are 
usually classed in the Strombidae from which Alarimclla 
differs in having an alatelv expanded, internally chan- 
neled outer lip. Alarimclla also lacks a strombid notch 
which perhaps suggests that its eves were on liiilbs at 
the base of the tentacles as in apporhaids rather than 
stalked as in strombids. 



Temperature implications of the three genera, Apor- 
rhais. Latiala. and Alarimclla, are somewhat equivocal. 
Undoubtedly the cUmate of the Santa Ana Mountiiins 
during the Turonian was warm; rare nidist fragments are 
found associated with some of these aporrhaids. Modem 
Aporrhais spp., however, occur from the Baltic to North 
Africa, and modern ArrhoQcs occidcntalis. arguably the 
closest hving relative oi Latiala, inhabits cool temperate 
waters from Newfoundland to North Carolina. As Cre- 
taceous aporrhaids were much more widely distributed 
than are the modem ones, modern Aporrhais and Ar- 
rhoges may not appropriately model the climate for Cre- 
taceous or early Tertiary species. Latiala is present in 
Japan in the Early Cretaceous, Pondoland, India, and 
the Gulf Coast during the latter part of the Cretaceous, 
and the Pacific Slope during the mid-Cretaceous, all in 
associations suggestive ot warm water R;;)i<7/rt-like gas- 
tropods are tropical to subtropical, and if Alarimella is 
related to them, it may indicate warm waters. The ab- 
sence of Laticda in Pacific Slope post-Turonian deposits 
coincides with the absence of several other thermophific 
molluscs (e.g., Trigonarca, Anthomja, and actaeonelhd 
gastropods) that had been present in the mid-Creta- 
ceous. 

Table 1 contrasts some characteristics of the species 
discussed in this paper. The wing dimensions are de- 
scribed as tall to short in the axial direction ( = height) 
and broad to narrow in the spiral direction ( = 
breadth). 

Institutional abbreviations used in this paper are: 
ANSP, Academy of Natural Sciences of Philadelphia; 
CASG, California Academy of Sciences, Geology: CIT, 
California Institute of Technology; LACMIP Natural 
HistoA Museum of Los Angeles County, Invertebrate 
Paleontology; UCBMP, University of California, Berke- 
ley, Museum of Paleontology; UCLA, University of Cal- 
ifornia, Los Angeles; USGS, United States Geological 
Survey; USNM, Ihiited States National Mii.seum, Col- 
lections of (California Institute of Technology and 
University of California, Los Angeles are at the Natural 
History Museum of Los Angeles Countv. 



L. R. Saul. 1998 



Page 121 




Figure 1. Iiulex map to areas from which eight aporrhaiid 
species ha\e been collected. Grave Creek, |ackst)n Countv', 
Oregon: PHonibrook Formation, late earh-niiddle Albian. Sis- 
kiyou Mountains, Jackson Count)', Oregon and Siskivou 
Coinit\', California: Osburger Gulch Sandstone Member of the 
Hornbrook Formation. Turonian. Redding area, Shasta 
Countw Califoniia: Frazier Silt and Melton Sandstone Mem- 
ber, Redding Formation, Turonian. Ono area, Shasta Countv, 
California: Bald Hills Member, Budtlen Canyon Fonnation, 
Cenomanian. Elder Greek, Tehama Count\', California: fos- 
siliferous boulder of Cenomanian or late Albian age in con- 
glomerate of Turonian age. Sunflower Valley, Kings Count^■, 
California: Panoche Formation. Cenomanian. Santa Ana 
Mountains, Orange Countv, Califoniia: Baker Canyon Sand- 
stone and lower Holz Shale Members of Ladd Formation, Tu- 



SYSTEMATIC PALEONTOLOGY 

Family Aporrhaidae Gray, 1850 
Genus Aporrliais da Gosta, 1778 

T>pe .species: Aporrhais pcspclicani (Linnaeus, 1758), 
Recent, European seas. 

Discussion: Aporrhais is a medium sized aporrhaid 



with a palmately e.xpanded v\'ing that lias at least three 
digitations. The digitations are more or less spikev, short 
to long, and channeled interiorly. The posterior digita- 
tion may be very short and entirely adnate to the spire 
or longer and iree for most of its length. The labia! dig- 
itations of which there are at least two, vary both in 
number and length even within a species. Each digita- 
tion prolongs a cord from the last whorl. The protoconch 
is multispiral with globose, smooth whorls. The earhest 
sculpture is of fine spiral cordlets which are soon crossed 
In fine curs'ed axial riblets. The riblets become stronger 
and commonlv fewer and each develops a node medially. 
The rov\' of nodes iorms the whorl shoulder and com- 
monly coincides with a strong cord as in Aporrhais iit- 
tingcriana (Risso, 1826). The whorl is subangulate, at 
least at the shoulder and basallv, the more anterior an- 
gulation is also noded and/or cordate. The whorl profile 
between the angulations is commonK' concave. At least 
one additional noticeable cord is commonly present an- 
terior to the anterior angulation. It usually produces a 
digation in some species, rarely in others. The rostrum 
is commonly bent, adaperaturally in some species, aba- 
perturalK' in others and extended by a rostral digitation. 

Aporrhais drachuki n. sp. 
Figures 2-7; Table 3. 

Diagnosis: An Aporrhais with two, nearly equal, 
strong, riiised and noded priman" cords on the body 
whorl, and a third somewhat weaker one on the base. 
Posterior-most cord iorming carina on spire whorls. 

Description: .Shell a\ erage sized for Aporrliais, mod- 
erately high spired, pleural angle about 44°; about six 
post-nuclear whorls, ungulate near mid whorl on spire, 
biangulate on body whorl; suture impressed; protoconch 
unknown; varices randomly present on early whorls; 
growth line antispirally concave on spire. Juvenile sculp- 
ture of fine collabral ribs; adolescent sculpture of about 
20 rounded arcuate ribs, strongly noded at the mid 
whorl angulation; ribs greatly reduced on body whorl but 
producing strong nodes on posterior and anterior an- 
gulation; three nearly equidistant spiral cords on body 
whorl, strongest on posterior angulation and weakest an- 
terior to anterior angulation; surface of body whorl cov- 
ered by fine raised spirals, about three per mm, becom- 
ing stronger and fanning out onto wing; posterior and 
middle cords (priman' cords) forming two angulations 
and digitations on wing. Outer lip e.xpanded palmatelv 
to form ■wing. Inner lip with finger of callus extending 
adapicaUy beyond the first suture, callus broad and rath- 
er thin parietalK', thicker and with more defined edge at 
basal sulcus. Rostiimi moderately short and narrow. 

Type specimens: Holotype LACMIP 11.37.3; para- 
n-pes LAGMIP 11375 ancl 11376 from GIT loc. 1058 
( = LAGMIP loc. 10890), LAGMIP 11374 from GIT loc. 
1065 ( = LAGMIP loc. 10891), LAGMIP 11377 from 
GIT 1064 ( = LAGMIP loc. 10893). 

Type locality: LAGMIP loc. 15295, south side of Sil- 



Page 122 


















THE NAUTILUS, 


Vol 


. Ill, 


No. 4 


Table 2. Some characteristics 


ot discussed aporrhaid 


species. 


Measurements and 


ratios 


are ax'erages from 


Tables .3- 


10. 








I'nstc- 

rior 


Wnit; 






























canal 


sliape 


\\'HA\'B 




A 




Dp/Hp 


R 




Rj 


C 


^f 


Mtrhais dracliuki ii sii 


'J 


palmate 


1.52 


+ 


0.26 


41° 


-(- 


4.8 


1.7 


± 0.2 


10 




24 


3 


Latiala n. sp. 


no 


quadrate 


0.94 


-t- 


0.1 


30° 


-(- 


1.5 


1.6 


± 0.2 


8 




20 





L. 


hcliaca x\. sp. 


no 


quadrate 


0.90 


4- 


0.05 


36° 


-t- 


3.2 


2.0 


± 0.3 


.3- 




28 





L. 


caUfomica (Gabb) 


no 


quadrate 


0.76 


-t- 


0.06 


35° 


-h 


4.0 


1.9 


± 0.2 


4- 




40 





L. 


si°ma n. sp. 


no 


fjuadrate 


0.S3 


+ 


0.06 


35° 


-(- 


3.2 


18 


± 0.2 


( 




52 





L. 


nodosa (Packard) 


no 


i(nadrate 


1.12 


-h 


0.20 


36° 


+ 


4.2 


1.0 


± 0.1 


" 




20 





Alariiui'lla anae n. sp. 


yes 


pointed 


0.44 


31° 


-(- 


2.3 


1,8 


± 0.1 


O'r' 




14 





A. 


vet a (Packard) 


yes 


triangular 


0.73 


+ 


0.15 


26° 


-(- 


2.8 


1.8 


± 0.2 


14 




y 





A = pleural angle: C = number of cords on body whorl; Dp/Hp = ratio ot whorl diameter to whorl height; R = number of iLxial 
ribs on bod\- whorl; - = ribs present only on back of whorl; Rj = number ot iL\ial ribs on juvenile whorl; WHAVB = ratio ot vying 
height to wing breadth. 



verado Canyon, 1025'N, 150'E of SW cor. sec.8, T5S, 
R7\V, El Toro quadrangle, Santa Ana Mts., Orange 
County, California. Ladd Formation, top of Baker Can- 
yon Sand.stone Member; Turonian. 

Dimensions: See Table 3. 

Age: Late Turonian 

Geographic distribution: Known onK Ironi the Ladd 
Formation, Baker Canyon Sandstone and lower Holz 
Shale Members, Santa Ana Mountains, Orange CounW, 
California. (LACMIP loc, 15295, 1 spec; CIT loc. 1064, 
1 spec; LACMIP loc. 10898, 2 spec; CIT loc. 1058, 3 
spec; CIT loc 454, 1 spec; CIT loc 1065. 1 spec; LAC- 
MIP loc. 10955, 2 spec). 




'^, 



■f. 



\,l 



/ 





•U^ 



1 



Figures 2-7. Afxtrrhai.s drarhiiki n. sp., all X 1 and whitened 
with anunonium chloride. 2-3. Holotype LACMIP 11373 from 
LACMIP loc. 15295, 2, aperture, 3, back view. 4-5. Paratyjie 
LACMIP 11375 from CIT 1058, 4, back view, 5, aperture.' 6- 
7. Paratype LACMIP 11377, from CIT 1064, 6, aperture, 7, 
back vdew. 



Remarks: Aporrliois dracltiiki mav be the earliest de- 
scrilied A/)()//7irt;.s. Wenz (1940) listed A/)<»;;7irt;.s as Sen- 
onian (Europe) to Recent (Europe), but Roy (1994) de- 
picts it as being present in the Cenomanian. Sohl (1967) 
reviewed North American species ascribed to Aporrhais. 
He concluded that Aponlwis hiauindata ( Meek and 
Hayden, 1856) ot Maastrichtian age Irom the upper part 
of the Pierre Shale ot Wyoming and Montana and an 
undescribed species of early Campanian age troni the 
Blutttowni Formation of Alabama were the onl\ actual 
Aporrhais from the Cretaceous of North America knovvii 
to him. Kase (in Kase and Maeda, 1980) described Pcr- 
issoptera elcofins Kase, 1980, of Barremian age, a species 
vyith an anterior digitation that is more spik"\' than lobate 
and not t\pical of Pvrissoptcra Tate, 1865. The spiky 
labral cbgitations are suggestive oi Aporrhais. but P. ele- 
oans appears to have no posterior digitation adnate to 
the spire. Pcrissoptcra cicgans is considerably earlier 
than Aporrhais drachiiki and has characteristics that may 
be intermediate between Drcpanochiihis imd Aporrhais. 
Aporrhais drachiiki differs from A. biangulata in having 
stronger nodes on its two prominent spiral cords, a more 
angulate whorl profile to the spire, and better developed 
;mal ribs. In having a well nodeil adapical cord and no- 
ticeable third cord, Aporrhais drachiiki is more similar 
to A. pespelecani than is A. biangulata. The primary spi- 
ral cords of A. drachiiki are relatively stronger and the 
nodes on them less prominent; the two primary cords 
are more nearly equal (the posterior cord is only slightly 
stronger); and indications are for a posterior apertural 
digitaion that is much shorter than in A. pespelecani. 
The over all shape and sculpture pattern are very similar. 
The posterior edge ot the outer lip In A drachiiki has a 
well developed posterior sinus that causes the lip edge 
to bend back'ward between the posterior adnate tligita- 
tion aTid apparent posterior labral digitation. A complete 
distal edge to the wing is not presened in any available 
specimen of A. drachiiki. None of the specimens pre- 
serves a digitation adnate to the spire. Paratypes have 
extensions of callus adapically from the posterior end of 
the aperture that may be remnants of an adnate poste- 
rior digitation. 



L. R. Saul, 1998 



Page 123 






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Rov (1994) suggests that m Aponiiai.s of Cretaceous 
age, the apertures are less heavilv callused than in A. 
pc.spchraui^ The htjlohpe of A. (Iracliiiki is onK Ughtly 
callused about the aperture, l)ut it mav not have been 
sufficiently mature to haw developed a complete callus 
deposit. Parat\pe 11375 has a thicker callus deposit on 
the inside of the outer lip, but it, too, is not as thick as 
that ol A pc.spclccdni. 

Etyinologj-: The species is named for Robert Dra- 
chuk, an amateur collector, who found the holotvpe of 
this species and donated it to VV. P. Popenoe. 

Genus Pcrissoptcra Tate, 1865 

T>pe species: RostcUaha parkiitsoni Mantell, 1822, 
by subsequent designation, Cossmann, 1904, Albian, Eu- 
rope. 

Discussion: The genera Pciissoptcro. DrcpanochciUis 
Meek, 1864, Arrlioncs Gabb, 1868, Latiala Sohl, 1960, 
and Gracihala Sohl, 1960 (See Figure 8) have been con- 
sidered to be closely related. Sohl (I960: 1967) placed 
Pcrissoptcra and Laiiala as subgenera of Arrhogcs, and 
derived Graciliala from Drcpauochcihis. Tate (1865), in 
proposing Pcrissoptcra as a subgenus oi Aporrbais. twice 
stated that the t)pe species of a section o{ Aporrhais was 
A. occklciitalis (Beck, 1836) (the species named bv Gabb 
(1868) as t)pe ol Arrhogcs) and UTote that this section 
corresponded with his new subgenus Pcrissoptcra, which 
had characteristics intermediate between Aporrhais and 
Alaria Morris and Lycett. 1850. If Tate had actualh- des- 
ignated A. occidcntahs as t\pe species of Pcrissoptcra. 
his action would ha\'e proceeded Gabbs ( 1868) clear and 
unequivocal designation o{ Chcnopus occidcntahs (Beck, 
1836) as type species oi Arrhogcs, and Arrhogcs would 
be an objective jiuiior sviionvin of Pcrissoptcra. 

Gabb (1868) commented upon Tate's paper m an ad- 
dendum. He quoted Tate's 'essential characters" of Pc;- 
issoptcra but considered it insufficientlv distinct from 
Anchura Conrad, 1860, and did not mention that Tate 
had named occidcntahs as t\pe species of am- group. 

Gardner (1875, p. 51), who made no mention of 
Gabbs work, stated that Tate had designated A. occi- 
dcntahs as type species oi Pcrissoptcra. Gardner argued, 
however, that there was no necessit\- for a supraspecific 
name for species better classed as Aporrhais. Group 1, 
and he did not use the binomen Pcrissoptcra occidcn- 
tahs. Gardner (1875) did not subsequently designate a 
t\pe species for Pcrissoptcra because he considered Tate 
to have named A. occidcntahs as tvpe species. 

Cossmann (1904) recognized Pcrissoptcra Tate as a 
useful taxon, in effect using it for Tate's Section 1, par- 
kinsoni group, and gi\ing the type species as RostcUaria 
parkin.wni Mantell, 1822, Albian. Cossmann's footnote 
that refers to Tate's (1865) "fig. 18 (sub. nom. Rost. 
Rciissi. non Gein.)" is part of the reference to Tate's 
proposal of the genus and apparently not part of Coss- 
mann's designation of the type species. Cossmann (1904, 
p. 95) stated that his diagnosis oi Pcrissoptcra was based 



Page 124 



THE NAUTILUS, Vol. Ill, No. 4 




'es 




Ferissoptera 



y Latiala 




Drepanocheilus 



Figure 8. Line drawings of five aporrhaid genera, all XL 
Species serving as examples for the genera are: Arrhoges oc- 
cidentalis (Beck, 1836), Recent, North Atlantic; Graciliala cal- 
caris (Wade, 1926) Cretaceous, Gulf Coast; Pcrissoptera par- 
kinsoni (Mantell, 1822), Cretaceous, England; Dn'pnnorlwilus 
evnnsi Cossniann, 1904, Cretaceous, Western Interior; Latiala 
lohatn (Wade, 1926), Cretaceous, Gulf Coast. 



on Ckinlru'r's (1875, pi. fi6) figures of the t\pe species 
and Rostcllaria mar^inota Sovverby, 1836. Rostcllaiia 
parkinsoni Mantell, 1822, is the first species and Ros- 
tellaria rcussi Geinitz, 1842, is the second species listed 
by Tate in his Section I, Parkinsoni group. Wenz (1940, 
p. 923) gave Aporrhais (Pcrissoptera) reussii Tate 1865 
non Geinitz = Rostcllaria parkinsoni Mantell as type of 
Perissiyptera. Dockery (1993) gave the type species as 
Rostcllaria rcussi Tate original designation. 



Tate (1865) did not directly designate any type species 
for Pcrissoptera. He wrote that Aporrhais pcspclccani 
was the t\pe species of the first section of Aporrhais and 
a second section of Aporrhais had for its type the li\'ing 
A. occidcntalis., and later stated that Pcrissoptera, "in- 
termediate between Alaria and Aporrhais, corresponds 
with that section of Aporrhais which has A. occidcntalis 
as its type". In presenting his arrangement of the fami- 
lies Strombidae and Aporrhaiidae he included in Apor- 
rhiiiidae "the e.xtinct genera Alaria. Diatihcma, and Pcr- 
issoptera ' As Tate considered Pcrissoptera to be extinct, 
the Recent A. occidcntalis could hardlv "serve as" its 
type species. 

Tate divided his new subgenus Pcrissoptera into three 
sections: Section I. Parkinsoni group; Section II. Cal- 
carata group; and Section III. Glabra group. Tate's topic 
was the classification of Rostellariae of the Cretaceous 
rocks, and, as might be expected, A. occidcntalis, a Re- 
cent species, is not included in any of Tate's three sec- 
tions oi Pcrissoptera. Thus, Tate not onlv did not directly 
designate occidcntalis as type species of Pcrissoptera, 
but he actuallv excluded it as a possible type species by 
characterizing Pcrissoptera as extinct, and he did not list 
A. occidcntalis among the species he included within 
Pcrissoptera. 

Mantell (1822) provided the name Rostcllaria parkin- 
soni for a specimen from the Albian of Blackdown fig- 
ured by Parkinson (1811), but he also referred to a spec- 
imen of early Cenomanian age from the Gre\' Chalk. 
Tate (1865) mistakenlv indicated that the Blackdown 
specimen was from the Grey Chalk and referred speci- 
mens from the Blackdown Greensand to Aporrhais (Pcr- 
issoptera) reussii (Geinitz, 1842), but these are not reu.s- 
si of Geinitz which is from the BacuUtenschichten of 
Zatschke and is of Turonian age (KoUmann, 1978). KoU- 
mann (1978) rexdewed Pcrissoptera parkinsoni, dis- 
cussed its occurrence, and considered the most repeated 
figure perporting to be P. parkinsoni (Mantell) (i.e.. 
Gardner, 1875, pi. 6, fig. 4; Wenz, 1940, p. 924. fig. 2705; 
etc.) to be an undescribed species, but both Kollmann 
(1978) and Tate (1865) quoted the same figures of Man- 
tell (1822, pi. 18, fig. 1, 4, 5. 10) as being Pcrissoptera 
parkinsoni (Mantell, 1822). 

Pcrissoptera has been widely used for moderately high 
spired aporrhaids having the axial sculpture much stron- 
ger than the spiral, and an expanded outer lip that is 
drawn out at its posterior e.xtremitv into a blade set off 
from the lobate medial distal margin bv a sinus (Figure 
8). The t\pe species has a relativelv short and wide ros- 
tnnn and is moderately carinate on the last whorl. In- 
cluded within Pcrissoptera are species with a longer ros- 
tnmi and stronger spind sculpture than are in Arrhoges, 
as well as a distinctive outer lip drawn out posteriorly 
and channeled interiorly Pcrissoptera differs from La- 
tiala in usualK haxing the spiral sculpture stronger and 
in haviTig the outer lip sinused on its distal margin, just 
anterior to the posterior extension, which is channeled 
interiorly. 

Pcrissoptera occurs more commonK in Europe than 



L. R. Saul, 199S 



Page 125 



in America, although the genus is present in the Western 
Interior (Perissoptcria prolabiata (White, 1876), Cenn- 
nianian) and the Gulf Coast (Pcrissaptciia prolahkita 
mississippicnsis Dockerv, 1993, Campanian). The onK' 
species described from Pacific Slope deposits is Pciis- 
soptera hannai (Allison. 1955) from the Aptian-Albian 
Ahsitos Formation of northern Baja Cahfornia, Mexico. 

Genus Laiiala Sohl, 1960 

Type species: Aiuhiira lohata Wade. 1926, hv original 
designation. Maastrichtian of the Gull Coast of North 
America. 

Discussion: S(^hl (1960) chfferentiated Latiala from 
Arrhogcs on the basis oi Latiala's broad, thick outer lip 
which is thickened and lobed at the anterior and pos- 
terior termini of its chstal margin, the more pointed and 
longer lobe directed posteriorward and the blunter one 
directed anteriorward. He found Latiala to be wideK' 
distnbuted [i.e.. New Jersey, Tennessee, Mississippi, 
Texas, India, and South Africa) during the late Late Cre- 
taceous. Abdel-Gawad (19S6) has added Latiala pelc- 
cyphora (Kaunhowen, 1897) of late Maastrichtian age 
from the Mitklle \'istula \'alle\- of Poland. Pacific Slope 
species resembling Latiala are, howe\'er, of Albian 
through Turonian age, and Kase (1984) reported Japa- 
nese Latiala of Aptian-Albian age. In Pacific Slope sed- 
iments, juvenile shells are more common and more 
abundant, especially in fine-grained, more off-shore de- 
posits, than are the mature shells. Juvenile shells of La- 
tiala appear to have been more readily transported into 
deeper water than shells of other gastropods with which 
the adult Latiala spp. are associated. Popenoe (1983), in 
assigning Pacific Slope specimens to Latiala, discussed 
only A. (L. ) califomicus. in which he included "Alalia" 
nodosa Packard and some specimens herein assigned to 
L heliaca n. sp. and L. sigma n. sp. 

Dockers' (1993, p. 61) raised Latiala to a genus based 
on its having a thinner, more fragile shell than that of 
Arrhoges, a heUcocone similar to that of Graciliala. and 
a body whorl without fine spiral firae, but Rov (1994) 
did not separate Latiala from Arrhogcs. Unfike Graci- 
liala, but as in Arrhogcs, the wang of Latiala is usually 
unchanneled. Specimens of Latiala lohata (Wade) from 
Coon Creek, Tennessee, show fine spiral striae under 
sufficient magnification even on the bod\' vs'horl, but 
sculpture of Pacific Slope species does differ from that 
of the type species in ha\dng a few noticeablv stronger 
lirae adjacent to the posterior suture. The Pacific Slope 
species herein assigned to Latiala varv in shell thickness; 
L. californica is thin shelled enough to meet Docker\'s 
specifications, but L nodosa has a thicker shell that is 
not more fragile than that of Arrhogcs. All of these spe- 
cies have some development of spiral firae on the bodv 
whorl, but in L. californica the lirae are so fine, and in 
L. nodosa the preservation of most specimens is suffi- 
cientlv poor, that the firae are commonlv obfiterated. La- 
tiala heliaca has the strongest lirae, but thev are not as 



strong as those oi Arrhoges occidciitalis. Latiala heliaca 
and Latiala sp. from Grave Creek are more slender and 
have a longer rostrum than Arrhoges. Of the five Pacific 
Slope Latiala species considered here, L. nodosa is most 
similar to Arrhoges in its stocky shape and thicker shell, 
liut L. nodosa has the anterior and posterior wing ex- 
tensions of Latiala. and its shape is similar to that of the 
thin shelled L sigina, which has the longer rostnnn and 
longer posterior wing extension of a Latiala. The most 
notable cfifference behveen these five species of Latiala 
lies in the sculpture of the spire and especiallv in the 
number and placement of iLxial ribs. A common feature 
of these species is the sculpture pattern on the spire of 
arched, somewhat opisthocfine ribs crossed bv fine spi- 
rals, a few of which are strongest adjacent to the pos- 
terior suture. 

This sculpture pattern is one of the features that La- 
tiala shares with, at least some species of Peiissoptera 
Tate, 1865. In its early whorls, sculpture, and shape, ex- 
clusive of the wing, Latiala resembles Peiissoptera. 
Wing development of more than one Perissoptcra spe- 
cies passes through a stage in which the shape of the 
wing outfine resembles that of Latiala or Arrhoges. A 
small specimen (Figures 9-10) in the LACMIP collec- 
tions labeled Peiissoptera rohinahlina (d'Orbignv, 1843) 
from the Aptian of Atherfield, Isle of Wight, Great Brit- 
ain (LACMIP loc. 11612), resembles Gardners, 1875, 
pi. 7, fig. 11 of P. rohinahlina. The distal margin of its 
wing is thickened anterior to a slight sinus at the base 
of the posterior wing digitation. 

Although Sohl (1960)"and Roy (1994) include Latiala 
as a subgenus of Arrlwges, and Sohl ( 1960) derives Ar- 
rlioges from Drepanoclieiliis. the species placed in La- 
tiala bv Kase (1984) and herein, suggest that Latiala ap- 
pears in the geological record before Arrhogcs, and Ar- 
rhogcs might more reasonablv be derived from Latiala. 
Dockerv (1993) notes that the heficocones oi Graciliala 
and Latiala are similar, but that Graciliala has a chan- 
neled outer lip and Latiala iloes not, a feature that lends 
support to Roy's (1994) placement oi Graciliala nearer 
to Drepanochcihts than to Arrhoges. Drcpanochcilus and 
Peiis.'ioptcra haxe both been recorded from the Barre- 
mian bv Rov (1994) and Kase (1980 in Kase and Maeda; 
but see remarks on Perissoptcra elegans under Apor- 
rhais drachiiki) respectively; Latiala is first recognized 
in the Aptian (Kase, 1984); Graciliala apparentlv is 
knowni onlv from the Campanian and Maastrichtian; 
Sohl ( 1960) intlicated that Cretaceous forms classed as 
Arrhoges belong either to Latiala or Peiissoptera, and 
Arrhoges is present in the Paleocene (KoUmann and 
Peel. 1983). 

Popenoe (1983) assigned Latiala from the Santa Ana 
Mountains to Arrhoges (Latiala) califomicus which was 
described from the Siskiyou Mountains (Gabb, 1864). 
He also included in A. (L.) califomicus specimens of 
Cenomanian to Turonian age which show considerable 
variation in number of axial ribs, height of spire, strength 
of spiral firae, and length of rostrum. He evidently in- 
tended further studv of this group as he had on hand 



Page 



126 



THE NAUTILUS. Vol. Ill, No. 4 



'i. 



\ '. 




20 




25 

Figures 9-;J(). All whiteneil with amiiioniiim chloride. 9-10. Prrissoptcni robinald'tna (d'Orbigny. 1842). h\pot)pe LACMIP 
11536, from Atherfield, Isle of Wight, Great Britain, x2. 9, back view, 10, aperturai view showing wing edge. 11-13, 19. Latinla 
sp., all X2, latex pulls from LACMIP loc. 24670. 11, LACMIP 7969. back \ie\\. 12, L.\CMIP 7968. aperturai \iev\. 13, LACMIP 
7966. back view. 19, two specimens, arrow points to LACMIP 7967. 14-18, 20-21. Latialn luiiaca n. sp.. 14, holotvpe LACMIP 
11.378 from LACMIP loc. 28788. xl, back \iew. 15, parat\pe LACMIP 11379 from LACMIP loc. 28788. X3. back \iew. 16, 
paratvpe LACMIP 11380 from LACMIP loc. 28788, X4, protoconch and first ju\enile whorl, 17, paratvpe LACMIP 11382 from 
LACMIP loc. 16838, X2, back view, 18, paratyi^e LACMIP 11383 from LACMIP loc. 16838, X2. left side. 20, paratApe 11381 
from LACMIP loc. 16838, X2, back view, 21, parat>pe 11384 from LACMIP loc. 16838. x2, wing in rock containmg abundant 
L. heliaca. 22-30. Latinla califomicn (C.abb), 22, lectotvpe ANSP 4772 from the Siskiyou Mountains?, X2, photo by T. Susuki, 
23, paralectotvpe ANSP 4772 from the Siskivou Mountains?, X2. photo bv T, Susuki. 24,'b\pot\pe LACMIP 11390 fro'm LACMIP 
loc. 10901. X2. back \iew, 25-2(>, hvpohpe'LAC;MIP 11388 from LACMIP loc. 10901, x2, 25, aperturai \iew, 26, back view, 27, 
hvjwtype LACMIP 1138.5 from UCIA loc. 4365. X2, back \iew, 28, hNp(>t\pe LACMIP 1 1387 from UCLA loc. 4365. X3. aperturai 
view, 29, hvpotvpe LACMIP 11386 from UCLA loc. 4365. X2, ai)erturarvicw. 30. hvpotvpe LACMIP 11389, from LACMIP loc. 
10901. Xl.5. back view. 



L. R. Saul, 1998 



Page 127 



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specimens of Cenonianian age from Clear Creek, Shasta 
County, and Sunflower Valley, Fresno County, that he 
had labeled Arrhoges (Latiala) new species. Specimens 
of Latiala of Turonian age from the Santa Ana Moun- 
tains re\eal differences in the sculpture ul' the early 
whorls and in onset of sculpture changes on the spire, 
as compared to specimens of L. califomica from the Sis- 
kiyou Mountains of the Oregon California border On 
the basis of these differences, L. califomica is discrimi- 
nated from L. nodosa and considered not to be found 
in southern California. 

Latiala sp. 

Figures 11-13. 19; Table 4. 

Diagnosis: A fusiform Latiala with about 18-24 axial 
ribs on spire whorls, reduced to about eight on body 
whorl; body whorl with rilxs not confined to back of 
whorl. 

Description: Shell of medium size, spindle shaped, 
except for expanded outer lip; whorls about twice as 
wide as high, numbering about seven or eight whorls, 
last whorl approximately half of shell height; pleural an- 
gle about .30°; whorl profile on spire roundly convex; 
profile of last whorl roundly convex except at axial nodes; 
outer lip expanded, broad, subquadrate, with short an- 
terior and longer more pointed posterior projection. 
Protoconch unknown. Sculpture of 18-24 arched, 
round-topped, axial ribs, crossed bv much weaker, fine 
spiral cordlets, strongest adjacent to posterior suture, 
two cordlets commonly strongest; sculpture on body 
whorl of eight to nine, short sharp, strong axiid ribs 
crossed by fine spiral cordlets; ribs on back of body 
whorl shorter, stronger, more nodular. Aperture elon- 
gate; inner lip thickly callused with well defined edge; 
rostmm narrow, of moderate length. 

Material examined: LACMIP 7628, 7966-7969 from 
LACMIP loc 24670 on Grave Creek, Jackson County, 
Oregon. 

Dimensions: See Table 4. 

Age: Late early to middle Albian. 

Geographic distribution: Studied specimens are all 
from the Hornbrook ? Formation on Grave Creek, Jack- 
son County, Oregon 

Remarks: This undescribed species is left unnamed 
because the specimens studied are all rock molds, and 
the figures are from latex pulls. The protoconch was 
present on some of these specimens, but the sandstone 
is too coarse-grained to yield a latex pull that replicates 
the original surface. Sculpture of the spire is similar to 
that of Latiala hayamii (Kase, 1984), but Latiala sp. has 
a broader pleural angle. Latiala sp. differs from L. he- 
liaca in having more axial ribs on the body whorl, some 
of which are on the apertural side of the whorl. Latiala 
sp. has a proportionately higher wing than Latiala liclia- 
ca, L. califomica, or L. sigma. 



Page 128 



THE NAUTILUS, Vol. Ill, No. 4 



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Latiala hcliaca n. sp. 

Figure.s 14-18, 20-21; Table 5. 

Arrhoges califomicus (Gabb) — Murphv and Rodda, 1960, p. 
841, pi 102, fig. 6-7. 

Diagnosis: A ,slender fusiform Latiala with ahout 16 
axial rib.s on spire whorls, greatlv reduced on final whorl 
to two or three on dorsal side of body whorl. 

Description: Shell of medium size, spindle shaped, 
e.xcept for expanded outer hp, of about nine whorls; 
whorls about twice as wide as high; last whorl approxi- 
mately half of shell height; pleural angle about 36°; 
whorl profile on spire roimdly convex; profile of last 
whorl roundlv convex except at axial nodes and with 
broad axial bulge on apertural face; outer lip ex-j^anded, 
broad, subquadrate, slightly thickened at distal edge, 
with short anterior and longer posterior projections. Pro- 
toconch of four or five rapidlv enlarging, rounded, glossy 
whorls; transition to juvenile sculpture gradual, marked 
bv spaced faint, arched, axial ribs and spiral striae; ribs 
rapidly strengthened, becoming crowded. Sculpture on 
first juvenile whorl of 26-36 arched axi;il ribs crossed bv 
much weaker fine spiral cordlets; sculpture of next 
through penultimate whorls of about 16 arcuate axial 
ribs crossed bv fine spiral cordlets, strongest adjacent to 
posterior suture, three cordlets commonK- strongest; 
sculpture on bodv whorl of three to four short, sharp, 
strong axial ribs crossed by fine spiral cordlets. Aperture 
elongate; inner lip broadly callused, callus angling from 
posterior of aperture across mid-whorl on apertural face 
and wrapping halfway around anterior rostmm; rostnnn 
narrow, straight, relatively short. 

Type specimens: Holot\pe, LACMIP 11378. Para- 
t>pes LACMIP 11379-11380 from LACMIP loc. 28788; 
UCLA 28625 = LACMIP 9825 , UCLA 28626 = LAC- 
MIP 9824 from UCLA loc. 3465, and LACMIP 11381- 
11384 from LACMIP loc. 16838, Ono area. Shasta 
County, Cahfomia. 

Type locality: LACMIP 28788, north end of Simflow- 
er Vallev ( = McLure Valley), Fresno Count)', Cahfoniia. 
Panoche Formation. 

Dimensions: See Table 5. 

Age: ?Late Albian-Cenomanian. 

Geographic distribution: Budden Canvon Forma- 
tion, Bald Hills Member, Ono area, Shasta Countv 
(UCLA loc. 3465; Great V;dlev Series, Elder Creek, Te- 
hama County (LACMIP loc. '24370); Panoche Forma- 
tion, Sunflower Valley, Fresno Countv (LACMIP loc. 
28788), Cahfomia. 

Remarks: No \arices were recognized on the earK' 
wliorls of L hcliaca. LACMIP 11380 consists of a pro- 
toconch of 4 polished, rounded whorls and the first te- 
leoconch whorl. The earliest axial ribs are irregular, low, 
and (hfficult to count, but on the second quarter of the 
first teli'oconch whorl the ribs become stronger, more 
(hstant and more regular Rib strength varies between 



L. R. Saul, 1998 



Page 129 



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iiidhiduals, and on some \vh(jrls the axial rihs inav be 
indiscemibie. 

Specimens from Elder Creek (LACMIP loc. 24370) 
are from a boulder containing species suggestixe of late 
Albian age in a conglomerate that is of Tunjnian age 
(Jones and Bailev, 1973). 

Laiiala hcliaca is larger and more slender than L. cal- 
ifoniica and has fewer axial ribs. Whereas L. califomica 
retains tlie fine axial ribbing of the juvenile whorl 
throughout most of the spire whorls, in L. hcliaca the 
intermecbate whorls have fewer, coarser ribs, and the 
body whorl is not angulate at the shoulder, although the 
nodose axial ribs can create the impression of a shoulder 
angulation. Latiala I}clica has more axial ribs on the spire 
and fewer on the body whorl, is more slender with a 
more fuisform shape, and has a slightK- liroader shorter 
wing than L nodosa. Latiala hclica is more slender, has 
less ot a shoulder, fewer axial ribs, and a shorter wing 
than L. si^ma. 

Latiala hcliaca is most similar in shape to Latiala haij- 
amii (Kase, 1984) from the Aptian-Albian of northeast 
Japan, but differs in having fewer axial rilis on the body 
whorl, 

Et}'niolog\-: The species name, derived from Greek, 
hclios, sun, refers to the type localitv' at the northwest 
end of Sunflower Valley. 



Latiala califomica (Gabb, 1864) 
Figures 22-30; Table 6. 

Aporrhais califomica Gabb, 1864, p. 128, pi. 29, figs. 2.30a, b. 
Arrhofies califomicns (Gabb)— Stewart, 1927, p. .36.3, pi, 21, 

fig. 1,5. 
Arrlwf^es (Latiala) cnlifomicus (Gabb). Popeiioe, 1983, p. 761 

(in part; not fig. 6F, I. See L. nodosa Packard). 
Not Arrhoges califomicns (Gabb) — Mui-phy and Rodda, 1960, 

p. 84!, pi. 102. fig. 6-7. See L. heliaca n. sp. 
Not Arrho^cs califoniiais (Gabb) — Jones, Sliter and Popenoe, 

1978, p. xxii.9, pi. 1, fig. 10. See L. nodosa Packard). 

Diagnosis: A small Latiala in which axial sculpture of 
spire consists of fine, closely spaced, almost sigmoid ribs. 
Sculpture of body whorl of three to four strong, short 
ribs, present only on dorsal side, nodular at the shoulder. 

Description: Shell, thin, of small to medium size, 
spindle shaped except for expanded outer lip; whorls 
nearly twice as wide as high, numbering about seven; 
last whorl appro.ximately two-thirds to three-quarters of 
shell height; pleural angle about 35°; whorl profile on 
spire rounded; profile of last whorl angulate at shoulder 
and with broad axial bulge on apertural face; varices rare 
(Pone juvenile varix per specimen), forming broad axial 
.swellings; outer fip expanded, broad and moderately tall, 
with very' short anterior and much longer posterior pro- 
jections. Protoconch multispiral of five or slx smooth, 
rounded whorls. Sculpture on body whorl of about four 
short, nodular axial ribs on dorsal side, a broad axial 
bulge on the apertural side and fine spiral striae, stron- 
ger spiral cordlets on shoulder and on collar; sculpture 



Page 130 



THE NAUTILUS, Vol. Ill, No, 4 



on piMiiiltimate wliorl of about 20 fine ni'arlv sigmoid 
axial rih.s and fine (about four per mm), regularly .spaced 
spiral cordlets, strongest near posterior suture; sculpture 
on earlier whorls finer than on pemiltimate whorl. Ap- 
erture elongate, outer lip thickened internalv; distal edge 
of outer hp relati\elv long; rostnnn short. 

T\pe specimens: Lectot\pe ANSP 4272 (designated 
bv Muipln and Rodda, 1960). Paralectotxpes (2) ANSP 
79470; CASG 61950.01 (one loose specimen + a rock- 
fragment containing several, two largely exposed). 

Hvpotvpes: LACMIP 11385-11388 from UCLA loc. 
4.365. Redding area, Shasta Count\-; and LACMIP 
11388-11.390 from LACMIP loc. lOyOl. Rancheria 
Gulch, Siskivou Countv, California. 

Type locality: Siskiyou Mountains?, Jackson Countv, 
Oregon or Siskiyou County, Cahfornia. Gabb (1864, p. 
129) listed this species from "Orestimba Caiion; Marti- 
fiez; Puerto Caiion, Stanislaus Countv; Siskivou Moun- 
tains." None of these was precise as to geographic or 
stratigraphic position. Stewart (1927) said that the label 
with Gabb's ANSP specimens read Siskivou Mts. and 
Martinez, but that all of the specimens were in a similar 
black hmestone matrix with small pebbles which he con- 
sidered more likely to have come from the Siskiyou 
Mountains than from Martinez. Natural Histon,- Muse- 
um of Los Angeles Countv specimens resembling Gabb's 
material are from the Osburger Gulch Sandstone Mem- 
ber of the Hombrook Formation in the Siskiyou Moun- 
tains, but are not identical in preservation to Gabb's Sis- 
kiyou M(nmtains specimens. 

Dimensions: See Table 6. 

Age: Turonian. 

Geographic distribution: Hornbrook Formation, 
Osburger Gulch Sandstone Member (LACMIP loc. 
109()l! LACMIP loc. 10876, LACMIP loc. 25422), Jack- 
son County, Oregon and Siskiyou Counts', California; 
Redding Formation, PFrazier Siltsone Member (UCLA 
4365) near Redding, Shasta Countv', California. 

Remarks: The CASG "sviity|")es" have a label th;it lists 
SUT 266, Calif Geol. Surv. 2144, and UCB 14907, m- 
dicating that the specimens were originallv at Universitv 
of Cahfornia, Berkeley, Museum of Paleontolog)' before 
they were moved to Stanford and from there to the Cal- 
ifornia Academv of Sciences. 

The lectotype is 15 mm high, nearly complete, and 
has an adult aperture, but is missing shell on the last 
two whorls. The absence of parts of the shell interfers 
with describing the sculpture. The ANSP and CASG 
type specimens are small, but some collections (r i; , 
L/VCMIP loc. 10901) contain a few larger indi\iduals 
(Figure 30), The above description depends in part on 
additional specimens from the rather generalized ty|ie 
localit) oi Siskiyou Mountains. Spec imens are commonly 
found locally in abundance as in the blocks of paratype 
material, but the very thin shell oi this species has made 



difficult the rec<)\er\' of specimens with shell adhering. 
This species exhibits juvenile sculpture onto the penul- 
timate whorl on small adults and up to the penultimate 
whorl on large adults. 

Ciabbs illustration (fig. 230a) has an outline more like 
that of Latiala hcliaca than of L. caltjornica and ma\' 
have been based on a specimen of that species. If so, 
the specimen appears to have been lost. 

Latiala califoruica differs from L. hcliaca in having an 
angiilate shoulder and more axial ribs on most whorls. 
It chffers from L. nodosa in having a very thin shell, 
retaining the fine axial ribbing onto or up to the pen- 
ultimate whorl. Latiala califoruica is smaller, has weaker 
spiral sculpture, and fewer axial ribs on the bod\' whorl 
than L. .si^iiia. Of the five species discribed in this paper, 
L ralifoniica luis the relativelv shortest wing. 

Latiala .si^ma n. sp. 
Figures 31-43; Table 7. 

Diagnosis: A relatively large Latiala with relatively 
strong spiral sculpture, numerous fine axial ribs on earlv 
whorls, and about seven ribs on bodv whorl. 

Description: Shell of medium size, stoutly spindle 
shaped, except tor expanded outer hp; whorls about 
twice us wide as high, nine in large mature inchviduals, 
seven in small mature individuals; last whorl approxi- 
mately two-thirds of shell height; pleural angle about 
36°; whorl profile on spire nearlv flat sided with narrow 
rounded shoulder; whorl profile of last tvvo whorls rather 
evenly rounded; varices poorly developed on early 
whorls; outer fip e.xpanded, tall and broad, subquadrate 
with verv short, blunt anterior and narrow posterior pro- 
jections; angle of chstal edge of outer lip to that oi sliell 
axis about 33°. Protoconch of four rounded, glossy 
whorls. Sculpture on winged body whorl of five to seven 
short axial ribs on dorsal side, on penultimate whorl of 
;ibout 20 to 30 cuned axiiil ribs and three narrow spiral 
cordlets at the sutiue; sculpture on first (juarter of first 
teloconch whorl of fine axial fines, at least eight per 1 
mm, axial lines double in size and bv third teleoconch 
whorl are four to 1 mm; second teleoconch whorl with 
about 40-50 fine axial ribs crossing about 20 spiral cor- 
dlets; cordlets weakest mid whorl and strongest poste- 
riorly. Aperture elongate, outer fip not nuich thickened; 
inner lip somewhat thickened; rostnnn relativelv long. 

Type specimens: Holohpe LACMIP 11391. Para- 
t\pes L.\C.V1IP 11.392-11398 from UCLA loc. 5421, 
LACMIP 11.399-11401 from CIT 1346, and LACMIP 
11402 from CIT 1266, all on Little Cow Creek, Redchng 
area, Shasta Countv, California. 

T\pe locality: UCLA loc. 5421, Little Cow Creek, 
south line sec. 9, T32N, R3W, MilKille cjuad., Shasta 
County, California. Redchng Formation, Melton S;md- 
stone. 

Dimensions: See Table 7. 

Age: Turonian. 



L. R. Saul, 1998 



Page 131 




Figures 31-52. All whitened with ammonium chloride. 31-43. Latiala sigrna n. sp., 31, holot\pe LACMIP 11391 from UCLA 
loo. 5421, xl.5, back \ie\v, 32, parat\pe LACMIP 11392 from UCLA loc. 5421, xl, apertural \iew, 33, paratApe LACMIP 11400 
from CIT loc. 1.346, xl.5, back view, 34, paratype LACMIP 11.393 from UCLA loc. 5421. Xl, back \iew, 35, paratvpe LACMIP 
11402 from CIT loc. 1266, Xl.5, showing jmenile to mature sculpture, 36, paratvpe LACMIP 11.396 from UCLA loc. .5421, X4, 
protoconch and two juvenile whorls, 37, paratope LACMIP 11398 from UCLA loc. 5421, X4, protoconch and two juvenile whorls, 
38-39, paratvpe LACMIP 11.394 from UCLA' loc. 5421, X2, juvenile whorls, 40, paratvpe LACMIP 11397 from UCLA loc. ,5421, 
X3, protoconch, tsvo juvenile whorls, and a varlx, 41, paratvpe LACMIP 11395 from UCLA loc. 5421, x2, protoconch and three 
juvenile whorls, 42, paratvpe LACMIP 11401 from CIT loc. 1346, Xl.5. showing mature sculpture on last whorl, 43. paratvpe 
LACMIP 11417 from UCLA loc. .5421, xl, angle of outer lip. 44-52. Lntiala nodosa (Packard), hvpotvpes, 44, LACMIP 11406 
from CIT loc. 1065, X3, protoconch and three juvenile whorls. 45, LACMIP 11403 from CIT loc. 1164, Xl.5, back view, 46, 
LACMIP 1140S from CIT loc. 1.532, XI, apertural view, 47, 50, 52, LACMIP 1080S ( = UCLA 5S43S) from CIT loc. 1212, Xl, 
47, wing angle, 50, apertural view, photo bv T Susuld, 52, back view, photo by T Susuld, 48, LACMIP 11407 from CIT loc. 1212, 
X2, juvenile to mature sculptru-e, 49, LACMIP 11405 from CIT loc. 1164, Xl.5, apertural view, 51, LACMIP 11404 from CIT 
loc. 1164, X 1.5, back view 



Page 132 



THE NAUTILUS, Vol. Ill, No. 4 



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Geographic distribution: Reddino Formation. Mel- 
ton Sandstone, Redtling area, Shasta County, California. 
[CIT loc. 1264; CIT loc. 1265; CIT loc. 1266; CIT loe. 
1346 = UCLA loe. 5421] 

Remarks: Latiola sigma has the strongest spiral seulp- 
ture and longest rostrum of the Latiala discussed herein. 
All of the a\ailalile mature specimens have the shell bad- 
ly leached and the adult whorl surface is not preserved 
on any of them. Winged adults range in size from about 
22 mm high to 46 mm high. None of the available spec- 
imens shows evidence of thickening along the tlistal mar- 
gin of the wing. Obscure swellings that may be indistinct 
varices are present on some early whorls. Latiala sigirui 
has a more gradate spire, more axial ribs on the imma- 
ture whorls, and the distal edge of the wing is at a great- 
er angle to the shell axis than in L. nodosa. It is larger, 
has a broader wing, and has more axial ribs on its bodv 
whorl than L. califomica and L. heliaca 

Etymology: Sigma, the eighteenth letter of the Greek 
alphabet, was W. P. Popenoe's manuscript name for this 
species. 

Latiala nodosa (Packard, 1922) 
Figures 44-52; Table S. 

Ainrin nodosa Packard. 1922, p. 430, pi. 36, figs. 5a-5b. 
Arr/iogc.v califontictis (Gabb) — Jones, Sliter and Popenoe, 

1978, p. xxii.g, pi. 1, fig. 10. 
Arrhoges {Latiala) californicns (Gabb) — Popenoe, 1983, p. 76 

(in part), fig. 6 F, I. 

Diagnosis: A thick-shelled Latiala with about four te- 
leoconch whorls having 14 obhque axial ribs per whorl, 
seven to eight axial ribs on body whorl not restricted to 
back of whorl, and a broad wing. 

Description: Shell thick for a Latiala. of medium size, 
spindle-shaped, except tor expanded mature labmm; 
whorls twice as wide as high, numbering about nine; last 
whorl a Uttle over hidf of shell height; pleural angle 
about 37°; whorl profile on spire rounded but ribs create 
incipient shoulder; profile of last whorl roimded but with 
apparent angulation caused by nodes; conuiionlv one 
varix per submature whorl; outer lip expanded, moder- 
ately broad, subquadrate with short anterior and poste- 
rior projections; angle of distal edge of outer lip to that 
of shell axis about 22°. Protoconch of four smooth 
rounded whorls. Sculpture on fifth and sixth whorl of 
about 20 low, obli(jue, gently concave to aperture, crest- 
ed axial ribs, on penultimate whorl of 14 axial ribs; bodv 
whorl sculptured by seven to eight axial ribs forming 
nockjse shoulder; all post[")rotoconch whorls with fine spi- 
ral striae, strongest near the posterior suture. Aperture 
elongate, outer fip thickened and not channeled; callus 
deposit over apertural face of bodv v\horl extending as 
broail tongue over pi'uultiniate whorl to posterior su- 
ture; inner lip thin t'dged, mediallv thickened; rostrum 
short. 

Type specimens: Holotvpe UCBMP 12297. Paratvpe 



L. R. Saul. 1998 



Page 133 





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UCBMP 12296 from UCBMP loc. 2142, Santa Ana 
Mts., Orange County, California. 

Hypotype,s: UCLA 58438 ( = LACMIP 10808) (A. (L.) 
calif'ornicus of Popenoe) from ClTloc.1212 ( = LACMIP 
loe.10735), LACMIP 11407 from CIT loc. 1221, and 
LACMIP 11408-11409 from CIT loc. 1532, Redclrng 
area, Shasta Count)-; LACMIP 11403-11405 from CIT 
loc. 1164, Santa Ana Mount;iin.s, Orange Count\', Cali- 
fornia. 

Type locality: UCBMP loc. 2155, 5.43 km S 65°E of 
B.M. 610, Corona .sheet. 0.8 km from mouth of Black 
Star Caiion. Chico float. 

Dimensions: See Table 8. 

Age: Turonian. 

Geographic distribution: Ladd Formation, Baker 
Canyon Sandstone and lower Holz Shale Members, 
Santa Ana Mountains, Orange Countv, California, (CIT 
loc. 80, 1 spec; CIT loc. 82, 12 spec; CIT loc 99, 2 
spec; CIT loc 454, 1 spec; CIT loc 1065, 4 immature 
spec; CIT loc. 1164, 59 spec; CIT loc 1290, 1 spec; 
CIT loc 1307, 8 spec; UCLA loc 2325, 2 spec; LAC- 
MIP loc. 10953, 1 spec; LACMIP loc. 29181, 1 spec); 
Redchng Formation, Frazier Silt, Redding area, Shasta 
County- California, (CIT loc 1212, 16 spec); CIT loc. 
1221, 3 spec; CIT loc 1532, 14 spec; UCLA loc. 4658, 
15 spec). 

Remarks: Precise stratigraphic position of the holo- 
tvpe was unknown to Packard (1922). but the parat\pes 
are from UCBMP loc 2142, which is in the lower Holz 
Shale. The species has been collected from numerous 
locahties in the Baker Canyon Sandstone and lower Holz 
Shale Members of the Ladd Formation. 

Popenoe (1983) included L. nodosa in A. (L.) califor- 
nicus, but the early whorls of L. nodosa have fewer a.xial 
ribs per whorl, the earliest of which are so low that the 
spiral riblets adjacent to the posterior sculpture is the 
more noticeable sculptural element. The earlv a.\ial ribs 
of L. nodosa, are characteristicallv chstant and crested 
with the sharp slope toward the aperture. Latiala nodosa 
has a\i;il ribs on the apertural face of the body whorl 
but L. californica does not, and L. califomica has a bet- 
ter-defined shoulder and fewer varices on the earlv 
whorls. Although L. nodosa is common to abundant at 
more than four localities, few specimens have the ear- 
liest whorls well-preserved. LACMIP 11406 and other 
specimens from CIT loc. 1065 have well-preser\ed earlv 
sculpture; LACMIP 11407 appears to have nearlv ;ill of 
the early whorls, but the shell surface of the first four 
whorls is not preserved. Remnants of fine axial ribs are 
present on the fifth whorl and 14 ribs are on the sLxth 
whorl. The fifth whorl has hvo ribs for each rib on the 
sixth whorl. Callus on the apertural face of the body 
whorl is thick enough to obscure some of the axial ribs 
and forms a broad lump at the base of the body whorl 
adjacent to the aperture. It extends apicallv from the 
posterior end of the aperture forming a callus tongue 



Page 134 



THE NAUTILUS, Vol. Ill, No. 4 



aero.s.s the penultimate wlioil. A thin and faiiK narrow 
inner lip i.s developed (ner the callii.s. 

Redding area specimens of L. nodosa are larger, more 
robust, and ha\'e a wider pleural angle than those from 
the Santa Ana Mountains. In the Redding area, L. no- 
dosa and L. sigma are of similar size, but L nodosa has 
a thicker shell, a more cjuadrate wing, the distal edge of 
which is at a lower angle to the shell ;L\is, and early 
whorls with fewer a\ial ribs and weaker spiral striae. La- 
tiala nodosa is from the Frazier Silt liut L. siguia is from 
the Melton Sandstone. Of the five species of Lotiala de- 
scribed in tliis paper, L. nodosa has the tallest wing rel- 
ative to breadth and to shell height. 

Genus AlaiimcIIa new genus 

T\pe species: Aporrhais vctiis Packard, 1922 from the 
Turonian of southern Cahfomia. 

Diagnosis: A RimcHa-\ike aporrhaid with sculpture of 
nearly aligned ;i.\ial ribs and finer spiral cords but with 
an expanded outer lip. Whorls becoming carinate on 
body whorl; carina e.xtending along trigonally expanded 
outer lip; outer lip with a posterior digitation pro.ximal 
or adnate to the spire; interior of outer lip channeled 
opposite the carina. 

Discussion: Alaiinwlhi is verv similar in shape and 
sculpture to some strombids such as the Rimclla-\ike 
gastropods (Clark and Palmer, 1923) of the Rostellarinae 
(Delpey, 1941). Certain species of Cah/pfraplionts Con- 
rad, 1857, e. g., Cahjptraphonis hinodifcnis PerriUiat 
and Vega, 1997, Maastrichtian, southern Mexico, also 
have similar shape and sculpture beneath their mature 
callus covering. The shape of the aperture and extension 
of the posterior canal up the spire of Alaiiinella also are 
similar in these strombids. The most obvious difference 
is in the outer hp of Alaiiniclla which is alateh- expanded 
and interiorly channeled but in the RimcUaAike strom- 
bids is not. Additionally the aperture of aporrhiiids has 
sinus areas on either side of the rostnnn and a concave 
columella which accomodate the anterior end of the foot 
and the snout, but the strombid shell is less excavated 
across the anterior end of the columella. Strombs char- 
acteristically have a strombid notch near the anterior end 
of the outer lip which is used by the stalked eve of 
strombs (Abbott, 1960). The eyes of aporrhaids are on 
bulbs at the tentacle bases (Yonge, 1937) rather than on 
long, mobile pedicle tips and are not extendible as in the 
strombs. Possibly stalketl eyes evolvetl in advance of the 
notch through which they extend, antl absence of the 
strombid notch does not assure aporrhaid eyes. The 
strombid notch is not always well defined in strombs 
(Savazzi, 1991), and some of the Rinirlla-\\ke gastropods 
are Siiid not to have a strombid notcli, v. g. Rinwlla Ag- 
assiz, 1840, Macilentos Clark and Palmer, 1923 (subob- 
solete fide Clark and Palmer, 1923, lacking fide Vokes, 
1939), Cowlifzia Clark and Palmer 1923. Nonetheless, 
the absence of a strombid notch in Alariinclla in con- 
junction with the similarity of the sculpture of AlaiimcIIa 



on the whorls of the spire to that of Lafiala. Pcrissop- 
fera, anil Craciliala, the development on the bod>- whorl 
of Alarimella of a modest carina, and the possession by 
Alarimclla of an expanded outer lip that is intemallv 
channeled opposite the carina suggest that it be included 
in Aporrhaiidae. 

Varices on spires of Alariinclla icta and A. anac are 
similar to those on species of Rimella. Varices are said 
to be of little or no systematic importance in Stromba- 
ceans (e. g. Davies, 1971, p. 328), and their presence 
does not aiil in differentiating between Aporrhaidae and 
Strombidae, but Clark and Palmer (1923) thought their 
absence on Macilentos characteristic. 

As in several aporrhaids (e. g., Latiala) the sculpture 
on the body whorl of Alariinclla not onK- thffers from 
that on the spire, but the back of the bod\' whorl has 
fewer and stronger ribs than the previous whorl, and on 
the apertural face, ribs may be virtually absent. A similar 
pattern of enlarged nodes on the back of the bod\ whorl 
appears to be present in Cah/ptraphonis hinodifcnis and 
in some of the Riinclta-form gastropods (e.g., Rimella). 
Savazzi (1991) suggests that such enlarged nodes assist 
an animal that has landed aperture up to right itself 

Length and deployment of the posterior canal is con- 
sidered a supraspecific characteristic among the Rimclla- 
hke gastropods (Clark and Palmer, 1923). None of the 
specimens of A. vcta has a complete spire, but the pos- 
terior canal appears to end on the juvenile whorls, and 
the canal of a specimen of A. anac, which has probably 
not reached its full length, does not extend to the tip of 
the spire nor does it change direction and extend across 
the back of the spire as in Cahjptraphonis hinodifcnis. 
Alarimclla is sufficiently similar to R/(?u'//«-like gastro- 
pods, except for the expanded outer hp, to suggest that 
Rimella, may be derived from Alarimclla by the reduc- 
tion of the outer hp and lengthening of the posterior 
canal. Similarity of sculpture of Cahjptraphonis hinodi- 
fcnis to that of A. vcta suggests that Cahjptraphonis 
Conrad, 1857, might also deri\e from an A/r;r//(i«7/rt-like 
form b\' reduction of the outer fip, lengthening of the 
posterior canal, and increase of callus deposition over 
the shell. Roy (1996) inchcated that strombids originated 
witliin die apoiThmds iluring Cenomanian-Turonian times, 
which places Alarimclla near the time of dixergence. 

Figures in Korobkox' et ill. (1960, p. 188) of Puffoptcra 
suhrccjiiicni Pchelintsev, 1953 from the Turonian of 
Transcaucasia are suggestive of Alarimclla, but Rostcllaria 
rcijuicniana d'Orbignv, 1842, the tvpe species of Pii<iiop- 
fi'ia PcheliutscN', 1953, is an .\ncluira C^onrad, 1860 (Ko- 
robkoN' et al., 1960, p. 183 ha\e Rostcllaria rcijucmana as 
type species, a misspelhng of R. rcqiiicniana spelled cor- 
rectK' on page 187 in the explanation for Figiue 453). 

Etymology: Ala wing, Latin and Rimella a gastropod. 

Alarimclla vcta (Packard, 1922) 
Figures 53-58; Table 9. 

Aporrli/iis vettis Packaril, 1922. p. 4.31, pi. 36, fisj 1. 

Diagnosis: An Alariiui-lla with the outer lip i-xpanded 
into a broad triangk'. 



L. R. Saul, 1998 



Page 135 




/ 

53 









% 



54 



55 



56 










62 



63 



59 



i 






65 




67 





Figures 53-67. All whitened with anmioniuni chloride. 53-58. Alariinclla veta (Packard), h\pot\pes, 53-54, USNM 494S20 
from USGS loc. 2759, XI, 53, back view, 54, labral side showing wing angle, 55-56, LACMIP 11415 from CIT loc. 978, XI, 55, 
back view, 56, apertnral \iew, 57, LACMIP 11416 from CIT loc. 1067, xl.5, ablabral side, 58, USNM 494821 from USGS loc. 
2759, Xl, apertural \iew showing channel in wing. 59-67. Alarimella aniic n. sp. from CIT loc. 1065, 59-60, paratvpe LACMIP 
11412, 59, X2, back \iew, 60, x'l.5, apertural \iew, 61-62, parat\pe LACMIP 11411, x2, 61, labral side, 62, apertural view, 63, 
65, paratope LACMIP 114L3, K,3, 63, back \iew, 65, ablabral side, 64, paratope LACMIP 11414, x2, apertural \iew, 66-67, 
holot\pe LACMIP 11410, ^2, 66, apertural \ieu, 67, back \iew. 



Description: Shell iiicdiuin .sized, high spired with 
pleural angle of about 26°; whorl profile on spire flativ 
convex becoming angulate on ultimate whorl; suture ap- 
pressed, shallow; bod\ whorl relati\eK' enlarged with a 
short, stniight rostnini; body whorl carina e.xtending onto 
expanded, triangulate outer lip of nearly equal height 
and breadth and having a short posterior cLgitation at its 
distal margin; posterior edge of outer hp extended up- 
ward along spire to form outer, thicked edge of posterior 
canal; inner lip thickened and extended to form inner 
edge of canal along spire; interior of outer lip channeled 
opposite the carina. Sculpture of spire dominated bv 
straight axial ribs, about 16 per whorl, nearK' alligned 
from whorl to whorl, narrower than the interspaces, axial 
ribs evanescing on apertural side of ultimate whorl but 
present on back; spiral sculpture of fine cords on spire 
and about four posterior to carina on ultimate whorl. 



nearly effaced on mid whorl, becoming strtjuger anteri- 
orly, about six on base. 

T\pe specimens: Holotype UCBMP 12298. 

H\'pot>pes: LACMIP 11415 from CIT loc. 978 and 
11416 'from CIT 1067; USNM 494820-494821 from 
USGS loc. 2759, all in the Santa Ana Mountains, Orange 
Ccnmtv, Cahfornia. 

Type locality-: UCBMP loc. 2171, "4 miles [6.44 km] 
SVV of Corona, Corona Sheet. At clav mine, 200 feet [61 
m] up the caiion from a cabin" (Packard, 1922), River- 
side Countv, C;diiomia. 

Dimensions: See Table 9, 

Geologic age: Turonian. 

Geographic distribution: Upper Baker Canyon 



Page 136 



THE NAUTILUS, Vol. Ill, No. 4 



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Member and lower Holz Shale Member of the Ladd 
Formation in the Santa Ana Mountains, Orange and Riv- 
erside Cos., California (UCBMP loe. 2142, 3 spec.; 
UCBMP loc. 2171, 1 spec; USCS loe. 2729, 2 spec; 
CIT loc 1065, 2 spec; LACMIP loc 15295. 2 spec; CIT 
loc 979, 2 spec; LACMIP loc 10898, 2 spec; CIT loc 
978, 4 spec; CIT loc 1067, 1 spec 

Remarks: Packard (1922) onlv mentioned the one 
rather poorly preserved specimen from UCBMP loc. 
2171 on the east side of the Santa Ana Mountiiins, Riv- 
erside County, Cahfornia. He was unsure of the strati- 
graphic position of this outcrop. Packard (1916) included 
a sketch map but cUd not include positions of fossil lo- 
calities on it. Grey's (1961) map co\ering the area rough- 
ly "4 miles [6.44 km] SW of Corona" indicates that part 
of the rocks considered Cretaceous by Packard are now 
mapped as Silverado Formation of Paleocene age. Al- 
though many of the cla\' mines of that area are in strata 
of Paleocene age, and AlarimcUa letci resembles Maci- 
lentos macilcnfus (White, 1889), an earlv to middle Eo- 
cene species (Squires, 1987), no undoubted Paleocene 
specimens with an expanded outer hp have been found. 
Furthermore Grev (1961, p. 22) lists Aporrhai.s cf A. 
vctit.s from his locality' 1 1 in the Baker Canvon Sandstone 
Member on the north side of Tin Mine Canvon, sec. 10, 
T4S, R7W, Corona South quadrangle (Grey,'l961. pi. 1), 
a localitv that is near the southern end of the outcrops 
of Cretaceous age that are roughly 6.44 km SW of Co- 
rona. Specimens of A. veto are uncommon but have 
been collected from several localities in the upper Baker 
Canvon Member and lower Holz Shale Member of the 
Ladd Formation in the vicinity of SiKerado Canyon, Or- 
ange County, Cahfornia. UCBMP loc. 2142, a locahty 
included in Packard's studv, contains several specimens 
of A. veto. 

Specimens of A. veto lacking the adult apertural char- 
acteristics of ha\'ing a posterior canal up the spire resem- 
ble Latiala nodosa (Packard, 1922), but they have stron- 
ger spiral scupltnre than does L nodosa, which also dif- 
fers from the former in having a shghtly more angulate 
whorl profile with somewhat nodose axial ribs. The wing 
of L. nodosa is taller and has a longer, straighter dist;il 
margin with a pronounced anterior projection. 

Ahiriinella vcta resembles illustrations of Coh/pfra- 
plionis binodifcnis of Maastrichtian age from Guerrero, 
southern Mexico, in scuplture and shape, but A. veto 
lacks the callus cowring of C. binodifcnis. and C. hin- 
odijcnis lacks the wing of A. veto. AlarimcUa vcta has 
stronger sculpture, a shorter posterior canal, and a nar- 
rower pleural angle than C. binodifcnis. 

LACMIP 1 1415 was encnisted on the back of the out- 
er lip and up the spire bv calcareous "worm " tubes, and 
ovsters. Tubes are also on the apertural base of the col- 
umella, and the shell was bored by sponges. The pen- 
nltiin;ite whorl has a predepositional hole and the an- 
terior portion of the outer lip has a cuned broken edge, 
both of which suggest tiiat the specimen ma\ ha\e been 
attacked bv a crab. 



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Ahiriuu'lla anac n. sp. 
Fissures 59-67; Table 10. 

Diagnosis: An Alaiiiuclla ha\ing a slender posterior 
spike on the outer lip proximal to the spire, a laterally 
direetetl, liroatl and relatively short wing, and rather 
coarse axial rihs on the spire. 

Description: Shell small, high spired with pleural an- 
gle ot al)out 31°; whorl profile on spire slightly convex 
becoming almost subangulate on ultimate whorl; strong 
randomly distributed varices present on spire; suture im- 
pressed; rostnmi apparently short and straight; outer lip 
expanded into t^vo parts: a posterior short thin spike ad- 
jacent at its base to the spire and a larger, carinate, short 
but broad wing extending laterally; sinus in outer Up 
margin adjacent to the rostrum; inner lip thin. Sculpture 
of spire dominated by rounded arcuate axial ribs extend- 
ing from suture to base, about 16 on penultimate whorl, 
axial ribs disappearing on back of ultimate whorl and 
replaced by fine sigmoid axial fines near outer fip that 
extend onto rostral neck; entire shell surface covered by 
fine, spaced spiral cordlets. 

Type specimens: Holotvpe, LACMIP 11410; para- 
t^pes LACMIP 11411 to 11414 all from CIT loc. 1065. 

Type locality: CIT loc. 1065, about 0.97 km north of 
confluence of Ladd and SiKerado Canyons, Black Star 
Canyon quadrangle, Santa Ana Mts., Orange County, 
Cafifomia. 

Dimensions: See Table 10. 

Age: Late Turonian. 

Geographic distribution: Known onl\- from the type 
locality west of Latld Canyon, Santa Ana Moimtains, Or- 
ange County, Cafifomia, in the Baker Canvon Sandstone 
Member of the Ladd Formation. 

Remarks: AlarimeUa anac may represent early whorls 
of A. veta. The posterior spike adjacent to the spire mav 
be an immature stage of a posterior canal along the 
spire. The largest specimen of A. anac has produced an 
ex-panded outer fip and is about half the size of the most 
complete specimens of A. vcta. Onh- one of the nine 
specimens from CIT loc. 1065 has the wing preserved, 
and it is thin and fragile. The lip ma\- not be fullv mature 
but its presence suggests that the animal would not be 
enlarging its shell significantly. Sohl (1960) considers rel- 
ative size a poor characteristic for specific cfifferentiation 
among aporrhaids, and both A. anac and A. vcta have 
about 16 axial ribs per whorl. The ribs of A. anac are 
somewhat more curved and the spiral cordlets are slight- 
ly stronger than those of A. veta. The fullv mature shape 
of the wing of A anac mav differ from that described 
here. The wing has a slightly cuned carina that is nearer 
to the posterior margin than to the anterior; the anterior 
side of the wing being about twice as tall as the posterior. 
Growth lines on the wing clearly confonn to the present 
blade-like shape and suggest that this species had a short 
but broad, pointed wing. 



Page 138 



THE NAUTILUS, Vol. Ill, No. 4 



Alarimdla anae reseinble.s illiistiatioiis of Gracilialo 
dcceinlirata (Conrad, 1858) of Maastiithtian age from 
the Riplev and Owl Creek Formations of the Gulf Coast 
(Sohl, 1960, pi. 11, fig. 5, 11). Young specimens of A. 
anae before the growth of the wing could easilv be mis- 
taken for immature G. dcccmlirota. which according to 
Sohl (1960) is not t\pical oi Graciliala in that the ante- 
rior border of the outer Up is not digitate. Sohl had no 
specimens of this species with the wing preserxed and 
could not find Conrad's hoIot\pe, but Conrad's illustra- 
tion (1858, pi. 35, fig. 11) shows a wing shght!\- taller 
but similar to that of A. anae and, as in A. anae, the 
anterior edge of the outer hp lacked the digitations typ- 
ical of Graeiliala calcaris (Wade, 1926) and other species 
ot Graciliala. 

Alarimdla anae differs from Anchura (H.) tricosa Saul 
and Popenoe, 1993, which has a somewhat similar pos- 
terior spike, in having fine spiral striae rather than about 
5 spiral cords on the spire whorls, fewer a\ia] ribs, and 
a less angulate whorl profile. The wing <jf Alarimdla 
anae is much shorter, less cur\'ed, and is without sec- 
ondare spurs on the shank. On the ultimate whorl of A. 
anae, the a-\ial scidpture is nearl\- effaced, and an axial 
bulge is present on the abapertural side of the apertural 
face. 

Et\Tnologj': The specific epithet, anae, refers to the 
name of the Santa Ana Mountains, Cahfomia. 



ACKNOWLEDGEMENTS 

I am very grateful to John Tavlor, Department of Zool- 
og)'. The Natural History Museum, for a photocopy of 
Tate (1865). Without it the Kpe species for Arr/iogra and 
Perissoptera could not be confirmed. Lindsev Groves, 
Natural Histon Museum of Los Angeles Countv, was 
extremely helpful in finding several other elusive pubU- 
cations. Marc Florence, Smithsonian Institution, and 
Elana Benamv, Academ\' of Natural Sciences of Phila- 
delphia, responded e.xpeditiouslv to specimen number 
questions. I thank the reviewers Richard Squires and 
William Elder and the editor Jose H. Leal for helping 
to make this paper much more readable. 

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L. R. Saul, 1998 



Page 139 



Jones. D L . W \: Sliter and \\'. P Popenoe 197S Mid-Cre- 
taceous (Albian to Turonian) liiostratigraphv ot nortlieni 
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LOCALITIES CITED 

CIT and UCLA localities have been given LACMIP 
numbers. The CIT and UCLA numbers for Redding 
area and Santa Ana Mountains localities are included 
here because thev have been plotted on published maps. 
Most of the CIT and UCLA localities of Turonian age 
in the Redding area were plotted on Jones et al. (1978: 
fig. 5). Most of the CIT localities of the northern Santa 
Ana Mountains were plotted on Popenoe (1942: fig. 2); 
these and UCLA localities were plotted on Saul and Bo- 
ttjer (1982: maps 1-3). 

80 CIT = LACMIP 8194: In sandstone above cgl., 

at fork of Silverado and Ladd Canyons on N 
side of Silverado Canyon, [NW 1/4, SW 1/4 
sec.8, T5S, R7W, El Toro quad.], Santa Ana 
Mts., Orange County', California. Coll: B. N. 
Moore, 1926. Ladd Formation, Baker Canyon 
Sandstone. Turonian. 

82 CIT = LACMIP 8195: Lnney sandstone bed 

near base of shale, S of roadcut at Holz's Ranch 
(locality may become obscured bv slides), Sil- 
verado Canvon [E edge SE 1/4, SE 1/4, sec. 7, 
T.5S, R7\\', El Toro quad.], Santa Ana Mts., Or- 
ange County, C;ilifornia. Coll: B. N. Moore, 
1927. Ladd Formation, Holz-Baker transition. 
Tiuonian. 

99 CIT = LACMIP 8180: Concretions in shale just 

above sandstone on south side of Silverado Can- 
yon coming in from south about 1 1/2 miles E 
of Ladd Canyon [appro.x. 0,12 km N, 0.07 km 
E of SW cor. sec.9, T5S, R7W, Santiago Peak 
quad.], Santa Ana Mtns., Orange County, Cali- 
fornia. Coll: B. N. Moore, 1928. Ladd Forma- 
tion, Holz-Baker transition. Turonian. 

454 CIT = LACMIP 10873: Cretaceous shale, fi- 
reline about Hough's 80 on S side of stream, 
about 400' above creek, Silverado Canyon 
]about 450'S, 875'E of NW cor. sec. 16, fsS, 
R'AV, Santiago Peak (juad.], Santa Ana Mts., Or- 
ange County, (-'alifornia. Coll: B. N. Moore, 
1929. Ladd Formation, lower Holz Shale. Late 
Turonian. 

978 CIT = LACMIP 10884: NE slope, and mar 
crest of bluff overlooking Santiago Canvon Jat 
about NE cor. sec.33, T5S, RTU''), approx. 1 1/ 



2 mi. SE of the dam just above the mouth of 
Harding Canyon, Santiago Peak quad,, Santa 
Ana Mts., Orange County, California. Coll: W. 
P Popenoe, 4/14/1933. Ladd Formation, Baker 
Canyon Sandstone about 100' aboxe gray basal 
cgl. Turonian. 

979 CIT = LACMIP 10885: 10' below 978, NE 
slope, and near crest of bluff overlooking San- 
tiago Canyon [at about NE cor. sec.33, T5S, 
R7W, Santiago Peak quad], Santa Ana Mts., Or- 
ange County, California. Coll: W P. Popenoe, 4/ 
14/1933. Ladd Formation, Baker Canyon Sand- 
stone, about 90' above gray basal cgl. Turonian. 

1058 CIT = LACMIP 10890: North side of Silverado 
Canyon Road, about 15' above rd., and 300'N 
72°E of Holz Ranch house, Silverado Canyon 
[1390'N, 210'E of SW cor. sec.8, T5S, R7W, El 
Toro quad], Santa Ana Mts., Orange County, 
California. Coll: W. P. Popenoe and others, 4/1/ 

1933. Basal Baker Canyon Sandstone about 10' 
above top of conglomerate, Ladd Formation. 
Turonian. 

1064 CIT = L.\CMIP 10893: Area S of Harding 
Canyon, 'Vulture Crags, just N of first large can- 
yon cutting across Cretaceous beds S of Har- 
ding Canyon, about 2 mi. S42°E of the dam in 
Hartling Canyon and 6800' N5.5°E of juncture 
of Santiago Creek and Traliuco Canyon Rds. 
[2600'N, 1625'E of SW cor. sec.34, T5S, R7W, 
Santiago Peak quad], Santa Ana Mts., Orange 
County, Cahfornia. Coll: W. R Popenoe, 10/14/ 

1934. Lower Holz Shale, Ladd Formation. Tti- 
ronian. 

1065 CIT = LACMIP 10891: Ss overlying basal Up- 
per Cretaceous cgl., from crest of scaip on W 
side of Ladd Canvon, about 0.6 mi. N of junc- 
ture of Ladd and Silverado Can\ons [1300'S, 
300'E of NW cor. sec.8, T5S, R7\V, Black Star 
Canvon quad.], Santa Ana Mts., Orange County, 
Califbrnia. Coll: W. R Popenoe, 3/3/1933. Ladd 
Formation, Baker Canyon Sandstone. Turonian 

1067 CIT: = LACMIP 10883: Immediately above 
base of gray ss overl\ing grav basal cgl. [1800'N, 
600'E of SW cor. .sec.2i, f5S, R7\V, Santiago 
Peak (juad], Santa Ana Mts., Orange County, 
California. Coll: W. P Popenoe, April 21, 1932. 
Ladd Formation, Baker Canyon Sandstone 
Member. Tiuonian. 

1164 CIT = LACMIP 10079: S side Silverado Can- 
yon near mouth of small N-flo\ving gull\-, and at 
top of lower fossififerous sandstone series, about 
at top of lower fossiliferous sandstone series, 
about 400 feet ( 120 m) SE of Holz Ranch house 
in SE cor .sec.7, T5S. R7W [1025'N, 150'E of 
SW cor. sec.8, T5S, R7W, El Toro quad.], Santa 
Ana Mts., Orange County, California. Coll: W. 
P. Popenoe, May 15, 19.'35. Lackl Formation, Ba- 
ker Canyon Sandstone Member. Turonian. 

1212 CIT = LACMIP 10735: Little Cow Creek, ap- 



L. R. Saul, 199S 



Page 141 



prox. 2 mile {3.2 km) NE of Fiazier's Corners, 
hard sandy concretions in shale, banks of gullies 
in pasture [about 2500'N, 750'W of SE cor. sec. 
4, T32N, R3W, Millville quad.], Shasta County, 
California. Coll: W. P. Popenoe, 1933. Redding 
Formation, Frazier Silt. Turonian. 
1221 CIT = LACMIP 107.38: about 1/4 mi. N of Al- 
turas-Redding hwy U.S. 299 and 1.6 mi. In rd 
NE of Frazier's Comers, S facing shale bank 
along Jim Creek [250'S, 700'W of NE cor sec.S, 
T32N, R3W, MilKille quad.], Shasta Count)', 
Cahfornia. Coll: Popenoe and Alilroth, June 29, 
1936. Redding Formation, upper Frazier Silt 
Member Turonian. 

1264 CIT = l^CMIP 10750: Massive, browai sand- 
stone cropping out in bed of small gullv tribu- 
tary to Little Cow Creek [appro.x. 1850'S, 
2250'E of NW cor. sec.9, T32N, R3W, Millville 
quad.], Shasta Counts', Cahtornia. Coll: W. P. 
Popenoe, April 12, 1937. Redding Formation, 
basal Melton Sandstone Member. Turonian 

1265 CIT = LACMIP 10751: Right bank of Litde 
Cow Creek, about 50' from creek [0.58 km S, 
0.48 km \y of NE cor sec.9, T32N, R3\V, Mill- 
ville quad.], Shasta Countv, California. Coll; W. 
P. Popenoe, 4/15/37. Redding Formation, Mel- 
ton Sandstone Member Turonian. 

1266 CIT = LACMIP 10752: Right bank of Little 
Cow Creek about 500 ft. dowaistream from CIT 
1265 [center of sec.9, T32N. R3W, Millville 
quad.,] Shasta Countv, California. CoU: W. P. 
Popenoe, April 15, 1937. Redchng Formation, 
Melton Sandstone Member Late Turonian. 

1290 CIT = LACMIP 10135: S side of prominent hill 
W of Mustang Spring, about 1500' nearly due 
N of Holz Ranch house, north side Santiago 
Canyon, sec. 7, T5S, R7\\', [Black Star Canvon 
quad.], Santa Ana Mountains, Orange County, 
California. Coll: Popenoe, Wells, Church, Hen- 
shaw, and Fiedler, April 6, 1937. Ladd Forma- 
tion, near top Baker Canyon Sandstone Mem- 
ber. Late Turonian. 

1307 CIT = LACMIP S169: About 25' E of CIT 
1290 and not more than 10' stratigraphicalK' be- 
low CIT 1290, W of Mustang Spring, north side 
Santiago Canvon, sec. 7, T5S, R7\\', [Black Star 
Canyon quad], Santa Ana Mountains, Orange 
County, Cahfornia. Coll: VV. P. Popenoe, April 
24, 1937, Ladd Formation, Baker Canvon Sand- 
stone Member Late Turonian. 

1346 CIT = UCLA 5421: LACMIP 107.54: Sand- 
stone nodules in shale, left bank of Little Cow 
Creek, about 70 ni NE (upstream) from inter- 
section of creek bed with S line of sec. 9 
[ISOO'N, 2200'E of SE cor ,sec.9, T32N, R3W, 
MilKille quad.], 10 mi. NE of Redding, Shasta 
County, California. CoU: W. P. Popenoe and 
Jane Hoel, 7/8/1937. Redding Formation, Mel- 
ton Sandstone Member. Turonian. 



1532 CIT = LACMIP 10815: Concretions in shale 
near right bank Salt Creek [approx. 600'N, 
700"W of SE cor. sec.4, T32N, R3W, Millville 
quad.], Shasta County California. Coll: Popenoe 
and Alilroth, Julv 10, 1936, Red(hng Formation, 
near top Frazier Silt Member Turonian 

2142 UCBMP: 2 1/4 miles NNW of B.M. 1271, Co- 
rona sheet. Left side of Silverado Canon, in 
shales undercut bv the stream about 200 ft, 
southeast of the road. Ladd Formation, lower 
IIolz Shale Member. Turonian 

2171 UCBMP: 4 miles SW of Corona, Corona Sheet 
[Corona South quad.]. At clay mine, 200 feet up 
the canon from a cabin. Riverside Countv, Cal- 
ifornia. Ladd Formation. Turonian. 

2325 UCLA = LACMIP 22325: Small gully entering 
Silverado Canvon from S, just W rjf the narrows, 
about 9000' upstream from the mouth of the 
canyon, and directly S from farmhouse on Holz 
Ranch, about 1025'N, 150'E of SW cor. sec.S, 
T5S, R7W, El Toro quad., Santa Ana Mts., Or- 
ange County', Cahfornia. Coll: W P. Popenoe, 
1946. Ladd Formation, top of Baker Canyon 
Member Turonian. 

2759 USGS: Near Silverado Canvon, in lower part of 
Ladd Canvon, Santa Ana Mountains Orange 
Counts', California, Coll: S. Bowers, April 24, 
1903. Ladd Formation, Baker Canvon Sand- 
stone Member Turonian. 

3465 UCLA = LACMIP 2.3465: Fossils in small kniob 
of sand)' cgl., on North Fork of Cottonwood 
Creek, about 1500'SSW of UCLA 3464, 0.38 
mi. N, 0.3 mi. E of SW cor sec. 16, T30N, R6W, 
Ono quad., Shasta Counts', California. Coll: P. 
U. Rodda and M. A. Murjihy, May 1955. Bud- 
den Canyon Formation, Bald Hills Member 
Late Cenomanian. 

4365 UCLA = LACMIP 24365: In fine grained ss 
with Roinaiiiccras, left bank of French Creek 
(north end Swede Basin), approx. 500'N and W 
of SE cor sec.5, T32N, R2W, Millville fjuad., 
Shasta Counts', California. Coll: W. P. Popenoe, 
8/25/1957, Redding Formation, basal bed of 
PFrazier Siltstone, "Turonian. 

4658 UCLA: Concretions at top of Frazier Siltstone, 
right bank of Salt Creek, about lOOO'N, 400'W 
of SE cor. sec.4, T32N, R3W, Millville (juad., 
Shasta County, California. Coll: W P. Popenoe, 
Aug. 1961. Redding Formation, Frazier Silt- 
stone Member, Turonian. 

5421 UCLA = LACMIP 25421: Sandstone nodules 
in shale, left bank of Little Cow Creek, about 
5' above the channel bottom, about 70 m NE 
(upstream) from intersection of creek bed with 
hue fence, and about 1/4 mi. downstream from 
old Walter Melton farmhouse [S hne sec.9, 
T32N, R.3W, Millville quad], Shasta County, 
California. Coll: W P. Popenoe, summer, 1937. 



Page 142 



THE NAUTILUS, Vol. 111. No. 4 



Rfildinii Foniiatioii, Melton Sandstone. Tnro- 
nian. 

10S76 LACMIP = CIT 1042: Limey len.se.s in ss crop- 
ping out on N hank of Rancheria (Juleh, about lfSS38 
1.5 ini. W of Henlev [approx. 21()S, SOOE of 
NW cor. sec.3(), T46N. HHW. Hornhrook quad.]. 
Sisldvou Countv, (Jaliiornia. Coll: Popenoe and 
Findlay, Sept. 8, 1933. Hornbrook Fm., Osbur- 
ger Gulch Sandstone Member Tnronian. 

10895 LaCMIP: North .side of Harding Canyon, north 24370 

of Modjeska Reservoir, at elev. 1705', 1775'N, 
1075E'of SW cor. sec.21, T5S, R7\V, Santiago 
Peak quad., Santa Ana Mts., Orange County, 
California. Coll: L. J. Czel, spiing 1958. Ladd 
F"ormation, Baker Can\'on Santlstone Member. 
Turonian. 

10898 LACMIP: North side of Harding Canyon, north 
of Modjeska Resenoir, at elev. 1750', 950'N, 
60()'E of SW cor. sec.21, T5S, R7\V, Santiago 24R70 

Peak quad., Santa .Ana Mts., Orange Coimt\', 
California. C(j11: L ). Czel, spring 1958. Baker 
Canyon Sandstone, Ladd Formation. Turonian. 

10901 LACMIP = CIT 1044: Right bank of Ranchena 
Gulch just over hill north from Scholz Ranch 
house, 1.5 mi. W of Henlev, 535'S, 1435'W of 25422 

NE cor. sec.30, T47N, R6W. Yreka quad., Sis- 
kivou County California. Coll: W. P. Popenoe 
and W. A. Findlay, Sept. 1933. Hornbrook Fm., 
Osburger Gulch Sandstone Member. Turonian. 

10953 LACMIP: Holz Ranch, north side of Silverado 

Canyon, on ridge top west of Laild Canyon at 28788 

approx. 1510' elev, approx. 197' above base of 
upper Baker Canyon Sandstone, 2550'N, 520'E 
of SW cor. sec.8, T5S,. R7W, Black Star Canyon 
(juad., Santa Ana Mts., Orange Countv, Califor- 
nia. Coll: R. G. Cassi, spring 1958. Ladd For- 
mation, upper Baker Canyon Member. Turoiii- 29181 
an. 

15295 LACMIP: South side of Silverado Canvon near 
mouth of small N-Howing gulK', about 400' SE 
of Holz Ranch house. 1025N, 'l50'E of SW cor. 
sec. 8, T5S, R7W, El Toro (juad., Santa Ana 



Mts., Orange Countw California. C>'oll: Robert 
Drachuk, 1979. Top of Baker Camon Sandstone 
Member, Ladd Formation. Turonian. 
LACMIP: Float— North Fork of Cottonwood 
Creek, 3/4 mi. downstream from month of Hul- 
ing Creek, Sec.21, T30N, R6W, Ono (jnad.. 
Shasta County, Cahfornia. Coll and date un- 
known. Budden Canvon formation. Bald Hills 
Member. Cenomanian. 

LACMIP (=CIT 1020; UCLA 4370): Sandstone 
boulder in heavv conglomerate, channel of El- 
der Creek, about 0.8 mi. N and E of SW cor. 
sec.l4, T25N, R6W, at east edge of Colyear 
Springs quad., Tehama Countw C^alilornia. Coll: 
W P. Popenoe and W. A. Findla\-, August 18, 
1933. Great 'Valley Series. Bndden Canvon For- 
mation. C^enomanian or late /\lbian boulder in 
Turonian santlstone. 

LACMIP = UCLA 4670: Sandstone cropping 
out in bed and on west bank of Grave Creek, 
W of road, 2400'N of SW cor. sec.5, T34S, R4W, 
Wimer 1954 15' <|uad., Jackson Connt\, Oregon. 
Coll: W. P Popenoe, Aug. 20, 1960. Late earlv- 
micklle Albian. 

LACMIP = UCLA 5422: Rancheria Gulch, 
about 1 mi. W of Henlev, and approx. 400'N, 
200()'W of SE cor. sec. 19, T47N, R6W, Yreka 
30' (juad. (1939), Siskivou Counts, C'alifornia. 
Coll: W. P. Popenoe, summer 1951. Hornbrook 
Formation, Osburger Gulch Member Turonian. 
LACMIP: ( = USGS loc. 14528) boulder m cgl, 
NW end of McLure Valle\- = Sunflower Valley, 
elev. 1420', 125()'N, 775'W of SE cor. sec.26, 
T23S, R17E, Reef Ridge quad.. Kings County, 
Cahfornia. Goll: Ralph Stewart. Panoche For- 
mation. Cenomanian. 

LACMIP: Hill north of Holz Ranch house at 
about 1500' elev, sec. 8, T5S, R7W, Black Star 
Canvon quad., Santa Ana Mts., Orange Count); 
California. Coll: W P. Popenoe. Ladd Forma- 
tion, Baker Canyon Sandstone Member. Turo- 
nian. 



THE NAUTILUS 111(4):143-14S, 1998 



Pasze 143 



Salinity Tolerance of the Freshwater Bivalve Dreissena 
polxjmoiyha (Pallas, 1771) (Bivalvia, Dreissenidae) 



Sha^Ti J. Wilcox 
Thomas H. Dietz' 

Department ot Biological Sciences 
Louisiana State Universit\' 
Baton Rouge. LA TOSO.IUSA 



ABSTRACT 

Tlie effects of ele\ated ion concentrations in the bathing me- 
dium on osmoregulation and mortalit\" rates were examined in 
the freshwater bi\al\e Dreissena polijinivpha. Animals accli- 
mated to artificial pondwater (APW) tolerated an acute transfer 
to approximately 100 mosniol kg ' artificial seawater {3.5^cc 
ASW), for months, with \irtuallv no mortalitv. However, there 
was substantial mortalits'. within da\s, when the animals were 
acuteh transferred to dilute ASW greater than 150 mosmol 
kg ' (5%f). Dreissena pc^li/nnotyha rapidlv ( — 12 h) became 
isosmotic to the bathmg niediiun when exposed to 100 mosmol 
kg'' ASW, although it took 3 d ol acclimation before the ion 
concentrations in the blood stabilized. Salinit)' tolerance was 
increased when the animals were acclimated for 3 d to a salin- 
itv of 54 mosmol kg'' ASW followed by 54 mosmol kg'' sahn- 
itv increases everv third day. Although some mussels acclimat- 
ed for 2 weeks to 377 mosmol kg' ASW could sur\i\e direct 
transfer to APW, sur\ival was impro\ed when the acute reduc- 
tion in salinitx' was less than 260 mosmol kg" ' Dreissena poli/- 
morplia has the capacity' to tolerate oligohaline water with 
small fluctuations in salinitv. 

Key words: zebra nmssel. estuarv, salinitv tolerance, fresh-wa- 
ter bivaKe, ion regulation, osmoretrulation. 



INTRODUCTION 

Dreissena pdlyiiun-pha re(juire minimal concentrations 
of Mg-\ Na^, Cl and K" in the bathing medium to 
ionoregiilate (Nichols, 1993; Ram & Walker, 1993; Dietz 
ct ah, 1994). Dreissena poh/morjjlui cannot survive a 
week when 45 mmol 1 ' NaCl is added to artificial pond- 
water (APW), or survive one dav it the K' concentration 
is increased to 1 mmol 1' (Horohox- et ah. 1992: Fisher 
et a]., 1991). However, these animals will survive in di- 
lute artificial seawater (Kilgour ci ah, 1994; Wright et ai, 
1996; Dietz ct aL. 1997), and ha\e been obserxed in 
brackish water and in estuaries where ambient salinities 
range from freshwater to greater than 12%o (Aladin & 
Potts. 1992; Khmowicz. 1958; Straver & Smith, 1993). 



' Author for correspondence. 
E-mail: zothom@lsu\m. sncc.lsu.edu 



The effects of environmental perturbation (tempera- 
ture, salinitv) on the physiological processes associated 
xxith ion regulation and salinitx' tolerance hax'e been 
stuched recently (Dietz ct a]., 1994; Kilgour et al. 1994; 
Scheide & Bonaminio, 1994; Wright ct al, 1996). Some 
reports suggest that the salinitv tolerance ot D. pohj- 
morpha from North American populations is influenced 
bv multiple factors including temperature, hie stage and 
the salinitv acchmation regime (Kilgour ct ah, 1994; 
Wright ct al, 1996). Recently, we have observed that 
mussels e.xposed to elevated Na^ and K* ( — 10% sea- 
water) will redistribute the ions between the intracellu- 
lar (ICF) and extracellular fluid (ECF) compartments 
(Dietz et al, 1997). When both Na' and K' are present 
in the media, the mussels maintained a normal ICF/ 
ECF K* ratio. In addition, the epithefial cells in the giU 
were able to partially recover cellular volume when ex- 
posed to a hxperosmotic bath cont;uning 45 mmol 1' 
Na* and 1 mmol 1 ' K*, but could not do so when K* 
was absent (Dietz ct al. 1998). 

In this study we report the effects of osmotic and ionic 
challenges on body fluid ion concentrations and siu"vival 
in D pohjmotyha. We identify the environmental safin- 
itv hmits and changes that are compatible with osmo- 
regulation and survival. 

MATERIALS AND METHODS 

Ajiimals: Specimens of Dreissena poli/moifiha (zebra 
mussels) were coOected from Lake Erie at the mouth of 
the Raisin River in Michigan and from the Mississippi 
River near Baton Rouge, Louisiana. Mussels were stored 
unfed in aquaria containing aerated artificial pondwater 
(APW in mmol 1 '; 0.5 NaCl, 0.4 CaCL, 0.2 MgSO,, 0.2 
NaHCO,, 0.05 KCl; Dietz ct al, 1994). Some of the 
animals were kept at 13°C and were step-wise acclimat- 
ed to 16°C and 22 ± 2°C for >5 d at each temperature 
before use. Only large animals (1.5-3 cm shell length) 
attached to the contiuner or to another mussel bv bvssal 
threads were used. Aquarium water vx'as replaced every 
2-3 d and waste water and containers were treated with 
1% chlorine bleach. 



Page 144 



THE NAUTILUS, Vol. Ill, No. 4 



Solute analyses: Blood samples (150-200 |xl) were 
collected by pericardial puncture from individual mus- 
sels and centrifuged at 15,000 g min before analysis 
(F\'bn & Costlovv, 1975). The osmolalit\' of the blood was 
determined by freezing point depression. Sodium and 
potassium concentrations were determined by flame 
emission photometry on diluted samples. Calcium and 
magnesium samples were diluted with LaCl/HCl and 
assayed using an atomic absorption spectrophotometer. 
Chloride was determined by electrometric titration. The 
difference between the total solute of the blood and the 
sum of the measured ion concentrations was identified 
as "other" and is mostly bicarbonate in APW acclimated 
animals (Byrne fie Dietz, 1997), but sulfate is a significant 
component in ASW (Potts, 1954; Withers, 1992). Net 
ion Rux (J„) was determined by measuring the change in 
bathing fluid ion concentration at specific intervals. The 
flu.x was e.xpressed as (xmol g ' dry tissue h '. 

Elevated bath solute concentrations: Some mussels 
were either acutely exposed or step-wise acclimated to 
a modified artificial seawater (ASW) diluted with APW 
(ASW in mmol 1'; 449.1 NaCl, 27.5 MgSO,, 24.4 
MgCl,, 9.9 CaCL, 6.6 KCl, 2.4 KHCO>, O.S KBr, 0.4 
H3BO3; 1076 mosmol kg ' total solute concentration; 
35%c; Chambers & De Armendi, 1979). No buffer was 
added to the seawater stock and the pH in diluted sam- 
ples varied from 7-8. The artificial seawater was diluted 
with APW to approximate natural mixtures of freshwater 
and seawater. 

We determined the effects of an acute transfer to el- 
evated salinity on the blood ion composition and mor- 
tality in mussels. Animals were transferred tlirectly from 
APW to a bath of diluted ASW that was changed daily. 
We calculated (see below) the time to 50% mortahtv- 
(LT^ii) and recorded the time of death of the last animal 
as 100% sample mortality- (SM,,,,,). The animals used for 
chronic exposure to a constant bath salinitv' concentra- 
tion were exposed to 54 mosmol kg ' ASW for 3 d fol- 
lf)wed by subsequent increases of 54 mosmol kg ' at 3 
d intervals until the specified bath solute concentration 
was attained, and the safinity held constant thereafter. 

Animals were exposed to cyclic changes in salinit\' by 
adding 9 liters of 161 mosmol kg ' ASW from a Mariott 
bottle, over 12 h, to one liter of APW resulting in bath 
concentration of 130 mosmol kg ' ASW. After 12 h all 
but one fiter of the bath was discarded and nine liters 
of APW was added over 12 h to return the salinity to 
about 14 mosmol kg ' After 6 d of the salinity cycles, 
samples of bathing medium and blood from 5 animals 
were collected at 4 h intervals for 24 h. 

Mortality studies: We determined the a!)ility of D 
■pohjmotyha to tolerate acute transfer to dilutions of 
ASW by noting their survival time and recording behav- 
ioral changes. Animals that gapeil widely, were unre- 
sponsive to stimulation by touching the valves or tissue 
near the siphons, and (Ld not hold the v;ilve closed when 
mechanically closed with forceps, were considered dead 
and the time noted. Some of the animals acclimated to 



APW, 108 
9 H- 




20 40 



I ' I 
80 100 120 140 160 



60 
Time (hours) 



Figure 1. Percent survival of APW-accIiinated D pohjmor- 
pha acutely transferred to artificial seawater diluted with APW 
(N = 30 for each group). The number adjacent each line rep- 
resents the total solute, in mosmol kg ' measured in the bath- 
ing medium. 

hyperosmotic solutions and acutek transferred to hvpos- 
motic solutions appeared to swell and were unable to 
completely close the valves, but we were able to deter- 
mine live or dead animals refiably bv judging motor ac- 
tivities. The time for 50%' sample mortality (LT,,,) was 
calculated from a probit model (Finney, 1971), using a 
linear regression of the logarithm of time and cumulative 
mortahty: probit (% dead) = A -I- B (log time, h or d). 

Statistical analyses: Data are expressed as mean ± 
s.e.m. Data were tested for homogeneity of variance and 
some of these data were log transformed before one- 
way analyses of variance (ANOVA) were performed at 
an alpha level of 0.05. A treatment that had a significant 
ANOVA was examined for similarities and differences 
using the Tukey Studentized range procedure at an al- 
pha level of 0.05 (e(jual sample sizes), or with the Tukev- 
Kramer modification when sample sizes were unequal 
(SuperANOVA, Abacus Concepts Inc., Berkeley, Cali- 
fornia). 

RESULTS 

Acute transfer to dilute artificial seawater: APW- 

acchmated D p(>h/nu>if>ha transferred directly into ch- 
lute ASW with solute concentrations greater than 151 
mosmol kg ' (5%c) suffered substantial mortality be- 
tween 5 and 21 h (Fig. 1). The LT,„ was —31 h for 
anim;ds transferred to 215 (7%ci) and about 36 h for an- 
imals in 194 (6.3%c) mosmol kg ' .•KSW. Survival time 
increased (LT,,, 74 h) tor the animals transferred to 172 
mosmol kg ' (5.6%o) ASW. The experiment was tenni- 
nated after 2 weeks and LT,,, values were not deter- 
mined for the four groups in salinities <151 mosmol 
kg ' as all or the majority of the animals survived. Mus- 
sels acchmated <108 mosmol kg ' ASW survived for 
months (LT^„ 3^ mo), unfed, as did the APW-accfimat- 
ed control animals (LT-„, 4—5 mo). A majoritv of the an- 



S. J. Wil 


co.x 


and T. H. Dietz, 1998 






Page 145 


Table 1. 


Ch 


mges in Drei.ssenti poli/morj)lict blood ion coneenti 


iitions tolldwinij acute transfer to 108 


mosinol kt; 


ASW (3.5%p). 




N 


niosniol kti ' 


Concentration (mniol 1 ' 


) 




(h) 


*Total solute Other Na* 


CI K 


Ca^- 


*Mg-- 




12 
24 

72 


5 
7 
6 
6 


49 ± 3' 11.0 ± 1.7' 18.5 ± 1.1' 
85 ± 2' 4.2 ± 1.7'' 38.8 ± 1.1'' 
97 ± 1'' 10.1 ± 1.7' 40.5 ± 0.7''' 
102 ± 1' 13.2 ± 1.4' 42.4 ± 0.9' 


14.4 ± 0.4' 0.6 ± 0.1' 
36.4 ± 0.8''' 0.8 ± 0.0' 
34.1 ± 0.5'' 1.2 ± 0.1'' 
37.3 ± 0.6' 1.6 ± 0.1' 


4.0 ± 0.1' 
2.3 ± 0.2'' 
2.0 ± 0.1'' 
2.5 ± 0.1'' 


0.5 ± 0.0' 
2.9 ± 0.1'' 
5.5 ± 0.1' 
5.1 ± 0.1' 



Mean ± sem. "Other" was calculated from the thfference between the total solute and measured ion concentrations Means within 
a column with different letters are significantK different b\ the Tuke\-Kramer Studentized range procedure, P < 0.05; {*) log,, 
transformed data for statistical analyses. 



imals in <108 mosmol kg ' ASW formed bvssal thread 
attachments within 48 h of the transfer. 

Changes in blood solute concentration: Drcissciw 
pohjmoi-pha transferred trom APW to 108 mosmol kg" ' 
ASW became isosmotic ■with the bathing medium be- 
tween 12 and 24 h (Talile 1). Sochum concentration in 
the lilood rapidlv tended to become isoionic, Init Cl 
remained hvpoionic to the medium. Although the ASW 
bath contained elevated Ca-" and Mg-* compared to 
APW, the blood concentration of Ca-* decreased 40- 
50%, but the Mg-^ rose 10-fold and became isoionic 
■with the bath. 

Exposure to elevated artificial seawater concentra- 
tions: Dreisscna poli/inorpha suffered considerable 
mortahty when acutely transferred to salinities approach- 
ing 15% ASW but tolerated more dilute solutions. 
Therefore, we examined the abilits- of mussels to sunise 
a step-v\ise increase in salinits' then subsequenth' held at 
a constant salinity. Several hundred mussels ■were trans- 
ferred to 54 mosmol kg ' (1.5%c) for 3 d and the salinifs' 
was increased bv 54 mosmol kg ' at 3 day intervals. At 
several salinities ranging from 269 to 486 mosmol kg ' 
groups of 30 mussels were transferred to separate con- 
tainers. The water was changed daily, but the salinity was 
held constant thereafter and the survival was monitored 
(Table 2). The LT-^,, for the mussels was inverseh- related 
to the concentration of the final bathing solution. How- 
ever, the total length of time mussels tolerated both the 



Table 2. Survival (LT-„) of Drcissena poh/inoqihii chromcall 
e\']:)osetl to tliintions ot artificial seawater 



Final bath solute 










concentration 


Pre 


acclimation 


time 




(mosmol kg ') 




(days) 




Davs LT,„ 


269 




12 




30.5 


325 




15 




21.3 


377 




18 




14.7 


486 




24 




5.4 



The pre-acclimation time was the total number of days the 
animals were in step-wise increasing salinitv (54 mosmol kg ' 
at 3 day intervals) until the specified final bath concentration 
was established. The LT-,, was determined on 30 animals at 
each salinitv. 



step-wise sahnitv adjustment and constant sahnih' accli- 
mation to reach the LT^,, ranged from 29 to 47 cl, with 
no consistent pattern. We typically observed greater than 
97% survival up to 269 mosmol kg"' (8.8%o) in large 
groups of mussels (~300) during the step-wise adjust- 
ment in salinitv. These data have some bias since the 
most tolerant animals survived the step-wise sahnity 
change and were used to determine the LT-,,,. In three 
separate studies, >70% of the mussels survived the step- 
wise increase in salinitA' up to about 380 mosmol kg ' 

Diurnal salinity- changes: Because the salinity of oli- 
gohaline water in coastal environments tends to change 
in a cvchc manner, we examined the ability of D. poh/- 
inorpha to tolerate cvclic changes in salinitv. We exposed 
the mussels to diurnal changes in salinitv ranging from 
14 to 130 mosmol kg'' for over a week and all of the 
animals survived and formed byssal attachments. The 
blood ion composition of the mussels paralleled the 
bathing medium during both the dilution and increasing 
salinih' phases of the cycle (Fig. 2). The blood ion com- 
position and concentrations lagged approximately 2-A h 
behind the changes in the ion composition of the bath. 
The mussel blood became isosmotic to the bathing me- 
dium during the high salinitv phase of the c\cle. During 
the initial 4 h of the dilution phase the total solute in 
the blood decreased rapidly, but the mussels osmore- 
gulated in the lower salinities. The concentrations of the 
measured ions also decreased as the bath became chlute, 
but the concentration of unidentified osmoKtes (other) 
was elevated. The control anim;ils held in constant chlute 
ASW or APW maintained constant blood ion composi- 
tion during the 8 d. In a similar studv, we extended the 
tidal cvcle for lid and all ot the animals sui'Aix'ed (data 
not shown). Most of the animals attached by byssal 
threads within 48 h in both experiments. 

Acute transfer from h\perosmotic to hyposmotic 
media: We anah'zed the blood ion concentration oi 
animals that survived 14 d acchmation to constant 377 
mosmol kg ' seawater and for 24 h after being re- 
turned to APW (Table 3). Animals that were acutely 
transferred to APW experienced a 79% reduction of the 
blood total solutes within 6 h (Table 3). The restoration 
of blood ion composition was not completed within 24 
h as onlv the concentration of Ca-\ K\ and total solute 



Page 146 



THE NAUTILUS, Vol. Ill, No. 4 



150-, Bath total solute, mosmol kg-1 



o 

E 
E 

c 
o 



c 
dJ 
o 
c 
o 

o 



loo- 




se - 



T 1 1 1 T" 

O O O O O 

o o o o o 

00 CM CD O O 
O T- T- CM O 



O O 

O O 

•>a- CO 

o o 



Time, h 



Figure 2. The changes in blood ion composition ot D poh/- 
niorpha over a 24 h salinitv cycle ransjing from 14 to 130 mos- 
mol kg"' solute (N = 5). The animals were e.xposed to the 
dumial changes in salinitA' for 6 d and blood samples collected 
dunng dav 7. The individual ion concentrations in the blood 
were added to form the stacked histogram. The filled circles 
represent the total solute concentration in the bath at the re- 
spective blood sampling times. 



were similar to APW control animals. The concentra- 
tions of Na* and Cl" were significantly less than con- 
trols, but Mg-* and the unmeasured solutes (other) re- 
mained elevated. It was likelv that the decline in solute 
concentrations was due to both the osmotic uptake of 
water as well as a loss of ions to the en\ironment, but 
these variables were not measured during this studv (see 
Dietz et at, 1997). In a separate study we measured net 
ion losses when mussels were direct]\' transferred from 
106 mosmol kg ' ASW to APW. The animals lost Na', 



Cl , and K' ions e.xponentially to the bathing medium 
during the initial 18 h in APW [regression coefficients 
(R) for Na', 0.998; Cl , 0.999; K\'o.994; P < 0.05 all 
regressions]. During the flax studv, the average net flux 
of' ions was; Na' /„ = -66.6 ± 8.1; Cl" /„ = -74.6 ± 
10.9; K* /„ = -7.5 ± 0.6 ixmol g' drv tissue h' (A' = 
5). The total solute oi the blood returned to values sim- 
ilar to APW controls bv 43 h. 

There was substantial mortality (50% in 48 h) when 
the animals were transferred from 377 mosmol kg' ' sea- 
water to APW, but similar groups of animals transferred 
to 161 and 108 mosmol kg ' ASW tolerated the hvpos- 
motic stress (>70% survival after 21 d). These animals 
rapidly became isosmotic to the new solute concentra- 
tion in the bath in 12 h (data not showm). In a similar 
experiment, animals acclimated tor 14 d to constant 269 
mosmol kg ' seawater were transferred into hvposmotic 
bathing media (APW, 108 and 161 mosmol kg ' ASW) 
and we observed a similar rapid reduction in blood ion 
composition within 12 h, but suni\al increased (>83% 
after 3 weeks) even in those animals transferred cUrectlv 
to APW. 



DISCUSSION 

An acute transfer of AP\\'-accIimated D poh/iDoiyha to 
172 mosmol kg ' was lethal to half of the animals in 3 
d. The observed intolerance was likeK' due to the limited 
abilitv of these animals to control the rapid rate of water 
and solute mo\ement liehveen the emironment and an- 
imal bodv fluid compartments. This inference is sup- 
ported by the observation that D. polymorpha survived 
step-wise acclimation to salinitv when the change was 
limited to increments of 54-108 mosmol kg '. Small 
changes in salinih and 1-3 d of acclimation beh\'een 
changes allowed the animals to survive salinities ap- 
proaching 269 mosmol kg ' before the onset of substan- 
tial mortalitv (Kilgour ct aJ.. 1994; Wright ct al. 1996; 
this studv). 

The response of D. polymoi'pha to ele\atecl salinity 
was to osmoconform. The isosmotic condition is accom- 
plished by the accumulation of salts, and partially bv the 
osTnotic loss of water (Dietz ct ai. 1997). The intracel- 
lular fluid compartment is subject to hmited regulation. 



Tabic 3. The blood ion composition of Drcisscna pi<liiiiH>rplui <icchmated to 377 mosmol kg ' artificial seawater for 14 davs and 
acuteK transferred to artificial pondudter (APW) 



mosmol kg 



Concentration (mmol 1 



Bathing medii 



N Total solute 



Other 



"Na- 



*C! 



K 



Ca^- 



*MS- 



APW controls 
377 mosmol kg 
APW, 6 hr 
APW, 24 hr 



39 ± 1' 
.386 ± 2' 

83 ± 3'' 

40 ± 1' 



2.1 ± 0.7'' 

-13.3 ± 4.5' 

14.2 ± 2.3' 

14.0 ± 1.5' 



16.7 ± 0.6' 

170.3 ± 2.0' 

28.2 ± 2.5'- 

10.5 ± 0.5-' 



15.2 ± 0.6 

206.8 ± 4.7' 

33.8 ± 0.8'' 

9.2 ± 1.7' 



0.5 ± 
5.2 ± 
1.0 ± 
0.5 ± 



0,1' 
01' 
0.1 



3.9 ± 0.3' 

4.6 ± 0.2' 

2.7 ± 0.2'' 
4.6 ± 0.4' 



0.4 ± 0.1' 
12.5 ± 0.2' 

3.0 ± 0.3' 

1.1 ± ().()■ 



Mean ± sem. "Other" was calculated from the difference between the total solute and measured .solutes. The negative "other " is 
due to complete ionization of NaCI and Ca-' salts when diluted for individual ion analyses. Means within a colunm with different 
letters are significanti) tlifferent by tlie Tukev-Kranier Studentized range procedure, P < 0.05; (*) log, transformed data tor statistical 
analyses. 



S. J. Wilcox and T. H. Dietz. 1998 



?dge 147 



These mussels have some abilih' to presene the ICF 
and make adjustments to celhilar \()hmie when thev are 
h\perosmoticall\ challenged (Dietz ct al. 1997; 1998). 
However, onlv mussels exposed to saline solutions con- 
taining sufficient K* are able to miiint;iin the ICF/ECF 
potassium gradient (Dietz ct al. 1997). Previous stutlies 
ha\e demonstrated that potassium a\ailahilit^- m the en- 
\nronment apparentlv is criticiil tor presening the intra- 
cellular volume, electrochemical balance, and survival 
(Dietz ct al.. 1994; Dietz ct al.. 1997). Drei.sscna pohj- 
morpha is similar to other freshwater bivalve species in 
that an increase in concentration of specific ions in the 
bathing medium results in a concomitant increase in the 
concentration of that ion in the blood (Deaton & Green- 
berg, 1991; Horohov ct al 1992; Wilcox & Dietz, 1995). 

A change in ASW of —100 mosmol kg ' at 3 d inter- 
vals allowed sufficient time for the animals to acchmate. 
This observation contrasts with an earher study where 
D. polijmorpha could not survive a week when 45 mmol 
1' NaCl was added to the APW (Horohov ct al. 1992). 
Drcisscna pohjmoiyha is the least tolerant of the fresh- 
water bival\'es when challenged with hvperosmotic me- 
dium of NaCl. In contrast, when the medium contmns 
45 mmol 1 ' NaCl and approximateh' 1 mmol I ' KCl, 
the h\perosniotic challenge is tolerated h\ this species 
(Dietz ct al, 1997; this stud\). Drcisscna pohjmoiyha 
will accumulate K* in the intracellular and extracellular 
fluid compartments from the bathing medium (Dietz ct 
al. 1997; 1998). The potassumi allows linutetl cellular 
volume regulation that likelv contributes to the animal 
survival in hvperosmotic conditions. 

Volume regulation in eui-vhaline and stenohahne h\- 
vaKe species is. in part, dependent on the rate of change 
of the free amino acid pool in response to a short-term 
osmotic challenge (Deaton & Greenberg, 1991; Silva & 
Wright, 1994; Neufeld & Wright, 1996). An acute trans- 
fer to a salinity of about 100 mosmol kg ' caused vir- 
tuiillv no short-term mortalitv in D, pohpnoiylia. Salin- 
ities between 120 and 150 mosmol kg ' were at the tol- 
erance limit for an acute exposure for this species. How- 
ever, freshwater bivalves depend primarily on 
electrolytes for \'olunie regulation as they ha\'e relativek 
low concentrations of free amino acids (Hanson 6c Dietz. 
1976; Potts, 1958; Dietz ct al. 1997). Although the free 
amino acid concentration is elevated with hyperosmotic 
challenge, it is a minor class of intracellular osmoKtes in 
freshwater bivalves (Potts, 1958; Dietz ct al. 1997). 

When mussels were exposed tor a week to a diimial 
cycle of saUnitv change of 14 to 130 mosmol kg ', thev 
repeatedly gained or lost ions from the blood depending 
on the direction of salinitv change. Most ot the ion flirx 
at higher ion concentration gratlients was attni)uteil to 
diffusion (see Wilcox & Dietz, 1995; Dietz ct al, 1997). 
Thus, the changes in body fluid solute concentrations 
with salinity c\'cles would be largely passive diffusion. In 
APW the mussels would be required to expend energy 
for osmoregulation, but less energy is expended as the 
animals osmoconfomi. Shumwav (1977) observed that 
the tissue water content in Mi/tihis cdiili.s remained rel- 



ati\elv constant during exposure to a sinusoidal full 
strength seawater/freshwater tidal cycle tor 7 d. It was 
suggested that animals exposed to an environment of 
constantK' fluctuating salinity would not continuously 
conform to the external sahnity but instead, maintain a 
tissue hydration level that required the least amoimt of 
energ\' expenditure. 

Drcisscna pohjmoiyha acclimatetl to 377 mosmol kg ' 
and transferred to APW lost ions and appeared to un- 
dergo a transient swelUng clue to osmotic water uptake. 
The reduction in ion concentration in the blood follow- 
ing transfer to APW was evident within 6 h, but the 
animals had not returned to control conditions by 24 h, 
as the concentrations of some ions were above and be- 
low those foimd in APW-acclimated controls. Many 
specimens could not tolerate the acute decrease in os- 
molalitv of 370 mosmol kg ' as 50% ot the animals died 
in 48 h. In contrast, the brackish-water bivalve, Corbi- 
ciila japonica, is capable of surviving the transition from 
425 mosmol kg ' seawater directly to freshwater with 
blootl total solute concentration returning toward control 
levels within a day (Matsushima, 1982). 

Several bivalve species exhibit a distinct hvpercalcem- 
ia upon exposure to dilute media and the response is 
inversely correlated with an increasing tolerance to 
freshwater (Deaton & Greenberg, 1991). The Ca-* in 
the blood of D. pohynoiyha acclimated to elevated sa- 
linitv, initially dropped upon acute exposure to APW, but 
was restored within 24 h to a concentration that was 
ecjual to that found in animals acclimated to APW (Table 
3), These animals Likely supplement the Ca-' concentra- 
tion in the blood with endogenous Ca-' sources (shell). 
However, D. yoh/moiyha blood Ca-' concentration 
changed very little with values ranging between 2 and 
4.6 mmol 1 ' under the conditions used in this study. 
Tliis species does not appear to utiUze Ca-* as a signif- 
icant extracellular osmolyte, but does regulate the con- 
centration within narrow limits. There appears to be a 
reciprocal relationship iietvveen the concentrations of 
Ca-* and Mg-' in the blood of D yohjmorpha. Mag- 
nesiinn was poorlv regulated at the higher concentra- 
tions tfjuntl in dilute ASW, but was hvper-regulated bv 
the annuals acclimated to APW (Dietz ct a/.. 1994). 

Drcisscna pohjmoiyha appears to be a freshwater 
mussel in transition from its brackish water ancestry. 
These animals evolved in freshwater connected to inland 
seas having minimal diurnal Sitlinitv changes. The ap- 
parent eunhahnitv' ot D. polijinoiyha suggests that these 
bivalves have some capacity to tolerate ohgohahne hab- 
itats. However, this species will likelv be restricted to 
low salinitv environments that have slowly changing sa- 
linity cycles. Drcisscna pohjmoiyha are not likelv to tol- 
erate habitats with rapid and large salinity fluctuations 
associated with daily tidal cycles. 

ACKNOWLEDGMENTS 

We thank S. J. Nichols for providing animals and Dr 
Harold Siherman for the many suggestions and com- 



Page 148 



THE NAUTILUS, Vol. Ill, No. 4 



inents. Diondi Lessard, Tim Smith and Julie Cherr\' pro- 
vided technical assi.stance. Thi.s work vva.s supported bv 
NSF grant DCB90- 17461 and Louisiana Sea Grant 
NOAA 46RG0()960 Project R/ZMM-1. 

LITERATURE CITED 

Aladin. N. V., and \\: T \\' Pott.s. 1992. Cluuiges in the Aral 
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B\Tne, R. A. and T. H. Dietz. 1997. Ion transport and acid- 
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Chambers. E. L. and J. De Aniiendi. 1979. Membrane poten- 
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Deaton, L. E. and M. J. Greenberg. 1991, The adaptation of 
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Dietz, T, H., D. Lessard, H. SiKerman and J. W. L\iin. 1994. 
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THE NAUTILUS 111(4):149, 1998 



Page 149 



Notices 



UPC:OMING MEETINGS, CONVENTIONS, AND 
WORKSHOPS 

1998 Annua] Meeting ol the North American Bentho- 
logical Society, Charlottetovvn, Prince Edward Island, 
Canada, 2-5 June 199S. Contact: Dr Jill Lanca.ster, 
j.lancaster@ed.ac.uk 

199S Conchologists of America Convention, Orlando, 
Florida, USA, 19-2.3 JuK 1998. Local contact: Lmda 
Koestel, lkoestel@magicnet.net 

1998 World Congress of Malacology, Washington, DC, 
USA, 2.5-.3() Julv 1998. Combining the 13"' International 
Congress, Unitas Malacologica, and 64''' Annual Meet- 
ing, American Malacological Union together with other 
participating malacological organizations. Local contact: 
Dr Robert Hershler, hershlerrobert@nmnh.si.edu 

The 8''' International Congress on Invertebrate Repro- 
duction and Dexelopment (ICIRD), Amsterdam, The 
Netherlands, 10-15 August 1998. Contact: 
VU_conference@dienst . vu . nl 



(Jlobal Biodiversit)' Forum, 29-31 August 1998, Mon- 
treal, Canada. Contact: celias@wri.org 

III Symposium "Fauna and Flora of the Atlantic Is- 
laiuls", Ponta Delgada, Azores Islands, Portugal, 21-25 
September 1998. Contact: simposio@alfnac.pt 

Annual Meeting of the Society of Integrative and Com- 
parative Biologv (iormerly The American Society of 
Zoologists), Denver, Colorado, USA, 6-10 January 1999. 
Held jointK' with The American Microscopical Society 
and The Cmstacean Society meetings. Contact: 
corinne_o'brien@sba.com 

Annual Meeting of the Societv' for Conservation Biology, 
College Park, Maryland and Washington, DC, USA, 17- 
21 June 1999. Local contact: David Inouye, 
di5@umiul.umd.edu 

IV Congreso Latinoamericano de Malacologia, Coqiiim- 
bo, Chile, 1999. Local contact: Dr Chita Guisado, 
cguisado@nevados.ucn.cl 



THE NAUTILUS 



Volume 111 
1998 




AUTHOR INDEX 

BiKi.ER. R 1 Okti;\, J 45 

BOUCMKT, P. 47 P\STI)Kl\<l, G 117 

Diet/.. T.H 143 Pi:n'< h, E.J 22 

EspiNosA, J 45 Saii., L.R 119 

Geickk, D.L 85 Vermeij. G] 47, 53 

Kantor, Y 117 WiEc:o\, S.J 143 

MiKKELSEN, PM 1 Wise. J.B 13 



NEW TAXA PROPOSED IN VOLUME 111 (1998) 

GASTROPODA 

Alarimella Saul, 1998, new genus ( Aporrhaidae) 134 

Alariiiwlla anac Saul, 199S, new species (Aporrhaidae) 137 

Aporrhais drachiiki Saul, 1998, new species ( Aporrhaidae) 121 

Latiala hehaca Saul, 1998, new species (Aporrhaidae) 128 

Latiala sionta Saul, 1998, new species (Aporrhaidae) 130 

Ficu.s villai Petuch, 1998, new species (Ficidae) 33 

Fusulciilns Bouchet & V'eniieij, 1998, new genus (Pseudolividae) 47 

Fiisulculu.s albus Bouchet & V'eniieij, 1998, new species (Pseudoli\idae) 49 

Fusuladas crenatus Bouchet & \"enneij, 1998, new species (Pseudoli\idae) 49 

Canccllaria mediamcricann Petuch, 1998, new species (Cancellariidae) 35 

Conus (Leptoconus) paschnlli Petuch, 1998, new species (Conidae) 36 

BrV'ALVIA 

Noetia [s.s. ) lindae Petucii, 1998, new species ( Arcidae) 37 

Plicatula miskito Petuch, 1998, new species (Plicatulidae) 38 

Mactra inccri Petuch, 1998, new species ( Mactridae) 39 

Micromactra miskito Petuch, 1998, new species (Mactridae) 40 

Petricola (Petricolaria ) donnae Petuch, 1998, new species (Petricolidae) 41 

REX'IEWERS FOR N'OLUME 111 

A. Bogan, R Bouchet, \V. P Elder, M. G. Harasewvch, D. G. Herbert, P M Mikkelsen, G Pastorino, E. J. Petuch, R. L. Squires, 
E. E. Strong, G. J. Venneij, A. Waren, J. B. Wise. 



INSTRUCTIONS TO AUTHORS 



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