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Volume 122. Numbcrl 
March 28, 2008 
ISSN 0028-1344 

A quaiierhj devoted 
to malacologij. 


Dr. Jose H. Leal 

The B alley- Mattliews Shell Museum 
3075 Sanibel-Captiva Road 
Sanibel, FL 33957 


Maiy Jo Bunnell 

The B alley- Mattliews Shell Museum 
3075 Sanibel-Captiva Road 
Sanibel, FL 33957 


Dr. M. G. Harasewych 
Depaitment of Invertebrate Zoology' 
National Museum of 

Natural Histoiy 
Smithsonian Institution 
Washington, DC 20560 


Dr. IJiidiger Bieler 
Department of Invertebrates 
Field Museum of 
Natural Histoi^v 
Chicago, IL 60605 

Dr. Aithur E. Bogan 

North Carolina State Museum of 

Natural Sciences 
Raleigh, NC 27626 

Dr. Philippe Bouchet 
Laboratoire de Biologic des 

Invertebres Maiins et Malacologie 
Museum National d'Histoii-e Natui-eUe 
55, rue Buffon 
Paris, 75005 France 

Dr. Robert H. Cowie 

Center for Consewation Research 

and Training 
University of Hawaii 
3050 Maile Way, Gilmore 409 
Honolulu, HI 96822 

Dr. Robert T. Dillon, Jr. 
Department of Biolog)' 
College of Charleston 
Charleston, SC 29424 

Dr. Eileen H. Jokinen 
8234 E. North Shore Road 
Sauk Ste. Marie, MI 49783 

Dr. Douglas S. Jones 
Florida Museum of Natiu^al Histoiy 
University of Florida 
Cainesydlie, FL 32611-2035 

Dr. Harry G. Lee 

4132 Ortega Forest Drive 

Jacksonville, FL 32210 

Dr. Charles Lydeard 
Biodiversity and Systematics 
Department of Biological Sciences 
University of Alabama 
Tuscaloosa, AL 35487 

Bmce A. Marshall 
Museum of New Zealand 

Te Papa Tongarewa 
P.O. Box 467 
Wellington, NEW ZEALAND 

Dr. James H. McLean 
Department of Malacology 
Natural History Museum 
of Los Angeles County 
900 E.xposition Boulevard 
Los Angeles, CA 90007 

Dr. Paula M. Mikkelsen 
Paleontological Research 

1259 TiTunansburg Road 
Ithaca, NY 14850 

Dr. Diarmaid O Foighil 

Museum of Zoology and Depaitment 

of Biolog}'" 
University of Michigan 
Ann Arbor, MI 48109-1079 

Dr. Gustav Paulay 
Florida Museum of Natural Histoiy 
Universitv of Florida 
Gainesville, FL 32611-2035 

Mr. Richard E. Petit 
P.O. Box 30 

North Myrtle Beach, SC 29582 

Dr. Gary Rosenberg 
Department of Mollusks 
The Academy of Natural Sciences 
1900 Benjamin Franklin Park^way 
Philadelphia, PA 19103 

Dr. Angel Valdes 
Department of Malacology 
Natural Historv Museum 
of Los Angeles County 
900 Exposition Boulevard 
Los Angeles, CA 90007 

Dr. Geerat J. Vermeij 
Department of Geology 
University of California at Davis 
Davis, CA 95616 

Dr. G. Thomas Watters 
Aquatic Ecology Laboratoiy 
1314 Kinnear Road 
Columbus, OH 43212-1194 


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VohDiic 122, Ntniihcr 1 

Marcli 2S, 2()()S 

r>,^xn^,7V-rc ISSN 0028-1344 


Philippe Boiic'het New species and new records nl stiutliwest Pacific C^anci'llariidai' 

Richard E. Pelil (Gastropoda) 1 

Vittorio Carilli On some Neogene to Recent species related to Galcodina Monterosato, 18'S4, 

Gdlcodinopsif; Sacco, 1895, and Massotia Bucquov, Dautzenl)erg, and Dollius, 
LSS4 (Caenogastropoda: Rissoidae) vwth the description of two new Alvaiiia 
species from the Mediterranean Pleistocene 19 

Diego G. Zelava Reallocation oi ('i/dinidcurdiinn cidssildhnuii Dell, 1964, into Pcniciiita 

Bernard, 1897 (Bi\'aK'ia: Cvamiidae) 52 

Notice 56 

MBLWHOI Library 

Massachusetts 02543 

THE NAUTILUS 122(1 ):1-1S. 2008 


New species and new records of sonthwest Pacific 
Cancellariidae ( Gastropoda) 

Pliilippe Boiichet 

Museum National il'llistoire Naturel 

55, me Button 

75005 Paris. FRANCE' 

Richard E. Petit 

SO(i St. Ciuules Itoad 

North N'hrtle Beach, SC 295S2 USA 


Fifteen species of Cancellariidae referable to the genera Zead- 
inetc, A^dinetidn. Fiisiaplwra. Nippoiinphcra, and Trigonostoma 
are reported from deptiis between 200 and TOO m in New 
Caledonia and other island groups in the southwest Pacific. 
TweKe ai'e new species: Zcadmete batln/oinoii new species, 
Xcadmcte plii/somon new species, Zeadmete bilix new species, 
Admctida affluens new species, Admctula marshalli new spe- 
cies, Adinetula bathijnoma new species, Adinctidd Iiiica new 
species, Admctida ciiiart^inata new species, Nipponaphcra argo 
new species, Nipponapliera agastor new species, Nippona- 
phcra tuba new species, and Trigonostoma tiiiblium new spe- 
cies. .\11 the Recent nominal species of Fusiaphcra described 
from localities throughout the Indo-Pacific area are considered 
to be conspecitic. the senior name being Fusiaphcra mac- 
rospira (Adams and Reeve. 1850), now with ten s)monyms. The 
ranges oi yippouaphera nodosivaricosa (Petuch, 1979) and 
Trigonostoma thijsthJou Petit and Harasew\ch, 1987, are ex- 
tended to the South Pacific 


The present paper is a continuation oi oin- studv of the 
deep-w^ater cancellariid fauna of the Southwest Pacific, 
based on the material originating from recent expedi- 
tions in New Caledonia, Vanuatu, Fiji, Tonga, Wallis & 
Futuna, and the Solomon Islands. In a previous paper 
(Bouchet and Petit, 2002), we described the new genns 
Mirandaphera and nine new species in the genera Afri- 
cotriton, Meiica. Sveltia, and Nipponapliera. We here 
deal with 15 species (12 new) in the genera Zeadmete, 
Admetula. Fusiapliera, and Trigonostoma, and add fur- 
ther species in Xipponaphcra. Oiu^ re\iew of the deep- 
water cancellariid fauna so far sampled in the southwest 
Pacific \^ill be complete after a third paper (in prepara- 
tion) dealing with the genera Brocchinia, Microcancilla, 
and Gcrgovia. In adchtion, the cancellariid fauna of New 
Caledonia includes shallow-water to offshore species in 
the genera Scalptia (5 species) Trifonoliaiya (several 
species), as well as the rediscovered endemic Mcrica 
semperiana, which we intend to deal with separately. 
Cancellariid radulae are not known to pro\ide dis- 

criminating species-level characters, and we tiius ditl not 
attempt to systematically examine them when we had 
live-taken specimens available. Much of the material re- 
ported in this series was collected in the 1980-1 991 )s and. 
at the time, fixed in iormalin and thi-n rinsed and dried. 
It is thus not adequate for nucleic-acid sequencing. More 
recent expeditions generate new material that is specifi- 
eallv put aside for barcoding. Oiu" treatment of the can- 
cellariid lamia is tlius currentlv restricted to a description 
of the shells, including the protoconch, but we mav ex- 
pect that in the future it will be possible to test some of 
oiu' species limits thi'ough molecular characters. 


In till.' lists of t)rpe and otiiei' material e.xaniined, indi- 
vidual lots in MNHN are unambiguously designated by 
the combination of cniise acronym (capitalized) and sta- 
tion number. DW (for Drague Waren) refers tfj dredge 
hauls, CP (for Chalut a Perche) to beam trawls; 1\' refers 
to live-taken specimens, dd to emptv' shells; spms to in- 
di\iduals that cannot be unambiguously assigned to one 
of these two categories (essentially commercially ob- 
tained specimens). Institutional acronyms are: AMNZ: 
Auckland Museum, Auckland, New Zealand; BAINH: 
The Natural Histon' Aluseum, London, UK; DMNH: 
Delaware Museum of Natural Histoiy, Wilmington, 
Delaware, USA; MNHN : Museum National d'Histoire 
Naturelle, Paris, France; NM: Natal Museum, Pieterma- 
ritzburg, South Africa; NMW: National Museum of 
Wales, Cardiff, UK; NSMT: National Science Museum, 
Tokyo, Japan; USNM: National Museum of Natural His- 
toiy, Smithsonian Institution, Washington, DC, USA; 
WAM; Western Austrahan Museum, Perth, Austraha. 


Family Cancellariidae Forbes and 
Genus Zeadmete Finlav, 192fi 


I lev. 


Type Specie.s: Ciincelhiria trailli Hutton, 1S73, bv 
original designation. Recent, New Zealand. 

Pase 2 

THE NAUTILUS, Vol. 122. No. 1 

Discussion: The genus Zcaclmete was proposed by 
Finlav (1926: 429) who later (1930b: 242) considered it 
to be a subgenus of Oaiiianiia Finlav, 1924 from the 
Lower Miocene of New Zealand. Powell (1979: 224) also 
treated Zeadmefe as a subgenus of Oamariiiii. However, 
Garrard (1975: 44) and Wilson (1994: 173) restored 
Zcadinete as a full genus, as did Petit and Harasewych 
(2000: 151), who gave a brief discussion of its possible 
relationship to other taxa. Among other differences, the 
type species of Oamaniia. Aclmctc sutcii Marshall and 
Murdoch, 1920, has strong columellar folds whereas 
Zeadmefe has weak, almost obsolete, folds. 

The genus Zeadmefe, as inteq^reted here, occurs in 
the Miocene to Recent faunas of New Zealand and in the 
Recent faunas of South Africa, Fiji and New C^aledonia 
(Z. kiilanda Garrard, from Australia, is probably an Ip- 
hiii(>j)sis.) Expedition material in MNHN also contains 
an undescribed species from the Solomon Islands and 
another one from New Caledonia, both represented by 
specimens too imperfect to be named. All live offshore in 
300-600 m, with New Zealand records as shallow as 65 

Zeadmefe bathi/omon new species 
(Figures 1-2) 

Description: Protocoucli glass\, suioolh, erect, oi 
about 1.3 whorls, diametei' 900 (xm. Teleoconch of four 
wliorls. Transition to teleoconch marked bv a sharp 
growth line and a spiral cord forming shoulder angle, 
shortly followed by onset of weak but shaip axial ribs, 
forming small nodes at strengthening shoulder angle. Ad- 
ditional spiral cords appear before end of first teleoconch 

whorl. Shoulder angle prominent, bearing minute coro- 
nations formed by intersection of spiral cords and axial 
ribs. Sutm-al ramp strongly concave, bearing five to seven 
fine, low spiral cords. Suture attachment orthogonal. 
Shoulder raised, bearing three to five closely spaced spi- 
ral cords. About 20+ wider, low spiral cords anterior to 
shoulder angle, extending down onto base. Fine, low, 
evenly spaced axial ribs extending from shoulder angle to 
anterior end of shell; about 25 on last whorl; interspaces 
between axial ribs bearing closely packed fine grow1:h 
lines. Low, narrow varices, formed onlv bv a slight raising 
of shell surface, at about 120° increments on final whorls. 
Outer lip smooth, faintK' sinuate adapically, cumng back 
in prosocline direction aiiapically. Aperture elongate, 
narrowly elliptical, without lirations uiside outer lip. Pa- 
rietal area with veiy thin, narrow callus. Columella with 
three folds, the posterior two weak; anterior fold broad, 
situated on small siphonal fasciole, forming edge of m- 
ductmal callus, which then forms edge of distinct sipho- 
nal canal. Exterior lieige, rims of varices pale straw- 

Type Material: Holotype MNHN 20498, 10.2 x 4.8 

Type Locality: Soutli of New Caledonia, 22°17' S, 
167°12' E, 390 m [VAUBAN 1978-79: sta. 3]. 

Material Examined: Norfolk Ridge, south of New 
Caledonia.— VAUBAN 1978-79: sta. 3, 22°I7' S, 
167°12' E, 390 m, I Iv (Figure 1).— BIOCAL: sta. 
DW77, 22°15' S, 167°15' E, 440 m, 1 dd.— BATHUS 2: 
sta. DW719, 22°48' S, 167°16' E, 444-445 m, 1 Iv.— 
SMIB 8: sta. DW166, 23°38' S, 167°43' E, 433-450 m. 

Figures 1-5. Zeddinclc. 1-2. Ztridmefe had^i/oinon new species, 1. lioli)t\pc, heit^lit 1(1.2 iiiiii: New Caledonia, Nortulk Kidge, 
22"17' ,S, I67'^12' E, 390 ni [Vfiiihan 1978-79 sta. .3]. 2. 23°38' S. 167°43' E, 4.3.3-450 iii |SM1B 8 sta. DW166]. 3, 4. Zcmliiwte 
plujsninoH new species, liolotyjie, height 13.1 mm, Fiji, 19°01' S, 17S°25' E, 500-516 m [BCJKDAU 1 sta. D\V14S8]. .5. Zcadinete 
Inlix new species, holotype. height 13.2 mm;. New Caledonia, Norfolk Ridge, 23°02' S, 168n6' E, 3.35 m [SMIB 5 sta. 15W98]. 

p. Bouchet 

H. E. Petit. 2008 


1 dd (Figure 2); Sta. D\\'167. 23°3S' S, 167°43' E, 430- 
452 m, 1 dd— NORFOLK 1: sta. D\\1666, 23°42' S, 
167°44' E. 469-860 ni. 2 dd.— NORFOLK 2: ,sta. 
DW2024, 23°2S' S. 167°5L E, 370-371 m. 1 dd. 

Et\ inologA": Fnini tlie Greek biitltus, deep, combined 
with (>iH(>\. .shoulder, for the deepK' eonea\e sutural 
ramp. To Ix' treatetl as a noun in apposition. 

Di.sti-ibutioii: Known onh' Irom south ot New Cale- 
ilonia, ali\e in 390—144 m. 

Remarks: In Zcaclincti' finhn/i Powell, 1940, the shoul 
der i,s also raised hut the sutural ramp is not conea\e. In 
Z. hathijomon, the shoulder is even with, or raised abo\"e, 
die suture. Also, Z.finknii has clathrate sculpture only on 
the adapical half of the last whorl, whereas the last whorl 
is entireK' clathrate in Z, baihijomon. The aperture is 
two-thirds of the shell height in Z. fiiilai/i, but in Z. 
batlujoinon the aperture is just o\"er hall the shell height. 
The onlv other Zeadmctc species we are aware of wth 
a sutural ramp that does not slope abapicallv is an vmde- 
scnbed species represented h\ two broken, worn speci- 
mens from a nearbv station on the Norfolk Ridge 
[BER-i-X 11: sta. DW35, 23°33' S, 167°16' E, 550-570 
m], which we leaxe undescribed because of the poor 
condition of the specimens. It differs from Z. bathi/onwn 
in haxing a shorter spire and a flatter sutural ramp that is 
de\"oid of spiral cords. It also has a largei' piotoconch 
\\ith a diameter of 1100 (im. 

Zeadmete phijsonioii new species 
(Figures 3^) 

Description: Protoeonch smooth, gloss\', ot 1.5 
whorls, diameter 1250 |jLm. Transition to teleoconch 
marked b\" weak axial rili and faint spiral cords, both of 
uhich increase in strength rapidh'. Teleoconch of four 
whorls, shell thiTi. Spiral cords flat, evenly spaced, \vith 
interspaces slighth" narrow-er than cords, about fi\'e on 
sutural ramp and 20+ anterior to shoulder angle. Axial 
ribs extending from suture to anterior end of teleoconch, 
interspaces considerably broader than ribs; about 20 ribs 
on last whorl. Sutural ramp slightly convex, sloping up to 
impressed suture. Aperture elongate-elliptical, only 
weakly angulate at shoulder, smooth within. Outer lip 
sinuous adapically but becoming prosocline anterior to 
peripherv'. Parietal wash indistinct on holot\pe, distinct 
on one parab.'pe. Columella almost vertical, bearing 
tliree folds, posterior fold on top of siphonal fascicle. 
Anterior folds callused, widely separated, anterior one 
almost obsolete, forming edge of short siphonal canal. 
Exterior cream with ill-defined pale brown band on pe- 
riphers' of spire whorls; some ribs on last whorl have 
brownish tint. 

T\-pe Material: IIolot\pe MXIIX 20519 (13.1 x 6.5 
m'm) and 6 parat\pes VINHN 20520-20521. 

T^pe Localitv-: Lau Ridge. Fiji, 19°01' S, 178°25' E, 
500-516 m [BORDAU 1: sta. DW1488]. 

Material Examined: Fiji. BORDAU 1: sta. DW 1486, 

19°01 ' S, 178°26' E, .395-540 m, 1 del parat\pe MNHN 
20520.— Sta. D\\T488, 19°0I' S, 178^25' E,' 500-516 m, 
6 M. holotxpe MNHN 20519, paratypes MNHN 20521 
(Figures 3—1). 

Etymology: From the Greek /j/»/.vrto, inflated, andonnw, 
shouldei', lor the appearance of the sutural ramp. To be 
treated as a noun in apjiosition. 

Di,stril)iilion: Known onlv from Fiji at the two stations 
listed above 

Remarks: Protoeonch diameter in paratopes ranges 
from 900 to 1300 jim. Zi-ddnicic phijsomon differs from 
Z. batlu/omon in having narrowei' spiral cords with wider 
interspaces, fewer, more wideK' spaced axial ribs, and a 
shghtly convex rather than strongK' concave sutural 
ramp. Also, in Zeadmete phi/sonum the axial ribs remain 
distinct below the peripheiy. 

Zeadnute hilix new species 
(Figin-e 5) 

Description: l^'otoconch prominent, smooth, glassy, 
of 1.1 whorfs, cUameter 875 fjim. Transition to teleoconch 
marked by onset of axial ribs and spiral cords. Teleo- 
conch high-spired, of five whorls. Spiral cords narrow, 
interspaces broader than cords. About tliree to foin^ fine 
spiral cords on sutural ramp; eight coi'ds anterior to 
shoulder angle on penultimate whorl. Last whorl with 
about 17 narrow spiral cords, one of which forms shoul- 
der angle and another, only slightly more prominent than 
those adjacent, is just posterior to periplieiy, giving te- 
leoconch a weakly biconic appearance. Axial ribs fine, 
spaced almost equal to spacing of spiral cords, rendering 
teleoconcli surface evenly reticulate; about 55 on last 
whorl. Sutiu-al ramp nari'ow. Apertru'e narrowly elliptic. 
Outer lip orthocline adapically, becoming prosocline 
only at anterior end. Columella almost straight, bearing 
three folds; adapical one on siphonal fasciole, other two 
on a broad hea\y callus; anterior one obsolete, forming 
edge of short siphonal canal. Periostracnm pale browai, 
shell pale straw \ellow. 

Type Material: IIolot)pe (dd) MNHN 2(J499 (13.2 x 
5.6 mm). 

Type LocalitA': Norfolk Ridge, south ot New Cale- 
donia, 23°02' S, 168°16' E, 335 m [SMIB 5; sta. DW98]. 

Material Examined: OnW known fmiii the holotxpe 

(Figure 5). 

Etymology: Latin adjective /);7/.v. lia\ing a donlile 
thread, with reference to the two strong spiral coi'ds 
around the shoulder angle. 

Distribution: Known only from New Caledonia at the 
t\pe locality. 

Remarks: Zeadmete hilix is placed in Zeadmete based 
on its cohnnellar stmcture, which is identical to that of 
other Zeadmete species, but it ma\ be immediately cBs- 

Page 4 

THE NAUTILUS, Vol. 122, No. 1 

tinguished from its congeners bv its elongate shape and 
relatively short aperture. 

Zeadmetc bilix superficially resembles the buccinid ge- 
nus Inrlahihi Finla\'. 1926, but species o{ Ire d alula have 
a recuned, notched siphoual canal and lack columellar 

Genus Adiucfnla Cossmann, 1889 

T)pe Species: Biicciiuiiii cvidsum Solander, 1766, by 
original designation. Eocene, British Isles. 

Discussion: Admetula is often placed in the synonymy 
u{' Boncllitia Jousseaume, 1887 (see discussions in Sacco, 
1S94: 42; Cossmann, 1899: 33; Davoli, 1982: 62; Ver- 
hecken, 1986: 33; but not Verhecken 2007: 286), but we 
regard BoiicUifia as distinct, based on the angled whorls 
and muricated sculpture. We refer to Landau, Petit, 
and Marquet (2006) for imther discussion. The genus 
Admetula is well represented in Paleogene and Neogene 
Tethvan faunas and in the Recent iauna along continen- 
tal margins at depths ranging from 75-700 m (Petit and 
Harasewych, 1991: 181).' 

Mam' species of Admetula appear superficially veiy 
similar, especially based on published illustrations, yet 
can easily be distinguished when directly compared. 
Three discrete "subgroups'" can be recognized in the ge- 
nus. One consists of small attenuate species such as A. 
eontidei (Altimira, 1978), A. epula Petit and Harasewych, 
1991, and A. afra Petit and Harasewych, 2000. Larger, 
wide, rather thin-shelled species such as A. haijeri Petit, 
1976, and A. cmarginata (described herein) form a sec- 
ond group, witli a third, intermediate group composed of 
thick shells with an anterior constriction of the last whorl 
such as A. vossi Petit, 1976, and A. dcroijae (Petit, 1970). 
Early Tertiaiy species usually have well-formed varices at 
irregular intenals, a feature not found on Recent species. 

Admetula afflueus new species 
(Figures 6-8) 

Description: Protoconch corroded on holotvpe, in 
other specimens consisting of 0.8 whorls, diameter 775 
[xm, shiny, smooth apically, with sl\ raised threads on 
aliapical part. Transition to teleoconch distinct, marked 
by onset of teleoconch sculpture. Teleoconch spire high, 
spire angle 47°; teleoconch consisting of 6.25 whorls, 
with sculpture ot evenly spaced, narrow axial ribs crossed 
by spiral cords forming small nodes at intersections. 
About 15 axial ribs on penultimate whorl, about IS on 
last whorl. About 12 primary spiral cords on last whorl, 
with spacing ecjual to that of axial ribs on shoulder and 
periplieiy, moi'e crowded on base; four to seven second- 
ary spiral cords in each interspace. Outer lip shai-p, 
prosocline. .'Vperture without lirations. Parietal area with 
very thinly applied callirs. Columella sloping, bearing two 
prominent folds extending to edge of inductural callus, 
witli a third broad siphonai fold. Siphonal canal shallow, 
well defined. Exterioi' white with thick, pale olive-brown 

Type Material: Holot\i:)e MNHN 20500 (22.7 x 13.0 
nim) and 4 paratApes MNHN 20501-20502. 

Type Locality: SW of Malaita, Solomon Islands, 
09°46' S, 160°53' E, 611-6.36 m [SALOMON 1: sta 

Material Examined: Solomon Islands. SOLOMON 1: 
sta. CP1749, 09°21' S, 1.59°56' E, 582-.594 m, 1 dd.— 
Sta. CP1750, 09°16' S, 159°55' E, 69.3-696 m, 1 dd.— 
Sta. CP1751, 09°10' S, 159°53' E, 749-799 m, 2 Iv (1 
paratype MNHN 20501).— Sta. CP1793, 09°13' S, 
160°d8' E, 505-510 m, 1 dd, 1 juv. dd (Figure 8).— Sta. 
CP1798, 09°2r S, 160°29' E, 513-564 m"2 Iv, 1 dd.— 
Sta. CP1808, 09°46' S, 160°53' E, 611-636 m, 1 Iv (ho- 
lotvpe. Figures 6-7).— Sta. CP1859, 09°33' S, 160°37' E, 
283-305 m, 1 Iv, 2 dd (3 paratopes MNHN 20502). Total 
of 13 specimens. (Largest specimen: 26.3 x 15.6 nun.) 

Distribution: Known onK^ from the Solomon Islands, 
alive in 305-749 m. 

Etymology: From the Latin afflnens, an adjective 
meaning abundant or copious, with reference both to its 
large size and relative abundance in the Solomon Archi- 

Remarks: Admetula affluens superficially resembles 
the specimen figured as A. garrardi (Petit, 1974) by Ha- 
segawa (2000: 585, pi. 291, figure 26) but differs bv bav- 
ins narrower axial ribs, a more constricted base and a 
mtjre twisted columella. It lacks the lirations within the 
outer lip that are present in A. garrardi. It is our opinion 
that the specimens figured as A. garrardi bv Hasegawa 
(2000) and Verhecken (1997: :306, figs. 11-13) are" not 
that species. Verhecken (1986: 34-35, figs. 1-2) exam- 
ined, redescribed and figured the holotype of A. garrardi 
and explicitlv described its nniltispiral protoconch 
whereas A. afflueus has a paucispiral protoconch. 

Of the other species of Admetula in the tropical south- 
west Pacific, A. affluens is more similar to A. marshalli, 
but differs by its larger adult size, less solid shell, much 
weaker spiral cords ;ind lack of apertural lirations. 

Admetula emarginata new species 
(Figures 9-11) 

Description: Protoconch glossy, white, of 1.1 whorls, 
tlianu-ter 1001) ixm, with live widely spaced spiral cords. 
Transition to teleoconch indistinct, protoconch cords 
continuing as teleoconch cords. Teleoconch of five 
whorls, sutuj-e impressed, wath sculpture of numerous 
prominent, well-defined axial ribs crossed bv spiral cords 
forming small nodes at intersections. About 14 widely 
spaced axial ribs on penultimate whorl and about 12 on 
last whorl, final one enlarged into terminal v'arix. About 
eight primai-y spiral cords on last whorl, nioic closely 
spaced than axial ribs, with one or more secondan' spirals 
in each intcispace and about sLx prominent secondan 
spiral coicls on spire whorls below rounded shoulder 
anc-le. Shell thin, a\i;il ril)s \-isibl(> tlirouHi last whorl. 

Figures 6-17. Admctula. 6-8. Admctula afflucns new species. 6-7. Holotvpe, height 22,7 mm, Solomon Islands, 09°46S, lfi0°53' 
E, 611-636 m. [SALOMON 1 sta. CP180<S].'8. Protoconch. Solomon Islands, 09°13' S, 160°0S' E, 505-510 m, [SALOMON 1 stu. 
CP1T93]. 9-11. Admetuh emarginata new species. 9-10. Holohpe, height 16 mm. Coral Sea, 20°03' S, 158°45' E. 315 in 
[MUSORSTOM 5 sta. 335]. 11. Protoconch, Coral Sea, 19°33.5' S, 158°.30.5' E, 230 m [CORAIL 2 sta. D31]. 12-13. Admctula 
marshalh new species, holot^pe, height 14.7 mm. Fiji, 16°39' S, 179°57' W, .591-596 m [BORDAU I sta. CP1396]. 14-15. Admctula 
lutca new species. 14. holot\pe, height 13.1 mm', Tonga, 21°19' S, 175°01' W, 22.5-2,33 ni [BORD.VU 2 sta. DW152]]. 15. 
Protoconch. Fiji. 1S°09' S, 178°39' \V, 290-.300 m [BORDAU 1 sta. DW1465]. 16-17. Admctula bathijnomu new species. 16. 
Holot\pe, heisjht 7.4 mm. New Caledonia, 22°52' S, 167°23' E, .590-600 m [MUSORSTOM 4 sta. DW225]. 17. Protoconch, New 
Caledonia, 22°52' S. 167°16' E, 5.30-541 m [BATHUS 2 sta. DW720]. All protoconchs at the same scale, scale bar 500 \i.m. 

Pase 6 

THE NAUTILUS, Vol. 122, No. 1 

Outer lip prosocline, sinuous, with everted stromboid 
notch just anterior to peripheiy and another everted 
notch near base. Apertiu'e wdth nine lirations, visible in- 
temalK' at level of terminal varLx, not extending to edge 
of lip or tlecplv into aperture. Parietal area with very 
thinlv applied callus. Columella sloping, with Uvo promi- 
nent folds extending to edge oi inductiuai callus and a 
third broad siphonal fold. Siphonal canal shallow, well 
defined. Shell white, with weakly defined bands of pale 
browai at shoulder and base, also browai behind outer lip. 

Type Material: Holotx^pe MNHN 20503 (16.0 x 10.9 
mm) and one parat)'pe MNHN 20504. 

Type Locality: Coral Sea, 20°03' S, 158°45' E, 315 m 
[MUSORSTOM 5: sta. 335]. 

Material Examined: Choral Sea. CHALCAL: sta. 1^31, 
19°33.5' S, 158°30.5' E, 230 m, 1 dd (paratvpe. Figure 
11).— MUSORSTOM 5: sta. 335, 20°03' S,15S°45' E, 
315 m, 1 dd (holotype. Figure 9-10).— EBISCO: sta. 
CP2571, 20°25' S, 158°45' E, 298-309 m, 1 dd. 

Etymology: From the Latin eiiiargiiiiitii.s alter the 
sinuous, everted outer lip. 

Distribution: Known onK bom the Coral Sea near the 
Chesterfield Islands, emptv shells in 230-315 m. 

Remarks: Achnetida cinarginata differs irom other de- 
scribed species of Admctula in having diffused colora- 
tion. It is much like A. Innjcri Petit, 1976, from the west- 
ern Cull of Mexico in lumng at the peripheiv a notch, 
which is not present in the similar species A. hathijiutind. 
Adnictida citwrginata also differs from A. hathijiioina in 
having less closelv spaced spiral cords and itxial ribs. The 
overall effect of the sculpture on A. emarginata is a strik- 
ing pattern of horizontal rectangles crossed by fine spi- 

Ailiiii'tidd iniirsli(dJ'i new species 
(Figures 12-13) 

Description: Protoconch glos.s)', brown, of one whorl, 
diairieter 825 |jLm, witli three widely spaced spiral cords. 
Transition to teleoconch marked by a change in color and 
onset of axial ribs. Teleoconch of 5.5 whorls. Suture im- 
pressed. Spiral cords closely spaced, about 14 priman- 
cords on last whorl with one secondaiy cord in most 
interspaces, forming small pointed beads where they 
cross axial ribs. One spiral cortl creates a modest angle 
between peripheiy and narrowly rounded shoulder. 
About 14 rounded axial ribs on last whorl, only filial one 
enlarged into a varix. Fine, densely packed growth lines 
on entire teleoconcli. Outer lip thin, veiy indistinctly 
notched just anterior to peripheiy and with a slight ever- 
sion of the siphonal canal. Aperture with nine lirations, 
visible interior to terminal varix but not extending deeply 
within aperture or to outer lip. Parietal area with thin but 
distinct callus. Columella concave, bearing three distinct 
folds that extend out to edge of inductural callus. Poste- 
rior fold largest, separated from central fold by a deep 
depression; two anterior fokls lonn a shelf bordering 

short, broad siplional canal. Shell white, periostracum 

Type Material: lIoloty^De MNHN 20505 (14.7 x 9.0 
mm) and 2 paratypes (all dd) MNHN 20506 (largest 
paratype: 17.0 x 9.7 mm). 

Type Loealitv-: Fiji, Lau Ridge, 16°39' S, 179°57' W, 
591-.596 111, [BORDAU 1: sta. 'CP1396]. 

Material Examined: Onlv known Iroin the tvpe ma- 

Distribution: Known oiiK Irom tlie Lau Ridge in Fiji, 
emptv shells in 591-596 ni. 

Etymology: Named after Bruce Marshall (Museum of 
New Zealand, Wellington), in appreciation for the stan- 
dards of his descriptions and illustrations of the moUus- 
ean fauna of New Zealand. 

Remarks: The New Zealand species A. siipcrsfcs (Fin- 
lav, 19.30b) is similar in shape and size, but differs in 
having a translucent white rather than browai protoconch 
with numerous fine, close spiral threads (versus 3 widely 
spaced cords), in having weaker axial eostae on the te- 
leoconcli, in lacking a terminal varix and internal lirations 
ln'hind tile apertural rim at matuiitv, and in that second- 
an' spiral sculpture is considerably weaker or entirely 
absent. Additionally, A. sitpcrstcs has a much more 
strongly developed periostracum, with prominent spines 
at the summits of the axial lamellae. Achnetida siipcrsttcs 
is endemic to northern New Zealand, living on muddy 
substrata in 79-550 m off the noi tlieastcni (northern- 
most record at 35°()8' S) and southwestern North Island. 
A similar species (possiblv a local variant) occurs off 
Three Kings Islands. There is no material of similar spe- 
cies from Norfolk Ridge south oi Norfolk Island in 

Achiichdii lilted new species 
(Figures 14-15) 

Description: Protoconch glassv', of slightlv more than 
one whnil. diameter 975 |JLm, worn but with traces of 
spiral cords visilile on terminal portion. Transition to te- 
leoconch rather indistinct. Teleoconch of about 4.7 
whorls. ^Vhorl profile regularl)' convex; suture im- 
pressed. Sculpture of prominent prosocline axial ribs 
crossed liv weaker spiral cords; about nine to ten a,xial 
ribs on last whorl, eleven on penultimate whorl, some 
forming irregularly placed varices; four priniaiy spiral 
cords on penultimate whorl and about 12 on last whorl, 
with one to four secondan spirals in each interspace; 
spiral cords forming elongate nodes where they cross 
axial rilis. Outer lip with veiy indistinct lirations, visible 
internally at level of terminal varLx, not extending either 
to edge of lip or deeplv within aperture. Parietal area 
with thin callus. Inductura almost vertical. Clolumella 
bearinii three folds; narrow anterior fold forniiim edge of 
short, recun'ed siphonal canal. Exterior vellow-browai . 
Periostracum thin, witli low incrniicnlal lamellae and 
liaiiy projections on spiral cords. 

p. Bouchet and R. E. Petit. 200S 


Type Material: HoIot>|3e MNHN 20507 (13.1 x 7.S 
mm) and one parat\pt" MXIIN 2050S. 

Type Locality-: T(ino;a, 2rM9' S, 175=01' W, 225-233 
m [BORDAU'2: sta. D\\1521]. 

Material Examined: Fiji BORDAU I: sta. ID^^■14fi5, 
18^09' S, 17S'39' W, 290-300 ni, 2 dd.— Tons^a. BOR- 
DAU 2: sta. DW 1521, 2ri9' S, 175°01' W. 225-233 m, 
1 dd (holot^•pe. Fignre 14).— Sta. CP1576, 19°42' S, 
174''1S' \\-, 253-26.3 ni. 1 dd (paratope. Figure 15). 

Ets'iiiology : From the Latin adjecti\e hifciis, meaning 
\elltiw. in reference to the eolor ol the shell. 

Di.stributioii: Known onl\- troni Fiji and Tonga at lo- 
calities eitetl abo\i'. 

• species cutters ironi its congeners 

Remarks: This new species dilt' 
b\' its strongh' prosocline a.\ial ribs forming irregularly 
placed \arices and its spiral sculpture with numerous 
secondan' cords. It is also distinguished from other Acl- 
metida species h\ its \ellow color. 

A specimen ot Admetula Iiitca new specie's m the Petit 
collection (No. 2872) is said to be from 50-200 m north 
of Tiiiwan, but this is a dealer's localit^ designation tliat 
needs to be confirmed. 

Aclinctiihi hatliiptonw new species 
(Figures 16-17) 

Description: Protoconch glassv, white, smooth, of one 
whorl, diameter 975 |jLm. Transition to teleoconch 
marked b\' onset of ;raal ribs and spiral cords. Teleo- 
conch of about four\vhorls. Last whorl bearing about ten 
to 14 rounded axial ribs, crossed by prominent spiral 
cords, about five to sl\ cords on spire whorls and about 
ten on last whorl, with line secondan* spirals in the in- 
terspaces. Final axial rib enlai-ged into a vailx. Small 
nodes formed at intersections whei-e spiral cords cross 
axial ribs. Sutural ramp narrow, almost flat; suture 
slightK' impressed. Outer lip prosocline. rounded, 
smoodi, witliout lirations within. Parietal area without 
wash or callus. Columella slightly inclined, bearing two 
prominent folds extending out to edge of inductnral cal- 
lus; a third descending fold forming edge of short siplio- 
nal canal. No siphonal fasciole. Last whorl well rounded. 
Shell white. 

Type material: Holot)pe MNHN 20509 (7.4 x 5.0 
mm) and one paratxpe .MNHN 20510. 

Type locality-: Norfolk Ridge, south of New Cale- 
donia, 22°52''S, 167°23' E, 59()-600 m [MUSORSTOM 
4: sta. D\\'225]. 

Material examined: Norfolk Ridge. BIOCAL: sta. 
D\V46, 22 = 53' S, 167°17' E, 570-610 m, 1 dd 
(parat\pe).— MUSORSTOM 4: sta. DW225, 22°52' S, 
167°2:3' E, 590-600 m, 1 1\- (holotvpe. Figure 16).— 
SMIB 8: sta DW193-196. 22=.59'-23°00' S, 168°21'- 
168°23' E. 491-558 m. 1 juv. dd.— BATHUS 2: sta. 
D\W20, 22'=52' S, 167°16' E, 530-541 m, 1 juv. dd (Fig- 
ure 17). 

Distribution: known onK hiini the Norfolk Ridge, in 
491-610 m. 

Elyniology: From the Greek Ixitliiis, deep, and no- 
iiuis. place ol lixing, treated as an adjective, 

Remark.s: Adituiulu htillniiiKiiKi resembles a ju\enile 
A. inarsliidii but differs b\' being proportionalK' nar- 
rower, with less con\ex whorls and a larger protoconch 
(chameter 975-1000 |xm versus 825 |JLm in A, nuirshaUi). 
Admctidn inarsJudli also diffei"s in having pointed beads 
rather than small nodes formed at the intersections of the 
axial libs and spiral cords. 

Kolm and Ania (1999: pi. 13, fig. 55) illustrated as C. 
atopodoiita Petit and Harasew^ch, 1986 an 8 mm high 
specimen of Admetula from the Early Pleistocene of \iti 
Le\T.i, Fiji. Their specimen closely resembles the present 
species except that the Fiji fossil has lirations within the 

Genus Fusiaphvra Flalie, 1961 

Type Species: CunccUariu inucrospira .A.dams and 
Reex'e, 1850, fixed herein to complv with ICZN Article 
70.3. Recent, Japan. 

Discussion: Habe (1961a) proposed the genus Fusia- 
plicr/i foi' a species he identified as Conccllaria mac- 
rospira Adams and Ree\e. Later in the same year, he 
(Habe 1961b) considered the species he had identified as 
C. macrospira to be a different species, which he tlien 
named Fusiaplicra macr'ospiratoides Habe, 1961. The 
fact that the type species of Fumaphero was based on a 
misidentified type species was noted by Verhecken 
(1986: 35), M/ho stated that there is some confusion in the 
matter of the type species but did not make a definitive 
statement aliout the resolution of the problem. He did 
list, howex'er, Fuskiphcra macrospira (Adams and Reeve) 
as t)pe species, as did Beu and Max-well (1987; 55), who 
then stated "but misidentified?, Preally = Fiisiaphcra 
macrospiratoidcs Habe, 1961." The contusion is demon- 
strated h\ the fact that Vlatsukiima, Ok-utani and Habe 
(1991: 179, pi. Ill, fig. 2) figured tlie t\pe specimen of 
F. macrospiratoidcs under the name F. macrospira. In 
the absence ot a definitive statement b)- Verhecken and 
the queries used by Beu and Maxwell, it appears to lie 
in'cessan' to fix a type species to comply with Article 
70.3. CanccUaria macrospira Adams and Reeve is here 
selected as the t\pe species of Fiisiajdicra Habe, 1961. 
Fusiaplicra is distinguished by a slender, non- 
umbilicate teleoconch with irregular varices and a thick 
outer lip with a well-defined posterior canal under the 
shoulder. The apeiiure is much like that of Scalptia but 
with a straighter columella. The genus Fusiaplicra ranges 
from South Africa across the Indian Ocean, north to 
Japan, to Australia, New Caledonia and Fiji. It thus ap- 
pears to be restricted to the Indo-Pacific area with the 
earliest known occurrence in the Miocene of Australia. 
Darragh (1970: 168) referred the Au.stralian Miocene 
species CanccUaria cpidromifonnis Tate, 1889 and C. 
cxaltata Tate, 1889, to Fusiaplicra, a placement accepted 

Page S 

THE NAUTILUS, Vol. 122. No. 1 

by Maxwell (1992: 166). However, three New Zealand 
Eocene species, Uxia (?) marshalli Allan, 1926, Uxia 
naronifonnis Finlay, 1930, and Fiisiaphera jenkinsi Max- 
well, 1992, placed in Fusiaphem by Ben and Mawvell 
(1990) and .Maxwell (1992), are not considered by ns to 
be correctly placed. Also, Plesiutriton paijtensis Olsson, 
1930, from the Eocene of Peni, was tentatively placed in 
Fusiaphcra by Beu and Ma\-\vell (1987: 55) bnt it is here 
excluded from the genns. It is possible that a new genus 
will have to be erected to contain these Eocene taxa. 
Many European Tertiaiy species of Unitas resemble Fii- 
siaplwra in foi'm but thev lack a stronglv delineated pos- 
terior canal. 

Fusiaplwra macrospira (Adams and Reeve, 1850) 
(Figures 18-25) 

Cancdlarid mticrospini Adams and Reeve, 1850: 41, pi. 10, 

fig. 2. 
CanccUarid wihncri G. B. Sowerby II, 1881: 6.37, pi. 56, fig. 2. 
Cancellaria pallida E. A. Smith, 1899: 31.3, text-fig. 4. 
Cancellaria producta G. B. Sowerby III, 1903: 220, pi. 4, fig. 5. 
Cancellaria (Trigonostoma) hiscinia Melvill and Standen, 1903: 

319, pi. 23, figs. 14-15. 
Cancellaria excpiisita Preston, 1905: 3, pi. 1, fig. 9. 
Cancellaria iosaensis Habe, 1961a: 72, Appendix 28, pi. 35, 

fm. 21. 

Figures 18-25. I'lisiaplicra inaiid^pira (Ailanis a}id tli'i'\i', 185(1). 18. Height 20.2 mm. I'hilippme.s, Buliea.sug 1., said t(i he Irom 
1.30-230 m. 19. Height 30.1 mm, Japan, Mikawa. 20. nuicnKpiraloidcs form, height 16.2 mm, Mikawa Issiki, Japan. 21. Holotxpe 
o\' Cancellaria nihneri (BMNH 1881.5.20.30), height 1 1.8 mm. 22. losaenda fonri, height 20.4 mm, Minabe, Japan. 23. Height 15.0 
mm, New Caledonia, 19°35' S, 163''25' E, 48 m ["lAGON sta. 1192]. 24, 25. Hnglil' 1 1 .0 mm. New Caledonia, 19°06' S, 'l63°10' 
v.. .50 m |LA(;ON sta. .542|. 

p. Bouehet and R. E. Petit, 2008 

Page 9 

Cancel lurid cizmniii Habe, 19(ila: 72, Apponilix 2.S, pi. 35, liij;. 

C(niccUiiri(i nwcrospiratcndcs Habe, 1961b: 4.33, pi. 23, fi^. 10; 

pi. 24, fig. 10. 
Fiisiaplicrii clampicrcnsis Garrard, 1975: 17, pi. 2, t'ii;;. 8. 
Fiisiaplicra cvii Petit, I9S0: 215, figs. 5, (i. 

Type Data: F. macrospira, China Sea, BMNH 
1969347, lectot^pe designated hy \'erlieeken (19Sfi: 36); 
illustrated bv Higo, Caflonion and Gcito (2001: 99). 

C. wihneri. Port Blair, .AiKLiiiian l.shnKls. Iiolotxpe BMNH 

1S81.5. 20.30, herein Fig. 21. 
C- pallida. 25 fins, oft Bonaparte Archipelago, NW Australia, 

hoIot>pe BMNH 1891.11.21.96. 
C. prDchictti. 40 fins, otf mouth of Uuihloti River, Natal, syii- 

t\pes BMNH 1903.7.27.76; SAM-A339 (Giles and Gos- 

liner, 1983: 28). 
C. (T.) luscinia. 40 fuis, Arabian Sea, 18°58' N, 71°45' E, 

s\iit\pes BMNH 1903.12.15.101-102 (2 .s\iit\pes); NMW 

C, exquisita. Ceylon, hoIot^pe BMNH 1905,10.4.75. 
C. tosaensis. Kochi Prefecture, Shikoku, |apan, holotvpe 

NSMT-Mo 13287; illustnited b\- Higo, Callonion ;uk1 Goto 

(2001: 99). 
C. macrospiratoides. Aiki Prefecture, Honshu, Japan, holotype 

NSMT-Mo 39781; illustrated bv Higo, Calloinon and Goto 

(2001: 99). 
C. aziimai. Aiki Prefecture, Honshu, Japan, NSMT-Mo 13285a 

(illustrated b\' Higo, Callomon and Goto 2001: 99) is la- 
beled as t\pe in .NSMT but is not the figured specimen 

and is smaller than dimensions given for holotvpe. 
F, dampierensis, Delambre Island, Dampier Archipelago, 

nortliwestem Australia, holot)pe WAM 550-71. 
F. cva, west of central Bazanito Island, southern Mozambifjue, 

holot^pe NM G4896. 

Description: Protoconch smooth, glassy, of two 
whorls with small initial nucleus, diameter 1000-1125 
)xm, indicating planktotrophic larval development. Tran- 
sition to teleoconeh marked by shaip axial rib followed by 
additional ribs and cords. Teleoconeh slender, of about 6 
\\'horls. .\xial ribs \ariable in number, 15 to 20 on last 
whorl of most specimens, extending adapically over 
slightlv channeled sutural ramp to suture. Some ribs 
forming varices at irregular intenals, others projecting 
slightK' abo\e ramp. Spiral sculpture of fine cords, about 
15 on last whorl of most specimens, with weaker seeond- 
arv cords in most interspaces. Cords form small nodules 
\\here the\- intersect axial ribs. Aperture narrowly ovate. 
Outer hp thickened into a vara, v\'ith about 14-16 strong 
Hrae that do not descend deeply into aperture. Internal 
hration beneath sutural ramp forms edge of well-defined 
posterior canal. Parietal shield thin but well developed, 
with pustules on its outer edge. Columella with three 
folds, anterior one forming edge of short siphonal canal. 
Last whorl slightlv constricted at base behind weak si- 
phonal fasciole. Exterior brown or white. Many brouii 
specimens with a weak band of white just below periph- 
er\- and white on the adapical ends of ribs and projections 
extending aboxe the shoulder. 

Material E.xamined: Japan. Off Mikawa, 30 fms, 1 
spm (figured bv Abbott and Dance, 1982). — Mikawa Is- 

siki, Aichi Pref., 50-70 m, 1 spm (Figure 20).— Off Min- 
abe, Wakayama Pref., 80-100 m, 2 spms (Figure 22). — 
Off Minabe, 100-200 m, 1 spm.— Off Mikawa, 50 fms, 1 
.spm (Figiu-e 19). — Off Tosa, 50 tms, 1 spm. — "Japan"', 1 
spm. — "Japan?", 1 spm (gift from Habe with "aziimai" in 
his hand). — Mikawa Bay, 50 m, 1 spm. — Off Mikawa, 40 
fins, 1 spm. — Enshu Kei, 2 spms. East China Sea. "180 
m", 1 .spm. Taiwan. Off Keelung, 50-110 m, 2 spms. — 
Off Keelung, 100-200 m, 5 spms.— Off SW Taiwan, 1 
spm. — Off Keelung, "deep water", 1 spm. — Off SW Tai- 
wan, 60 fms, 1 spm. Philippines. Off Aliguay Island, 
Mindanao, "240 m", 2 spms. — Off Aliguay Island, 
"80-120 m", 1 spm.— Off Balicasag Island, "240 m", 1 
spm. — Off Balicasag, "130-230 m", 1 spm (Figure 18). — 
MUSORSTOM 3: sta. DR140, ir43' N, 122°34' E, 93- 
99 m, 1 dd; Sta. CP 141, ir45' N, 122°45' E, 40-44 m, 
1 dd. Vietnam, no locahty, "50 m", 1 ,spm. Indonesia. 
Masalembo, Java, ca. 20 ims, 1 spm. (All above in Petit 
collection). Solomons. SOLOMON 1: sta. DW1760, 
8°47' S, 160°01' E, 172-179 m, 1 dd. Coral Sea. CHAL- 
CAL sta. Dll, 20°3r S, 16r06' E, 83 m, 1 dd. New 
Caledonia. LAGON: sta. 375, 22°32' S, 167°08' E, 67-71 
m, 1 dd; Sta. 517, 19°09' S, 163°35' E, 42 m, 2 dd; Sta. 
542, 10°06' S. 163°10' E, .50 m, 3 Iv (Figures 24-25); Sta. 
1129, 19°29' S, 163°49' E, 40m, 3 Ivi 2 dd; Sta. 1163, 
19°11' S, 163°22' E, 48m, 2 dd; Sta. 1168, 19°16' S, 
163°09' E, 50 m, 1 Iv; Sta. 1192, 19°.35' S, 163°25' E, 
48m, 1 K, 1 dd (Figure 23).— MUSORSTOM 4: sta. 
DW151, 19°07' S, 163°22' E, 200 m, 1 dd. Fiji. SUVA2: 
sta. DW44, Viti Levu, 17°51.7' S, 177°13' E, 33 m, 1 dd. 

Dinicnsion.s: 19.3 x 8.4 nmi (Coral Sea, CHALCAL 
sta. Dll), 17.9 X 8.3 mm (New Caledonia, LAGON sta. 
1192), 14.8 X 7.3 (New Caledonia, LAGON sta. 1192), 
30.2 X 14.0 mm (Japan). 

Distribution: Natal, Mozambique, Arabian Sea, An- 
grias Bank, India, Ceylon, Andaman Islands, northwest 
Australia, Japan (from Izu Peninsula and Yamagnchi 
Pref. southwards), Taiwan, the Philippines, Vietnam, In- 
donesia, Solomon Islands, Queensland, Australia, New 
Caledonia and Fiji. Offshore from 20 to ca. 250 m. 

Remarks: Of the eleven nominal species that we in- 
clude under Ftis'uiphcra imicrospira. t\vo have type lo- 
calities in the southwest Indian Ocean, one in the Ara- 
bian Sea, two in the Bay of Bengal, two off northwestern 
Australia, one off Borneo and three off Japan. We have 
examined representative material from Japan, Taiwan, 
the Philippines and New Caledonia anil fail to recognize 
more than one species. All specimens have in common a 
naticoid multispiral protoconch inchcating planktotrophic 
dexelopment and we inteipret the different names as 
individual rather than geographical variants, connected 
1)\- intermediates. Of these, the nominal species F. nwc- 
wspiratoidcs represents a form where the axial and spiral 
sculpture are of equal prominence, whereas F. tosaowis 
represents a form with much stronger axial sculpture. 
Hasegawa (2000: 585) was of the same opinion when he 

Page 10 

THE NAUTILUS, Vol. 122, No. 1 

stated that "[F. macrospiratoiclcs] and other related spe- 
cies, such as F. aznmai Habe, 1961 and F. tosaensis 
Habe, 1961 mav be intraspecific forms of F. macrospira 
(Adams and Reeve. 1850)." We did not examine ex- 
tensive material fiom the Indian Ocean but published 
descriptions and illustrations suggest that the nominal 
species from this area also fall within the range of x-aria- 
tion of F. inacmsj)ira. Specimens from northwestern 
Australia describecl by Garrard (1975: 17-19) are, how- 
ever, distinctly smaller witli average heights of 10.5 nnn 
to 14 mm. 

Genus NipponapJjcra Habe, 1961 

Type species: Nipponaplwrd Itahci Petit, 1972 by 
ICZN Opinion 1052; Recent, Japan. 

Discussion: Species lA' Nipponaplwra haw tlic angleil 
outline of Triponostoma but are less tabulate, have only 
a small umbilicus if one is present, and ha\'e a different 
cohunellar moi-phologv'. The genus has been utilized pri- 
marily to include species having an angled last whorl, a 
triangular aperture, and three colmncllar folds. Here we 
also place in Nipponaplicra species with a rounded last 
whorl and rounded apeiture, but which are united with 
those taxa trachtionallv included in the genus by the pe- 
cuhar cohunellar moipholog)' consisting of two anterior 
columellar folds situated on a slightK- raised shelf, much 
like a widely bifurcate single fold. 

The genus Mistcia Janssen, 1984, from the Miocene of 
The Netherlands, is similar in outline to the angled spe- 
cies ofNipponap]uTa, but has only t\vo weak folds on the 

Axclclla Petit, 198S (a replacement name for tlie pre- 
occupied Olssonclhi Petit, 1970) has been considered to 
be confined to the Americas (Petit, 1970: 84; 1972: 104). 
What appears to be a neat division of genera between the 
Americas (AxelcUa) and the Indo-Pacific [Nippoiinphcra) 
is blurred b\' the enigmatic species CanccUarut iii^ahna 
Melvill and'Standen, 1901, from the Gulf of Oman, a 
species that appears to possess the characters of Axelella. 
Despite that species, which has not been studied in de- 
tail, it is our opinion that the similarities betvveen Axclclla 
and Nipponaplicra are superficial and that thev can be 
separated b\' their columellar moqihologv". 

Recent species of Nipponaplicra have previously been 
known from South Africa to the northwestern Indian 
Ocean, eastward to the Philippines and north to japan. 
The range is now extended to New Caledonia. In addi- 
tion to the species treated by Bouchet and Petit (2002), 
we now include in Nipponaphera the following: A', scuii- 
pelliicicla (Adams and Reeve, 1850) [described in Can- 
ccUaria: previously placed in Cancellaria by Habe 
(1961b) and other Japanese authors]; N. tcramaclm 
(Habe, 1961) [described in Trigonaphera, placed in 
Scalplia and Trigoiiosfoina bv \arious authors; placed in 
Nipponaplicra by Habe (1961a: pi. 36, fig. 4) on the plate 
caption although in the text it is placed in Trigonaphciri]; 
N. nodosivaricosa (Petuch, 1979) [described in Agatrix 
(Olssonclla), placed in Nipponaplicra by Bouchet and 

Petit (2002)]; N. qiiasiUa (Petit, 1987), new combination 
[described in Cancellaria], N. kastoroac (Verhecken, 
1997) new combination, and N. snclniraiiti (\'erhecken, 
1999) new combination [the last two described in Axe- 

The fossil record has not been completely searched for 
Nipponaplicra but we have recognized Cancellaria 
chinenensis MacNeil, 1961 of the Okinawa Pliocene and 
C. yonabaniensis MacNeil, 1961 of the Okinawa Mio- 
cene as belonging here. Also, Ovama, Hirose and Nish- 
imoto (1995) described the new species Nipponaphera 
tagiichii from the Miocene of Japan and at the same time 
transferred Cancellaria souloi Ilatai, 1941, to the genus 

Nipponaphera (Petuch, 1979) 
(Figures 26-29) 

Agatrix (Ols.'ionella} uoelosiraricosa Petuch, 1979: 11, 
figs. 26, 27. 

Description: Protoconcli pale bi'owai, of about one to 
1.2 wliorls with fine spiral sculpture on final third. Tran- 
sition to teleoconch marked bv prominent axial rib fol- 
lowed by onset of wide, rounded spiral cords and weak, 
poorly defined axial ribs. Teleoconch of 3.5 to 4.2 
rounded \vhorls. Spiral sculpture of 12 to 15 rounded 
primaiy spiral cords, with weaker secondary cords in 
each interspace; about sLx priman cords on penultimate 
whorl. Primaiy and secondan- cords all bear extremely 
fine spiral threads. Axial sculpture of eight to 12 promi- 
nent, elevated ribs on last whorl, more munerous on 
earlier whorls; final one or t\vo ribs becoming wider, 
forming \arices. Spiral cords and interspaces crossed by 
fine, closelv spaced growth lines, giving surface a linen- 
like appearance under low magnification. Suture im- 
pressed, sutural ramp convex. Aperture elongate, 
rounded. Last whorl slightly constricted behind siphonal 
fascicle. Outer lip only shghtlv prosocline, edge thin. In- 
terior of outer lip with ten to 14 strong lirations extend- 
ing deeply into aperture. Stromboid notch manifested by 
veiy slight indentation in outer lip. No parietal caUus; 
some specimens with a thin wash on parietal area. In- 
ductural area cov'cred with thin callus, which extends 
back o\er chink-like umbilicus. Columella with three 
folds, posterior one most prominent, almost pei"pendicu- 
lar to axis; two anterior folds shaiply descending, situated 
on ends of u-sh;iped platform, anterior-most one forming 
edge of short liut well-defined siphonal canal. Exterior 
cream to pale yellow-brown, many specimens with ir- 
regular markings: most specimens with hvo or three 
white spiral cords at periphen' of last whoil, with two or 
tlu'ee dark binwn cords ;ib()\c ;in(l below. 

Type Material: Ilolotvpe (11 x 9 nnn, fide Petuch; 
12.9 X 8.2 uun.fide N'erhecken), DMNH 126397. 

Type Locality: Off Balicasag Islanil, Philippines, from 
.300 111 depth.' 

Material E.xamined: New Caledonia. BATHUS 1: 
sta. D\\'672, 20^48' S, 165°21' E, 347-366 m, 1 Iv (Fig- 

Figures 26-3S. Xippcmaplicrn. 26-29. Sijjpiuuiphcni nodcisiiiiiicosci (Petueli. lUTVJ). 26. Jluiglit Id.iJ nun. Nrw Cakxlunia, 
20°1T' S, 163°50' E, 500-600 m [BATHUS 4 sta. DW89S]. 27. Height 17.0 mm. New Caledonia, 20°48' S, 165°21' E, 347-366 m 
[BATHUS 1 sta. DW672]. 28. Protocoiich, 2r45' S, 166°37' E, 250 m [BATHUS 1 .sta. CP713]. 29. Teleoconch microsculpture, 
same specimen as 28, same scale. 30. Nipponuphcra argo new species, ho!ot)pe, height 8.6 mm; Coral Sea, 22°48' S, 159°24' E, 450 
m [MUSORSTOM 5 sta. 300]. 31-.36. Nipponaplicra c;g( new species 31-32, holot)pe, height 19.3 nmi, Solomon Islands, 9°21' 
S, 160°24' E, 357-359m [SOLOMON 1 sta. CP180(.)]. 33. Height 12.4 mm, Fiji, 19°52"' S, 174°40' VV, 38.3-,393 m [BORDAU 1 sta. 
CP1561]. 34. protoconch, \'anuatu, 20°20' S, 169°49' E, 400-440 m [MUSORSTOM S sta. CP963]. 35. Teleoconch microsculptnre; 
same specimen as 34. same scale. 36. Height 17.9 mm, Philippines, ITOl' N, 124°04' E, 214-246 rn [MUSORSTOM 3 sta. CP145]. 
37-38. Xippoiwphcra tuba new species, holot\pe, height 20.7 mm, Vannatn, 15°10' S, 167^14' E, 394-421 m [MUSORSTOM S: 
sta. CP10S7]. 

Paee 12 

THE NAUTILUS, Vol. 122, No. 1 

ure 27), 1 del— Sta. CP713, 21°45' S, 166°;37' E, 250 m, 
1 Iv (Figures 28-29). BATHUS 2: sta. DW717, 22°44' S, 
167°17^ E, 350-393 m, 1 Iv. BATHUS 4: sta. CP897, 
20°16' S, 163°52' E, 305-350 ni, 1 Iv.— Sta. DW898, 
20°17' S, 163°50' E, 500-600 m, 1 del (Figure. 26).— 
Sta. DW901, 19°03"S, 163°15' E, 297 m, 1 dd.— Sta. 
CP905, 19°02' S, 163°16' E, 294-296 m, 1 Iv.— Solomon 
Islands. SOLOMON 1: sta. CP1801, 9°25' S, 160°26' E, 
264— 273m, 1 Iv. (Dimensions of largest New Caledonia 
specimen: 18.5 x 11.6 mm.) 

Distribution: At this time Nippoiuiphcra nodosiiaii- 
cosa is knovsTi only from New Caledonia, the Solomons 
and the Philippines (Springsteen and Leobrera, 1986; 
Verhecken, 1999). The Indonesian specimen figured as 
Axelello cf nodosivaiicosa by Verhecken (1997: 299, figs. 
5-7) was not attributed by him in his 1999 work to either 
N. nodosivaiicosa or N. suduirauti (see below). In the 
New Caledonian dredgings, live specimens were taken 
from depths of 25(J-393 m and empt)' shells from as deep 
as 600 m. 

Remarks: Verhecken (1999) described tlie species 
AxelcUa suduirauti, here placed in NipponapluTa, distin- 
guished from N. nodosivaiicosa based on protoconch 
characteristics. It was stated that N. suduirauti lias a 
mnltispiral protoconch as opposed to tlie paucispiral pro- 
toconch of N. nodosivaiicosa. Our specimen of N. no- 
dosivaricosa from the Solomon Islands has a protoconch 
that is difficult to attribute to one or the otlier of the two 
species and we believe that the separation between N. 
nodosivaiicosa and N. suduirauti should be reevaluated, 
perhaps using molecular characters. We refer to Ver- 
hecken's (1999) work for his discussion on the question. 

Nipponaj)lu'ra argo new species 
(Figure 30) 

Description: Protoconch smooth, of 1.1 whorls, diam- 
eter 1050 fjim. Teleoconch of about 3.1 whorls, high- 
spired. Whorl profile angulated at shoulder. Sculpture of 
low, broad axial ribs and much finer spiral cords of rather 
even strength, except for one on sutural ramp and two at 
peripheiy of last whorl, which are more prominent than 
others; 14 axial ribs on penultimate whorl, seven on last 
whorl (specimen with severe growth scar and regrowth, 
distorting sculpture of last whorl); about 15 spiral cords 
on penultimate whorl and abf)nt 35 on last whorl, crossed 
bv tliin incremental riblets. Suture shallowly impressed. 
Last whorl slightly constricted behind siphonal fascicle. 
Outer lip thin, shaqi, smooth within, lacking lirae. Inner 
lip with well-developed parietal shield, extending slightly 
over narrow umbilicus. Columella only slightly concave, 
with three folds; anterior two close together, much like 
one large bifurcate fold. Siphonal canal short, indistinct. 
Exterior uniformly veiy pale vellowisli-white. 

Type material: Holotype MNHN 20511 (8.6 x 6.5 
mnil and one paratype MNHN 20512. 

Type Locality: Argo Bank, Coral Sea, 22°48' S, 
159°24' E, 450'm [MUSORSTOM 5, sta. 300]. 

Material Examined: Coral Sea. MUSORSTOM 5, 

sta. 299, 22°48' S, 159°24' E, 360-390 m, 1 dd 
(paratype).— Sta. 300, 22°48' S, 159°24' E, 450 m, 1 dd 
(holotype, Figure 30). 

Etjanologj': Named for the Argo Seamomit, a promi- 
nent topographic feature of the Coral Sea, from which 
tlie specimens were collected; to be treated as a noun in 

Distribution: Coral Sea (Argo Seamount), dead in 
390-450 111. 

Remarks: Nippoiuiphcra argo differs from N. goniata 
Bouchet and Petit, 2002 by its sculpture of broad, low, 
non-lamellar axial ribs. Also, the spiral cords of N. argo 
are of more even strength, except for one on the shoulder 
and two on the peripheiy, which imparts a slightly bian- 
gular aspect to shells of this species. 

Nipponaphcra agastor new species 
(Figures 31-36) 

Description: Protoconch iioriiialK smooth, corroded 
on holotyiJe, of 0.9 whorls, diameter 800 jjim. Proto- 
conclVteleoconch boundaiy indistinct due to corrosion, 
but distinctly marked by onset of teleoconch sculpture on 
specimens from Vanuatu. Teleoconch of five rounded 
whorls; spire angle 64°; suture deeply impressed. Axial 
sculpture of prominent, regularly spaced ribs, eleven on 
last whorl, eleven on penultimate whorl. Ribs rounded 
over steep, narrow sutural ramp. Spiral sculpture of 
evenly spaced, prominent spiral cords, eight on penulti- 
mate whorl, 13 on last whorl, witli three to five secomlaiy 
cords in each interspace; spiral cords rise over axial ribs, 
forming small nodules on primaiy cords at intersections. 
Numerous fine growth lines cross spiral cords, creating 
small imbrications. Last whorl slightly constricted behind 
siphonal fasciole. Outer lip prosoeline. Inner margin of 
lip smooth apart from 16 prominent lirae extending into 
aperture; two indistinct lirae on parietal area. Columellar 
callus well developed, forming shield over chink-like um- 
bilicus. Columella with three almost equal folds; anterior 
one sloping sharpK down at edge of small but distinct 
siphonal canal, wliicli recurves abaxially. Exterior chalky 

Type Material: I lolotvpe 20513 ( 19.3 x 1 2.3 mm) and 

4 parat\]X's MNHN 20.514. 

Type Locality: Hclween Gu:idalcaiial and Florida Is- 
land, Solomon Islands, 9°21' S, 160°24' E, 357-.359 m 

Material Examined: Vanuatu. MUSORSTOM 8: sta. 
CP963, 20°20' S, 169°49' E, 400-440 m, 1 Iv (Figures 
34_35),_Tonga. BORDAU 2: sta. CP 1561, 19°52' S, 
174°40' W, 383-393 ni, 1 ild (Figure 33).— Solomons. 
SOLOMON 1: sta. CP1746, 09°23' S, 159°57' E, 302- 
396 m, f dd; Sta. CPf800, 9°21' S, 160°24' E, 357- 

p. Bouchet and R. E. Petit, 20()(S 

Paee 13 

.359ni. 4 1\ , 1 (111 (holotxpe. Figures 31-32, and 
paratxpe.s),— Philippine.s. MUSORStOM 3: .sta. CP145, 
ll"()i' N, 124°04' E, 214-246 m, 1 dd (Figure 36).— 
Piinta Eiigano, tangle net.s, approxiniateh' 60 Inis, 1 ,spni. 

EtAinologA-: From the Greek agastor. a noun in appo- 
sition, meaning near kinsman or brother, to highlight the 
similarit\' to .Y. iiodosivarico.'ia. 

Di.stribiition: OiiK known Iroiii the material exam- 
ined: Philippines, Solomons, X'anuatu and Tonga. Depth 
range in the southwest Paeifie 360-41)0 ni; in the Phihp- 
pines possihK' shallower. 

Remarks: Specimens from the Philippines and the So- 
louions are ihstinetK" larger with adult sizes ranging trom 
16.8 mm to 19.5 nun, whereas specimens from Vanuatu 
and Tonga are nuich smaller with adult sizes at 12.2 mm 
and 12.3 mm respectixeh'. 

Nippoiiaplicra i/oa.stor is sxinpatric with A', nodosivari- 
cosa in the Philippines and the Solomon Islands but dii- 
fers bv its spiral sculpture with more numerous (3 to 5) 
secondaiv cords, not separated by an incised groove as in 
A', noclosivaiicoso. It ilifiers trom N. tuba, with which it 
is s\nnpatiic in \'annatu, b\- being more slender ami lack- 
ing a stromboid notch. 

Xipponaphcro tuba new species 
(Figin-es 37-.38) 

Description: Protoconch smooth, of 0.9 whorls, diam- 
eter SOO |jLm. Protoconcli/teleoconch boundary' distinctly 
marked bv onset of teleoconch sculpture. Teleoconch of 
fi\"e rounded whorls: spire angle 75°; sutin-e deeply im- 
pressed. Axial sculpture ot prominent, regvdarly spaced 
ribs, ten on last win)rl, 14 on penultimate whorl; ribs 
rounded on steep, narrow sutui'al ramp. Spiral sculptiu'e 
of e\"enlv spaced cords oi se\eral strengths; priman- cords 
e\enl\- separated b\- one secondaiy cord, resulting inter- 
spaces fOled with t^vo or three tertiary cords; spiral cords 
rising over ;rxial ribs, forming small nodules on primary 
spiral cords at intersections. Numerous fine growth hues 
cross spiral cords, creating small imbrications. Last whorl 
shghth" constricted behind siphonal fasciole. Outer lip 
prosocUne. with distinct stromboid notch adapical to cen- 
ter of lip. Inner margin of lip smooth or slightK^ crenu- 
late, with 19 prominent lirae extending into aperture, 
four adchtional lirae beneath sutural ramp. Columellar 
callus well dexeloped. forming shield over chink-like um- 
bilicus. Columella with three almost equal folds, anterior 
one sloping shaiplv down at edge of small but distinct 
siphonal canal, which recunes abaxially. Exterior yellow- 
browii with a band oi white below^ peripheiy, bordered 
b\' indistinct bands of darker lirown. 

T%pe Material: Holotxpe MNHX 20516 (20.7 x 15.0 
mm) and one parat\pe VIXHX 20517. 

T\pe Localifrv-: \'anuatu, 15n()' S, 167°14' E, 394- 
421 m [MUS(5r,ST0M S: .sta. CP1()S7]. 

Material Examined: \ anuatu. MUSORSTOM S: sta. 
CP1US7. 15=10' S. 167°14' E. .394-421 m, 1 Iv, 1 dd 

(liolot\pe. Figures 37-3S, and paratxpe). — Sta. [no data, 
mixed lot], 1 dd. 

Etymology: From the Latin tuba, a war trumpet, 
which this new species can, with some imagination, be 
reminiscent ol; used as a noun in apposition. 

Distribution: Kuown ouK Iroin N'anuatu. 

Remarks: Nipponapltcra tuba diiters trom N. ci/- 
plioiiiii liouchet and Petit, 2002 in being more rounded 
and robust. Also, the spiral sculpture never appears as 
incised lines as in .V, ci/pli(uiia and A', nodosivaricoaa. 
Nipponaphera luba also has a thick outer lip that is not 
present in N. ciiphoiua. 

Genus Trigonostoina l^laimille. LS27 

Trifj^ona Periy, 1811: pi. 51, T\pe .species: Trigona pellucida 
Pern-, 1811, by monotypy. Not Trigona Jurine, 1807 (Hv- 

Trigonostoma Blainviile, 1827: 652. Tvpe species: Dclphiiiiila 
trigonostoma Lamarck, 1822 (?= Bucciniim sailarc Gnie- 
lin, 1791), b\' nionot\p\'. Recent, Indo-Pacific. 

Remarks: Trigoiuistoina has a ditferent taxonomic 
composition for various authors. We liere use Trigono- 
stoma sensu Into to encompass the nominal genera Ven- 
tiilia Jousseaume, 1887, Arizelostoma Iredale, 1936, 
Ovilia jonsseame, 1887, and Extractrix Korobkov, 1955. 
We do not include Scalptia Jousseaume, 1S87, Trigona- 
pliera Iredale, 1936 and Cancellaphera Iredale, 1930, 
which are sometimes treated as subgenera of Trigono- 

Trigdiio.stiniia Injblium new species 
(Figures 39-44)" 

Description: Protoconch smooth, glassy, of one whorl, 
dianu'tci- 1050 \l\\\. Transition trom protoconch to teleo- 
conch abiTipt, denoted by onset ot axial and spiral sculp- 
ture. Teleoconch of 2.5 whorls; spire depressed; umbili- 
cus broad. First teleoconch whorl with 16 axial ribs. Sec- 
ond whorl with about IS axial ribs that ha\e become 
somewhat obsolete. Spiral sculpture of broad, closely 
spaced cords witli narrow interspaces; about seven to ten 
spiral cords on sutural ramp and 25 between shoulder 
angle and umbilical rim. Sutural ramp weakly concave, 
but forming deeplv channeled shoulder between shoul- 
der angle and impressed suture. Apertin"e narrowh' 
ovate. Outer lip smooth. Inner lip forming parietal 
shield, partly covering deep, wide umbilicus. Columella 
with two descending folds. Anterior canal not con- 
stricted. Holot\pe exterior chalk)' white, with two broad, 
ill-defined browni bands, best seen througli shell. 

Type Material: Holotxpe MNHN 20518 (7.0 x 6.5 

Type Locality-: North of Makira Island. Solomon Is- 
lands, 1()°13' S, 16r29' E, 381-383 m [SOLOMON 1: 
sta. CPLS37]. 

Material Examined: Taiwan. TAIWAN 2000, sta. 
13W36. 2P'54.8 X, 120"36.2 E, 305 m, Bashi Channel. 1 

Page 14 

THE NAUTILUS, Vol. 122, N( 

Figures 39-47. Trii^onosloiiKi. 39—44. Tiigoitustitnui tn/hliiiiii new species. 39. Height H.3 nini. Taiwan. 15aslii Channel 21°54.S' 
\, 120"36.2' E, 305 m [TAIWAN 2()()() sta. d\\'36|. 40-41. Holohpe, heiglit 7.0 mm, Solomon Islands, I()"I3' S, I6I°29' E, 381-383 
111 [SOLO.VION I sta. CPI837]. 42. Height 5.0 mm; New Caledonia, 23°03' S, I66°58' E, 397-400 m [BATHUS 2, sta. D\V730]. 
4.3-44. Protoconcli and teleoconch microsculpture, same specimen as 42. 45-47. Trii^diioslomii tlu/stlilnii Petit and Harasew>'ch, 
1987. 45. Height 16.9 mm. New Caledonia, 22°47' S, 167°28' E, 241-245 m [BATHUS 2 sta. CP728]. 46. Height 15.0 mm, Fiji, 
16°50.4' S, 178°12.5' E, 200-215 m [MUSORSTOM 10: sta. D\\T333]. 47. Protoconeh, New Caledonia, 22°43''s. 167°16' E, 300 
m [SMIB 1 sta. DW6]. 

dd (Figure 39). Solomons. SOLOMON 1: sta. CP1837, 
10°13' S, 16r29' E, 381-383 m, 1 Iv (holotyioe. Figures 
40-41). New Caledonia. BIOCAL Sta. DW77. 22°15' S, 
167°15' E, 440 m, 1 juv. Iv.— BATHUS 2, sta. DW730, 
23^03' S, 166°58' E, 397-100 m, 1 ju\-. dd (Figures 42-44). 

Etj'inology: From the Latin tn/hliiiiiL incaiiing cup, a 
shape that the new species is SDiiuwhal rcmiiiiscrnt iil; 
used as a noun in ajijiosition. 

Di.strihiition: kiidwii oiiK' troiii Tai 


New Cale- 

p. Bouchet ami H. E. Petit, 2008 

Page 15 

donia ami the SoloiiiDiis. Ali\e in 383-440 iii; eiiipt) 
shells from 305 in. 

Remarks: ()1 tlie knowni Recent species of Trigono- 
■stoina, Trigoiio.sioiiia tn/bliuni resembles onlv T. seinicl- 
isjiincta (Sowerbw 1S49) in ha\ing primariK' spiral sculp- 
ture. It differs from that species in being more de- 
pressed, with a broader umbilicus. In T. scmkhisjnncta 
there is no spiral sculpture on the sutural ramp. .\lso, tin- 
spiral sculpture ot'T. seinidisjnncta is arranged in groups 
of cords separated hv wide fin-rows. 

Tiigonostomci tlii/sthloji Petit and Harase^\'^■ch, 1987 
(Figures 45-47) 

Trigonostoiud tlii/.stlilon Petit and Harasew\'ch, 1987: 79, figs. 5, 

Trigonostoma ant'uinalii — Ilabe, 1961a: 435, pi. 24, tig. 14; pi. 
23, fig. S; 19(Slb: 73, pi. 36, fig. S: Lan, 1979; 9.5, pi. 41, 
figs. 93, 93a; Abbott and Dance, 1982: 299 (second figure 
in bottom row); Habe and Okutani, 1985; 233 (second 
figure in bottom row); Bosch, et al., 1995: 157. fig. 687. 
[not Cancellaria nntiqunta Hinds, 1843] 

Trigonostoma antiquatum — Ovama and Takemura. 1963: 
Trigonaphero (2) plate, fig. 5. [not CanccUiirki (intitjudid 
Hinds, 1843] 

Trigonostami (sic) tlu/sthlon — Hasegawa, 2000: 581, pi. 290, 
fig. 11. 

T^pe Material: Holot^pe, USNM 747301 (17.3 x 12.7 

T\pe Locality': Off west coast of Wasir Island, West 
\Vokam, Aru, Moluccas (5°30' S, 134°12' E) in 56-73 m. 

Material Examined: New Caledonia. LAGON: sta. 
387, 22°39' S, I67°07' E, 225 m, 1 dd.— SMIB 1: sta. 
DW6, 22°43' S, 167°16' E, 300 m, 1 dd (Figure 47).— 
BATHUS 2: sta. CP728, 22°47' S, 167°2S' E, 241-245 
m, 1 1\- (Figure 45).— Dredged at "300-400 m". off 
Belep, north New Caledonia, 1 spm. Fiji. MUSOR- 
STOM 10:sta. D\V1333, I6°50.4' S, 178°12.5' E, 200- 
215 m, 1 dd (Figiu-e 46). Solomons. SOLOMON 1: sta. 
D\MS50, 10°28' S, 161°.59' E. 1.39-261 m, 1 Iv. 

Description (of a specimen from New Caledo- 
nia): Protoconch smooth, glassy, of 1.8 whorls, diam- 
eter 1050 Jim. Transition to teleoconch marked by axial 
rib followed by both axial and spiral sculpture and flat- 
tening of the sutural ramp. Teleoconch of up to sb; tabu- 
late whorls. Sutural ramp flat, bordered b\' cord-like 
shoulder angle. Suture impressed. Axial ribs prominent, 
extending from suture across ramp, over shoulder angle, 
where the\' form recuned spines on manv specimens, 
dowii and o\"er siphonal fasciole, then inside umbilicus. 
Shoulder spines not formed on al! ribs, but man\' ribs 
lacking spines form short, sloping buttress against pre- 
ceding whorl; about 12-15 ribs on last whorl, more nu- 
merous on earUer whorls; two thick, closely spaced ribs 
mark end of growth in adults; a few vailx-like rib occurs 
earher on some specimens. Fine, closely packed growth 
hues of \'aning number in axial interspaces sHghtly over- 
lap each other, producing a scabrous appearance. Spiral 

sculpture of veiy closely spaced cords, rising over ribs to 
form small nodes. Sutural ramp with about 12-15 spiral 
cords of uniform prominence; about 15 primary spiral 
cords on last whorl bct\veen shoulder angle and siphonal 
fasciole; about ten fine secondan' cords in each inter- 
space. Spiral cords of equal strength continue inside um- 
bilicus. Outer lip slightly prosocline. Outer lip thickened 
h\ \-ari,\, slightlv serrate on margin, interior with eight to 
ten short irregular lirae, not extending (juite to outer 
edge of lip and not extending deeplv into aperture. Ap- 
erture triangulai', witli small postericir notch under su- 
tural ramp. Posterior p(jrtion of inner lip adpressed 
against siphonal fasciole and anterior quarter of penulti- 
mate whorl. Inner lip with two descending folds, poste- 
rior one slightly larger than anterior; third incipient fold 
present in some specimens. Umbilicus extends to proto- 
conch. Siphonal canal sliort. Exterior white with ven' 
faint orange-brown at shouldei'. Specimens over 24 nun 
in htight are known. 

Distribution: Ciull ol Oman to the Moluccas, Japan, 
the Philippines, Solomon Islands, New Caledonia and 

Remarks: Tlic mibricati' sculpfiu'e resulting iioin the 
overlap of growtli lines, which is so noticeable on the 
New Caledonian specimens, is absent from T. antiqua- 
tum and is much less evident on T. tliijstldon from other 
areas. However, we do not consider this difference to be 
of taxonomic significance. This sculpture is a prominent 
feature of T. sailair (Gmelin, 1791), the ty^^e of the 
genus but that species has a more angular shape and its 
whorls are barely attached. 


We thank Bertrand Richer de Forges, the indefatigable 
companion of the first author on many expeditions, for 
his skill and determination in exploring the deep-water 
bendios of die tropical Pacific. Ahmed Abdou did SEM 
illustrations of protoconchs and micr(jsculptm"e; Del- 
phine Brabant and Philippe Maestrati did the digital 
photographx- and plate assembling. Bruce Marshall 
helped us inteipret our material of Achnvtula from Fiji. 
Alan Beu helped us iinpro\'e the manuscript b\- tracking 
internal inconsistencies. 


Abbott, R. T. and ,S, P. Dance. 1982. Conipendnnn olSeasliells. 
E. P. Dutton, Inc., New York, L\ + 411 pp. [2'"' printing, 
revised, 1983; 3"' printing, revised, 1986; see also Habe 
and Ohitani, 1985]. 

Adams, A. and L. Reeve. 1848-50. Mollusca. In: A. Adams (ed.) 
The zoology of the voyage of H.M.S, Samarang; under the 
command of Captain Sir Edward Belcher, C.B., F.R.A.S., 
F.G.S. during the years 1843-1846. Reeve, Benham. and 
Reeve, London, x + \-\' + 87 pp., pis. 1-24. [i-x (i-ii plate 
ex-planations to pis. 1-9; iii-x Introduction to Mollusca by 
Adams only), 1-24, pis. 1-8, 1 No\ ember 1848; 2.5-14, pis. 

Pase 16 

THE NAUTILUS, Vol. 122, No. 1 

10-17, 27 April 1850; i-,w, 45-87, pis. 18-24, 31 August 

.Allan, R. S. 1926. Fossil Mollusca f'roiu the M'aihao Green- 
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THE NAUTILUS 122(1):19-51, 20()S 

Pao-e 19 

On some Neogene to Recent species related to Galcodina 
Monterosato, 18cS4, Galcodinopsis Sacco, 1895, and Massotia 
Bncqnov, Dantzenberg, and Dolllns, 1884 (Caenogastropoda: 
Rissoidae) with the description of two new Alva nia species from 
the Mediterranean Pleistocene 

\'ittorio Garilli 

APEMA Research and Kdueatioiial Senice 
\'ia Alia F;ilconara. 34 
1-90136 Palermo, ITALY 


SLx species, related to the subgenera Galcodina, Galeoclinop.sis, 
and Massotia. are re-anaK'zed. Ahaiua francescoi new species 
iSE SiciK'l and A. rosariac new species (SE Sicilv and NW 
Peloponnesus) are described from Mediterranean Plei.stocene. 
Galcoclinopsis is regarded as the appropriate genus for Oli- 
gocene-Recent taxa ha\'ing a quite conical shell close to that of 
some Alvania species and showing Manzoma-\]ke combination 
of two microsculptiu'al characters; the pitted surface on the 
spiral cords and the arrangement of the roughK' prismatic ele- 
ments forming nvnnerous and ven- fine spiral threads. Its t^'pe 
species, Rissoa tibcriana (previously known from Mediterra- 
nean Mio-Pliocene), li\es along the tropical \\' African coasts, 
where it is kno^ni under the name A. fariai. As suggested bv the 
oldest record oiGalcodinopsis, the European Oligocene Rissoa 
diiboisi, tliis genus \er)' likely originated from a pre-Neogene 
Ahania group. The svntspe of Rissoa pnisi, a scarcely known 
species (Pleistocene of Rhodes), and material (also from type 
locality') of the almost unknown jR. cingnlata (from Sicily) and 
of its close relatixe, R. tenera (Mediterranean, Atlantic Moroc- 
co and Canan' Islands), is sho\\^l. These three taxa and the t\'pe 
species of Galcodina and Massotia, are here tentatively con- 
sidered as belonging to Ahania sensu lato. With the exception 
of A. cinoulata, all die discussed .species ha\e a multispiral 
protoconch. Generally, the protoconchs studied exhibit a sculp- 
tural pattern known in odier rissoid taxa. Protoconch I of the 
txpe species oi Massotia , A. lactca, is characterized b\' a coarser 

Additional Keywords: Rissoidae, taxonom\', VIediterranean-W 
.Atlantic, Xeogene-Recent, new species 


The genus Aliania Risso, 1826, comprises one of the 
most di\-ersified groups in the caenogastropod famil\- 
Rissoidae, especialK" when considering the tropical east- 

ern Atlantic and the MecUterranean provinces. Thev in- 
habit a large variety' ot en\ ironirients, Irom littoral to 
l)ath\al, and their geographical tlistribution is extensi\e, 
including the VIediterranean, Atlantic, Indo-Pacific, and 
the temperate Australian coasts (Ponder, 1985). Their 
stratigraphical distribution might extend back to the Late 
Cretaceous, but the first well-estabHshed records date to 
tlie early Tertian' (Ponder, 1985). With regard to the rich 
Mediterranean and Fluropean Tertian' Alvania assem- 
Ijlages, some of the most informative analyses are those 
of Sacco (1895), Seguenza (1903), Cossmann (1921), Lo- 
zouet (1998), Kowalke and Harzhauser (2(:)()4), and Chirli 

Many authorities, e.g. Monterosato (1SS4), Bucquoy et 
al. (1884), Weinkauff (1885), Kobelt (1888), Locard 
(1886), Nordsieck (1968, 1972), Jeffreys (1867, 1869), 
Pallaiy (1920), Wenz (1938), War^n (1973, 1974), Gofas 
and Waren (1982), Van Aartsen (1982a, 1982b), Moolen- 
beek and Hoenselaar (1989. 1998), Van der Linden and 
Wagner (1989), Moolenbeek et al. (1991), Van der Lin- 
den (1993), Bouchet and Waren (1993), Giannuzzi- 
Savelli et al. (1996), Palazzi (1997), Gofi^s (1999), A\ila 
(2000). and Arduino and Arduino (2001), ha\'e contrib- 
uted to the kmowledge of the rich Recent Eastern Atlan- 
tic (especially the W Africa and the Macaronesian Prov- 
ince) and Mediterranean fauna. Ponder's rissoid re\ision 
(1985), listing ine ,\k'riiua subgenera, provided addi- 
tional perspective to the taxonomy. 

Vlv attention is focused here on a relatively large, in- 
lormal group of species oi Ahania sensu lato character- 
ized by shells with wide and ovate aperture, lacking an 
internal denticulation of the outer lip, and otten bearing 
varices on a well-developed, inflated body whorl. These 
species have been historically assigned to the subgenera 
Galcodina Monterosato, 1884, Galcodinopsis Sacco, 
1895, and Massotia Bucquoy et al, 1884. This subgeneric 
settlement was partially rejected by Ponder (1985), who. 

Page 20 

THE NAUTILUS. X'ol. 122, No. 1 

modifying the .systematic arrangement of Monterosato 
(1884) and Bucquo\- et al. (18S4X stated that grouping of 
the numerous species oi Ahaiiia was \'en' difficult at the 
subgenus le\el. The same author inchided Massotia (t)iDe 
species: Ri.ssoa lactea Michaud, 1830) and Galeodinopsis 
(tvpe species: R. tiberiano Coppi, 1876) in the Alvania 
sensu stricto group and doubtfully considered Goleodina 
(t\'pe species Turbo carinatiis Da Costa, 1778) as a valid 
subgenus on the basis of shell characters. Piani (1979) 
raised Galeaclina to generic level and placed Ri.ssoa cin- 
gulata Philippi, 1836, and R. tenera Philippi, 1844, 

The principal aim ol this work is to proxide, tor the 
first time, as complete as possible a taxonomic dataset 
based on shell features. Species such as Rissoa cingulata, 
[often misidentified as Alvania carinafa (Da Costa, 
1778)], R. pnisi Fischer, 1877. and R. tibcriana Coppi, 
1876, are very poorly known or, in the case of the last 
taxon, the generic attribution to Alvania appears incor- 
rect. I also describe two new species from the Pleisto- 
cene of the central Mediterranean area. 


Most of the examined material, all consisting of shells, is 
housed in the Museum National d'Histoire Naturelle, 
Paris (Departement S\stematique et Evolution), in the 
private collections of Maurizio Forli (Prato, Italy) and of 
Stefano Palazzi (Modena, Italy). Remaining material is 
housed in the private collections (made 
a\'ailable bv Nino Adorni Sbrana, Grosseto, Italv), Ste- 
fano Rufini (Anguillara), in the Museum f'iir Naturkunde 
(Humboldt University, Berlin), the Museo Geologico 
G.G. Gennnellaro and the Dipartimento di Geologia e 
Geodesia (both Uni\ersit\' of Palermo, Italv), in the Gou- 
landris Natvu'al Histon- Museum (Kifissia, Athens), the 
Dipartimento di Scienze della Terra of the University of 
Catania, Italy, the Zoologisch Museum of Amsteixlam, 
the Museum National d'Histoire Naturelle of Paris (De- 
partement Histoire de la Terre), and in my personal col- 
lection. Many other private collections were visited. 

In the parts dedicated to each species, a list f)f the 
material is provided with all information given in the 
original labels. An abbreviated list of essential s\'non\'my 
and/or citations is provided. 

The fossil material (all from the Mediterranean area), 
from the deposits of Dattilo (NW Sicily), Cartiera 
Mulino (Vittoria, SE Sicily, type locality of Alvania 
francescoi new species and Alvania rosaiiae new spe- 
cies), and Kyllini (Elea, NW Peloponnesus, Greece), was 
obtained by washing liulk samples on a serial sieves (0.5, 
1, and 2 nnri square meshes) and sorted using a stereo- 
microscope. The same procedure was followed for the 
Recent bulk samples collected from Vlagnisi (Siracusa, 
SE Sicily, the type localits' of Rissoa ciiigulafa J'liilipjii. 
1836) and Mondello (Palermo, NW Sicily) specificall) to 

recover that species. Geological, stratigraphic and paleo- 
ecological information on the deposits of Cartiera 
Mulino, Dattilo and Kyllini are taken from Costa (1989), 
Garilli (1998; 2004), Garilli et al. (20()5a; 2005b) and 
Garilli and Galletti (2007). The stratigraphic information 
on the Sicilian deposits of Birgi (Trapani) and Tonnnaso 
Natale (Palermo, originally attributed to the upper 
Pleistocene ThyiTenian by Ruggieri and Milone, 1973) 
are from Ruggieri and Unti (1988) and Heaity et al. 
(1986), respectively. For the remaining fossil ma- 
terial (mainly from Coll. Forli c.v-coll. Palazzi), I followed 
the stratigraphic attributions reported bv the col- 

For all the discussed species, at least five shells were 
studied by the scanning electron microscope (SEM) us- 
ing a Philips XL3() ESEM, except for the (sole) syntyjDe 
of Rissoa pnisi and R. cingulata, of which there were 
only three shells available. Specimens examined by SEM 
were cleaned in a Bransonic 5 ultrasonic machine using 
distilled water. Particular attention was given to proto- 
conchs and teleoconch microsculptures as potential 
sources of taxonomic characters at species level. The 
mnnber of protoconch whorls were counted according to 
Verduin's method (1977). 

Shells were measured using a stereo microscope pro- 
vided with a cross-line micrometer eyepiece. The posi- 
tion of any varix on the body whorl is indicated in degrees 
of the angles formed h\ the plane of the \An\ and of the 
outer lip. 

Geographic, bathvmetric, and stratigraphic distribu- 
tion of each discussed species is based on the examined 
material and the literature. Published records were criti- 
cally evaluated on the basis of good illustrations or sat- 
isfacton' descriptions. This t\pe of dataset allows for just 
a rough representation of the geographical distribution, 
especially for the Adriatic Sea and the eastenrmost Medi- 
terranean basin, of which I found veiy little material 
from the collections studied. 

Abbre\dations are used as following: DGUP: Diparti- 
mento di Geologia e Geodesia. LTniversita di Palermo, 
Itah'; DSTC: Dipartimento di Scienze della Terra, Uni- 
versita di Catania, Italv; GNHVI: Goulandris Natural 
Histoiy Museum, Kifissia, Athens, Greece; MGUP: Mu- 
seo Geologico G.G. Gemmellaro, Universita ch Palermo, 
Italy; MNHN-DHT: Museum National d'Histoire Na- 
turelle, Departement Histoire de la Terre, Paris, France; 
MNHN-DSE: Museum National d'Histoire Naturelle, 
Departement S)stematique et Evolution, Paris, France; 
MPOB: Dipartimento del Museo di Paleobiologia e 
deirOrto Botanico, Universita di Modena e Reggio 
Emilia, Modena, Italy; MSNCS; Museo Regionale di 
Storia Naturale e Mostra Pernianente de! Carretto Sicil- 
iano, Terrasini, Italy; ZMA: Zoologisch Museum Amster- 
dam, Holland; ZMB: Museum fiir Naturkomde, Hmn- 
boldt Universittit, Berlin, Germanv; Coll.: collection; 
Coll. MF: Maurizio Forli collection, Pi'ato. ItaK'; Coll. 
P.'VL: Stefano Palazzi collection, Modena, Italy; Sh(s): 
.shell(s), used in the Material Examined sections only. 

V. GariUi, 2008 

Pa^e 21 


FamiK' Rissoidae C.raw 1847 
SubtaniiK" Rissoiiiac C>vav, 1847 
Genus Alvania Risso, 1826 

Tj'pe Species: Ahiinia ciiropcn Risso, I82(i [smioiinui 
of A. cimcx ^Linnaeus, 1758)], suhseijuent designation bv 
Ne^^ll, 1885 

Ahania caritiata (Da Costa, 1778) 

(Figin-es 1-14) ^, 

Turbo ctniiuitiis Da Costa, 177S; 1()2-1()3. pi. S, li^. 10 
Risson trocJiIca Midland. 18.30: 16, iig. 4 
Galeodiim cingulata (Philippi, 1836). — Piani, 1979: 70-71, iigs. 

Description: Shell small, sturdy, conical and keeled to 
slender and turrited, reaching about 5 nun (rarely 6 mm) 
in height; 3.9 — 1.1 nun in width. Protoi'oni. 1 1 multispiral, 
conical, consisting ot about 2.2 con\'e,\ whorls. Proto- 
conch I \\ith O.S whorls, sculptured with sl\ veiy line 
spinJ lirae and microscopic granules between them. Pi-o- 
toconcli/teleoconch transition well-marked and sinuous. 
Protoconch II sculptured \w\i\\ sparse, spirally arranged 
microscopic pimples, stronger in adapical direction, and 
one to t^\■o spiral ridges, one of them always veiy close to 
lower suture. In the largest shells, teleoconch formed by 
4—4.5 moderateK' convex whorls. Common moqjh (Fig- 
ures 2, 6-7) wth teleoconch whorls markedly dominated 
b\" spiral sculpture, which consists of veiv strong cords 
(numbering 2-.3, 3-6, 6-12, and 12-17 on first, second, 
third, and last teleoconch whorl, i^espectivelv). Second- 
aiT, less conspicuous cords may occurr on last whorl. 
More marked spiial cords on adapical portion of wliorls, 
at a certain distance from suture, give a characteristic 
keeled shape. Cords progressively less strong on basal 
area. Unkeeled moi-ph (Figures 1, 4-5) characterized by 
a slender shell shape, usually bears more spiral cords 
(18-20) on last ^^'horl. Axial sculpture alwa\s formed by 
numerous (32-50 on penultimate whorl), occasionally 
\er\ narrow and lamella-like ribs, becoming obsolete to- 
ward base. Intersection of ribs with spiral cords gives an 
almost general clathrate pattern in mikeeled moiph. In- 
tersection of spiral and axial sculptures nodular, usually 
forming squares (Figure 14), with exception of last whorl 
\\-here a rectangular pattern occurs (Figure 13). Micro- 
sculptiu'e consists of veiy fine spiral tlireads (Figures 
13-14). coxering all teleoconch surface, with exception of 
main spiral pattern. On earlv teleoconch \\'horls, spiral 
hrae often alternate with spiral alignments of micro- 
scopic pimples. Sutures slightly inclined. Last whorl well 
expanded, comprising .3/5 to 3/4 (rarely more than 3/4, 
Figure 3) of shell height, often bearing one or two varices 
(mainK" in keeled moi-ph) with angles of 10°-340°. Ap- 
erture wide, o\"ate, sliglitly rounded to angled in the pos- 
terior part, comprising .3/5 to 3/4 of last whorl height. 
Outer lip slightK" prosocline, internally smooth, exter- 
nall\- markedK- thickened by a strong rim veiy close to hp 
edge, and co\ered b}- spiral cords. Inner lip moderately 

arcuate and lathei' thickened, with a \e\y narrow to dis- 
creteJN' expanded (Figure 3) callus delimiting a xcn- small 
umbilical chink. 

TApc Locality: (Cornwall, southwestern England. 

Material L.vaniincd: (Jreat Britain: Cornwall, Fal- 
moutii, 1 sli., cull. MF, 1974, E55A; Channel Islands, 
Henii, 11 shs, coll. MF, 09.1974, E54A. Atlantic France: 
Normandy, Carteret, 3 shs, coll. MF, Jul, 1973, E28A; 
lower Normandv, St. Pair, 7 shs, MNIIN coll. Denis, 
1945; Brittany, Finistere Anse de Bertheaume, 20-30 m, 
industrial dredging, 4 shs, MNHN coll. S. Gofas, 1978; 
Brittany, Cote-du-Nord Plomanac'h, fissures of infralit- 
toral rocks, 1 sh., MNHN coll. S. Gofas, 1973-78; Brit- 
tany, Finistere Roscoff, "les Coehons Noirs", sand and 
conchiferous gravel, 20 m, 27 shs, MNHN coll. Gofas, 
Jul. 1994; Brittany, St. Lunaire, 20 shs, MNHN Coll. 
Fischer, 1898; Brittany, St. Servan, 10 shs, MNHN coll. 
Staadt, 1969; Brittany, St. Lunaire, 2 shs, MNHN coll. 
Ph. Dautzenberg (figured in Bucquoy et al, 1884, pi. 35, 
figs. 1, 2); St. Lunaire, 5 shs, MSNCS, 7173 and 7174, 20 
Jun. 1970, on the beach at low tide; Brittany, Morlaix, 
Saint Michel en Greve, 2 shs, coll. MF, 1976, E25A; 
Brittany, Saint Jacut, 3 shs, coll. MF, 06.1975, EI2A; 
Brittany, Saint jacut, 14 m, 6 shs, coll. MF, 04.1974, 
E12B; Brittany, Carnac, Quiberon, 4 shs, coll. MF, 1970, 
E16B; Brittany, Carnac, Quiberon, Pointe de Couquel, 2 
shs, coll. MF,' 1970, Ell A; Brittany, Saint Malo, 7 shs, 
coll. MF, 07.1973, E13A;Brittan\-, St. Malo, Lizardrieux, 
1 sh., coll. MF, Aug. 1982, E69A. Atlantic Pyrenees, 
Aquitaine, St. Jean de Luz, Cote Basque, infralittoral 
rocks, 1 sh., MNHN coll. S. Gofas, 1980-81; Atlantic 
Pyrenees, Aquitaine, St. Jean de Luz, 73 shs, MNHN 
coll. H. Fischer, 1898; Aquitaine, Soulac, 1 sh., MNIIN 
coll. A. Dolfus; Aquitaine, Hendaye, 2 shs, coll. VIE, Jul. 
1976, E57A. Portugal: Algan'c Sagres, Baie de Baleeira, 
(37°00.7' N, 08°55'.0' W)! tide zone, 1 sh., MNHN, Mis- 
sion Algarve, May 1988; Algaive Sagres, Ponta da Ba- 
leeira, "(37°00.3'' N, 08°55.5' W)," 17-23 m, 5 shs, 
MNHN, Mission Algai-ve, May 1988; vUgan-e Sagres, 
Pontal dos Coi-vos, (37°01.3' N, 08°58.3' W), at the'foot 
of fiilaise, 17-22 m, 5 shs, MNHN Mission Algan'e, May 
1988; Albufeira (southern coast), Ponta de Castelo, 3-6 
m, 2 shs, coll. MF, OS Aug. 1985, E50B. Atlantic Moroc- 
co: Asilah, mouth of Oued el Helou, conchiferous depos- 
its, beach, 6 shs, MNHN coll. S. Gofas, 1971-72. Strait of 
Gibraltar: Tanger, Grande Plage, conchiferous deposits, 
beach, 3 shs, '^MNHN coll. S. Gofas, 1970-81; Cadiz, 
Getares, beach, 3 slis, coll. MF, ex coll. G. Bogi, legit 
Ilanselaar, 2230 GET; Cadiz, Barbate, conchiferous de- 
posits, beach, 6 shs, MNHN coll. S. Gofas, 1976-81; 
south Ceuta, Punta del Desnarigado, (35°53.6' N, 
05°16.8' W), 16-20 m, 1 .sh., MNHN coll. Bouchet, Go- 
fas and Lozouet, May 1996. Mediterranean Spain: Cas- 
tellon, Columbretes Islands, Espinosa Island, 5 m, 1 sh., 
coll. MF, 26 Jul. 1974, M273A; Malaga, 15 m, 7 shs, coll. 
MF, ex coll. Cesare Bogi, 2127IMA(V002G); Malaga, 
Algeciras, Torre del Almirante, 3-5 m, I sh., coll. MF, 28 

Pacre 2^ 

THE NAUTILUS, Vol. 122, No. 1 

V. Garilli, 200S 

Page 23 

Figures 9-14. Alvania carinata (Da Costa. 1778), piiitiiriiiRli and details ot Sfulptiiic. 9. Sicil), l^ilenno, Terrasiiii, "Mai^au;<;;iare- 
Ciucca di Cinisi" beach, coll. PAL, (146E), protoconcli. 10. Pnn ence, Mar.seille, La Baule, small beach at 25 km west from Marseille, 
coll. P.\L (212SBAU-V008C), protoconch. 11-12. Shell trom the same locality and collection, sculpture of protoconch I and II (11) 
and a detail of protoconch I (12). 13. Same shell as Figure 2, detail of teleoconch sculpture on the penultiiuate whorl, coll. PAL 
(307B). 14. Detail of teleoconch sculpture on die last whorl, NW Sicily, Palermo, Terrasini, "Magaggiare-Ciucca di Cinisi" beach, 
detritus of Miniacina, coll. PAL (146E). Scale bars; 100 |jLm in Figures 9-11 and 13-14; 50 ^.m in Figure 12. White arrows indicate 
the protoconcli/teleoconch boundary', respectively. 

Sep. 1976, .\I2()7B; Malaga, Algeciras, 3 shs, coll. MF, 23 
Sep. 1976, M207A: Malaga, Fuengirola, 0.5-1 m, 1 sh., 
coll. MF, 20 Aug. 1973, M6SB; Malaga, Algeciras, Playa 
Getares, 6 m, 1 sh., coll. MF, Aug. 1983, M207E; 
Malaga, Cabo Pino, detritus, 10 m, 2 shs, coll. S. Riifini, 
(41.80g). Mediterranean Morocco: M'diq "(anc. 
Rincon)"" conchiferous deposits, beach, 1 sh., MNHN 
coll. S. Gofas, 1971. Algeria; Alger, 1 sh., MNHN coll. 
Locard. Mediterranean France: Languedoc, Carnon, 
conchiferous deposits, beach, 1 sh., MNHN coll. S. Go- 
fas. Aug. 1976; Languedoc, Roussilion, 3 shs, MNHN 
Coll. Dolfus, 1903; Languedoc, Roussilion, 1 sh., MNHN 
coU. Ph. Dautzenberg (Moll, du Roussilion); Provence, 
Marseille, La Baule, small beach at 25 km west from 
Marseille, 3 shs, coll. MF, ex coll. C. Bogi, Oct. 1986, 
2128BAU (\'008C); Provence, Marseille", Cape Cou- 
ronne. 4 shs, coll. MF, ex coll. C. Bogi, Oct. 1987, 
2129COU (VOOSE); Provence, Marseille; Grand Con- 

gloue, (43°10.6' N, 05°24.2' E), 33 ni, 31 shs, MNHN 
rec. H. Zibrowius Jun. 1996; Pro\ence. lies Embiez, con- 
chiferous deposits, beach, 10 shs, MNHN coll. S. Gofas, 
1968-70; Provence, lies Embiez, Petit Rouveau, dredg- 
ing ol sandy conchiferous bottom, 3-5 m, 5 shs, MNHN 
coll. S. Gofas, 196S-70; Piwence, St. Glair, infralittoral 
rocks, (43°08.2' N, 6°23.2' E), 0-1 m, 1 sh., MNHN rec. 
S. Gofas, Sep. 1992; Provence, St. Raphael, 2 shs, 
MNHN coll. Locard; Provence, 9 shs, MNHN coll. Petit, 
1873; Provence, Sanaiy, 1 sh., MNHN coll. Locard; Cor- 
sica, Ajaccio, 3 shs, MNHN coll. fousseaume, 1921; Cor- 
sica, Galeria, Punta StoUo, 6 m, 1 sh., coll. MF, 07 Aug. 
1984, M52D; Corsica, Pianottoli, Anse de Chevanu, 2 
shs, coll. MF, Jun. 1988, M77A. Tunisia: Djerba, Aghir, 
5 m, Posidonia bed, 1 sh., coll. MF, S. Palazzi legit 06 
Aug. 1993, M79A; Djerba, Al Jazii'ah. 1-2.5 m, 1 sh'., coll. 
MF, Aug. 1974, M9B. Italy: Friuli Venizia Giulia, Tri- 
este, beach, 7 shs, coll. MF, D. Di Massa legit 1976, 

Figures 1-8. Aliania ciniiiata (Da Costa, 177S), variation in shell shape and sculpture. 1. "Form" cciirincita Bucquoy et al., 1SS4, 
shell from coll. Ph. Dautzenberg (Moll, du Roussilion), MXIIX-DSE. 2. T\pical, keeled moiph. Liguria. Geneva, Camogli, 42 m, 
coU. P.\L ;307B). 3. MarkedK- keeled morph, corresponding to Rissoa trochlea Michaud, 1830, Strait of Gibraltar, Cadiz, Getares, 
beach, coll. PAL (2230 GET). 4-5. Small, unkeeled morph. La Spezia, Monterosso, 1.5-30 m, coll. PAL (70A). 6. Typical moiph with 
varice, Sardinia, Sassari, Capo Caccia, Gala deUa Calcina, 6 m. coll. PAL, (112A). 7. Profile xdew of the same shell as Figure 2. 8. 
Juvenile shell fitting well with the concept oi' Aha nia cinnulata (Philippi, 1836) sensu Piani (1979, figs. 2-3). Scale bars: 1 mm in 
Figures 1-7: 0.5 mm in Figure 8. 

Pase 24 

THE NAUTILUS, Vol. 122, No. 1 

307B; Liguria, Geneva, Camogli, 42 m, 1 .sh., coll. MF, 
06.1981, 271B: La Spezia, Monterosso, 15-30 m, detri- 
tus, 1 sh., coll. MF, A. Lugli legit Oct. 1978, 70A; La 
Spezia, Portovenere, 5-25 m, 3 stis, coll. MF, Oct. 1978, 
329A; La Spezia, Riomaggiore, 30 m, 1 sh., coll. MF, 
Aug. 1978, 312A; Liguria, La Spezia, Punta Mesco, 35 m, 

1 sii., coll. MF, Jul." 1987, 176C; Tuscany, Livorno, San 
Vincenzo, Borraccia, 12 m, 1 sh., coll. MF, 12 Jul. 1987, 
190A; Tuscany, Livomo, Secche della Meloria, 6-17 ni, 2 
shs, coll. MF, 1974, 34E; Tuscany, Livorno, Castiglion- 
cello, 4 shs, coll. MF, 1972, 21A; Tuscany, Livorno, 
Romito, off mouth of the torrent Chiona, 30-35 m, 2 shs, 
coll. MF, ex coll. C. Bogi, 36A; Tuscany, Livorno, Bagni 
Fiume, 20 shs, coll. Mf'~ 1977, 34B; Tuscan Archipelago, 
Island of Capraia, 100/400 m, 2 shs, coll. MF, ex coll. C. 
Bogi, 2126CAP(V()()5C;); Island of Capraia, Punta della 
Fic^a, 29 m, I sh., coll. MF. 15 Sep. 1985, 28R; Island of 
Capraia, Punta Civitata, 40 m, 1 sh., coll. MF, 19 Sep. 
1985, 28T; Tuscan Archipelago, Gorgona Island, 35-40 
m, 11 shs, coll. MF, Aug. 1978, 69A; Gorgona Island. 40 
ni, 18 shs, coll. MF, legit C. Bogi, 194; Tuscanv, Siena, 1 
sh., yellow sands from unknown layer, lower Pliocene, 
coll. MF ex coll. PAL, F22A; Tuscanv, Grosseto, Fmita 
Ala, Baia N'erde, 5 m, 1 sh., coll. MF,'G. Terzer legit OS 
Sep. 1974, 27A; Tuscany, Grosseto, Punta Ala, Punta 
Hidalgo, 2 shs, coll. MF, Jun. 1975, 27B; Tuscany, d'os- 
seto, Follonica, Gala Felice, 7 m, I sh., coll. MF, 25 Aug. 
1987, 55B; Tuscanv, Grosseto, Island of Palniaiola, 28 m, 

2 shs, coll. MF, 14 Sep. 1986, 172A; Grosseto, Island of 
Elba, Gapo Galamita, Scogh Corbelli, 46 m, 1 sh., coll. 
MF. Sep. 1972, 5E; Island'of Elba, Scoglio Remaiolo, 35 
m, 1 sh., coll. MF, Ma\- 1980, 5K; Island of Elba, For- 
miche della Zanca, 20 m, 1 sh., coll. MF, Apr. 1984, 5P; 
Lazio, Rome, Civitavecchia, 2 shs, coll. MF, 1975, 137B; 
Sardinia, Sassari, Capo Caccia, Gala della Calcina, 6 m, 
conchiferous detritus at upper limit ot Poskhviia bed, 10 
shs, coll. MF, S. Palazzi legit 19 Aug. 1983, 112A; Sar- 
dinia, Sassari, bay at E of Faro di Capo Testa, 7-19 m, 1 
sh., coll. MF, 22 Aug. 1983, 125B; Nuoro, Capo Comino, 
Ruia Island, 0-2 m,"l sh., coll. MF, 1974, 75A; Sardinia, 
Cagliari, Island of San Pietro, channel of San Pietro, 
2— t m, Posidnnia bed, 2 shs, coll. MF, G. Liuzzi legit 
10 Oct. 1976, 264A; Sardinia, Cagliari, Island of 
Sant'Antioco, Gala de Saboni, 1 sh.. coll. MF, 07 Aug. 
1983, 115B; Sardinia, Cagliari, Island of Sant'Antioco, 
Gala de Saboni, 14 m, I'sh., coll. MF, 10 Aug. 1983, 
115E; Campania, Napoli, Island of Capri, Pimta Vivara, 
6 m, 1 sh., coll. MF, 08 Sep. 1978, 254B; Campania, 
Napoli, Island of Procida, Marina Grande, 2.5-9 m, 
2 shs, coll. MF, 1974, 105A; Puglia, Bari, off Palese, 
12-13 m, 1 sh., coll. MF, 11 Sep. 1979, 77 A; Pugha, 
Taranto, 2 shs, coll. MF, 1973, 84B; Pugha: Taranto, 
Campomarino, 2 shs., coll. MF. 12 Feb. 1977, 309A; 
Puglia, Taranto, Maniggio, 1 sh., coll. .VIF, 1977, 280A; 
Puglia, Brindisi, "Batteria Bi'in" beacli, 14 shs, coll. MF, 
G.'Oriolo legit 08.1970, 193ID; Puglia,, Punta 
Croce, 15 m, 19 shs. coll. MF, G. Oriolo legit Aug. 1974, 
262A; Pu<j"lia. Biindisi, Torre (iuacetn, 5 in. I sli.. 

coll. MF, 1976, 42A; Brindisi, Lendinoso, 10-20 m, 1 sh., 
coll. MF, 1977, 279A; Puglia, Lecce, GaUipoH, Costa 
Brada, 2 shs, coll. MF, Apr. 1978, 9IB; Lecce, Porto 
Gesareo, 2 shs, coll. MF, Oct. 1977, 46B; Lecce, Porto 
Cesareo, Torre Lapillo, 1 sh., coll. MF, 06 Jun. 1978, 
46C; Lecce, Marina di Ugento, 30 m, 1 sh.,' coll. MF, 
1977, 68G; Lecce, Gallipoh, La Vecchia Torre, 2-6 m, 2 
shs, coll. MF, 1976, 91D; Lecce, San Cataldo, 1 sh., coll. 
MF, Apr, 1973, 260C; Lecce, Specchinlla, 1 sh., coll. 
MF, Oct. 1974, 28IA; Calabria, Reggio Calabria, Pen- 
timele, from fisherman nets, 8 m, 1 sh., coll. MF ex coll. 
Sciano, V060A 194; Reggio Calabria, Lam-eana di Bor- 
rello, Peeoraio, 2 shs, (layer 2), lower Pleistocene, coll. 
MF ex coll. PAL, F24A; Sicily, Trapani, San Giuliano, 10 
shs, coll. MF, 25 Jul. 1978, 297A; Trapani, Scopello, Ton- 
nara, 12-20 m, 1 sh., coll. MF, 06 Apr. 1986, 333F; Sicily, 
Trapani, Egadi Islands, Favignana, Secca del Toro, 30 m, 
20 shs, coll MF, A. Lugli legit 02 Jun. 1983, 272H; Egadi 
Islands, Fa\'ignana, 4 m, 17 shs, coll. MF; Egadi Islands, 
Favignana, Gala Rotonda, 20-30 m, 3 shs, 04 Jun. 1983, 
coll. MF, 272G; Egadi Islands, Favignana, Scoglio Cor- 
rente, 30 m, 3 shs,"coU. MF, 01 Jun. 1983, 272F; Egadi 
Islands, Favignana, Punta Sottile, 30-40 m, 4 sirs, coll. 
MF, 30 Mav 1983, 272E: Egadi Islands. Marettimo, 
Punta Bassana, 41 ni, 2 shs, eolj. MF, 04 Jun. 1983, 140A; 
Egadi Islands, Levanzo, Faraglione, 7-12 m, 3 shs, coll. 
MF, 03 May 1979, 325B; Sicilv, Pantelleria Island, Baia 
dei Fichi dindia, 22 m, 1 sh., coll. MF, Jul. 1983, 154M; 
Pantelleria Island, Punta Capace, 31 m, I sh., coll. MF, 
Jul. 1983, 154N; Sicily, Palermo, 1 sh., coll. MF ex coll. 
C. Bogi, (V056A) 194;' Palermo, Bagheria, Aspra, 18.5 m, 
2 shs,"coll. MF, Apr. 1973, I24C; Palermo, Punta Raisi, 
Marina Longa, 1 sh., coll. MF, 28 Feb. 1979, 320A; Pal- 
ermo, Teri'asini, "Magaggiare-Ciucea di Cinisi" beach, 
detritus oi Miniaeiiia. 14 shs, coll. MF, S. Palazzi legit 23 
Sep. 1977, I46E; Palermo, Island of Ustica, Scoglio del 
Medico, 25 m, 1 sh., coll. MF, Aug. 1980, 286K- Sicily, 
Messina, Milazzo, Capo Milazzo, Gala Sant'Antonio, 2 
shs, upper yellow sands, upper Pleistocene, coll. MF ex 
coll. PAL, F5A; Messina, Eolie Islands, Lipari, Secca del 
Bagno, 38-40 rn, 3 shs, coll. MF, 05 Sep. 1979, 338A; 
Sicily, Catania, Acitrezza, 3-25 m, 2 shs, coll. MF, Sep. 
1975, 22A; Sicily, Siracusa, Vendicari, ba\', 2 shs, coll. 
MF, 13 Sep. 1977, 20A; Siracusa, Portopalo di Gapo 
Passero, 2-3 m, 1 sh., coll. MF, Jun. 1976, 63A; Siracusa, 
Peninsula ot Magnisi, southern side, 1 sh., coll. \'. Ciarilli, 
Jun. 2006; Sicily, Palermo, Tonnnaso Natale, 1 sh., late 
middle Pleistocene, MGUP 166/2/49; Sicily, Trapani, 
Birgi, 3 shs, uppt'r Pleistocene, Tyrrhenian Stage, 
MGUP I76.5/36/I4; Sicih', Trapani, 12 shs, upper Pleis- 
tocene, T)rrhenian Stage, MGUP 358/3/42. Isle of 
Malta: Make, 8 ,shs, MNHN coll. jousseannn', 1921. 
Croatia: Istria, Rovinij, between lit. Nbuitrax, llr. Mun- 
trav and Hr. Azino, 8-15 in, detritus Irom bottom, 1 1 shs, 
coll. ,\ll\ S. Palazzi legit 19 Jun. 1978, MI88B; Istria, 
Savudrij;!, beach. 8 slis, coll. MF, S. Palazzi legit Oct. 
1975, M364A; lslii;i, lioMuij, (S-18 m, 19 shs, coll. MF, 
20 |uii. 1078, .VIISSA; Isti-Ki, \is:,r, 1 sh., coll. MF, 03 

\'. Garilli, 2008 

Pase 25 

Apr. 1988. M76A; Lstria, Umag, 1 ,sli., coll. Ml-", 1978. 
M109A; Lstria, Umag, Ta\-erna Loro, 0.3 ni, 1 sli.. 17 Jun. 
1978, coll. MF, i\ll()9B; Kaciack, dam ol' Cigalc, 4 m, 1 
sh., 29 Jul. 1983, coll. MF, M14A. Greece (Aegean Sea): 
NW .Aegean, Island of Limnos. near Moudro.s, 3-5 m, 48 
.shs, coll. MF, A. Lugli legit Aug. 1991, M8()A; Sithonia, 
Ormos Panagia.s, 2-3 m, 1 sh., Aug. 1982, coll. MF, 
M.31B; Sithonia, Nesis Dhiaporos, 33-34 m, 2 shs. coll. 
MF. 01 .-Vug. 1986, M31D. Unknowi locaUty from Medi- 
terranean Sea: 120 shs, MNHN coll. \'a\'ssiere. 

Habitat: Tlie species is usualK' found at depths com- 
patible with the upper part of the shelf, the infralitoral 
stage ot Peres and Picard (1964). Rarelv, I found material 
collecteti from the titlal zone. According to Gofas and 
Ponder (1991), Ahaiiia caiiiuita fixes deeply buried un- 
der stones. It seems also to be linked (in the Mediterra- 
nean) to phanerogam beds. The finding of two shells 
from the Island of Gapraia (Tuscan .\i-chipeIago), col- 
lected at a depth of 100-400 m, is much probabK due to 
lo\\'er shelf-slope transport. 

Distribution: Occurs probabK thi'oughout the Medi- 
terranean, but its presence in the easternmost coasts, 
from which I did not see any material, needs to be con- 
firmed. In the Atlantic it is recorded from tlie coasts of 
Great Britain soutli to Morocco. As fossil, it is rare in tlie 
Mediterranean Neogene where it is recorded from the 
lower Pliocene \ellow sands near Siena. It becomes more 
frequent dming tlie .Mediterranean QuaternaiT, where I 
found it from the lower Pleistocene of Reggio Calabria, 
and the middle-upper Pleistocene of SiciK- (Tommaso 
Natale, Capo Milazzo and Trapani). 

Remarks: A complete sviionyniy list was pnnided by 
Piani (1979) and \'an Aartsen (1982). In various collec- 
tions, I found different lots of this species contahiing 
ju\enile shells (not higher than 3 mm, see Figure 8) 
identified as Alvania cinoulata (Philippi, 1836), species 
hereafter re-described and discussed. This wr(.mg deter- 
mination \'eiy likely follows the misitlentification of Piani 
(1979, figs. 2-3) and Giannuzzi-Savelli et al. (1996, figure 

T\picall\-. the shell of this species has a characteristic 
keeled shape due to the presence of well mai'ked spiral 
cords that become \eiy strong on the well-de\eloped last 
whorl. The number of cords on achdt whorls is shahtlv 
\ariable. 3^ in the penultimate whorl. The not-keeled 
niorph. characterized by having a slender and turrited 
shape and more numerous cords (5-6 in the penultimate 
whorl), is die "form" ecariimta Bucquoy et al, 1884 (Fig- 
ures 1. 4—5), also knowii as mirwr-ecarinata Vlontero- 
sato. 1884 (probably corresponding to Rissoa hiciiUana 
var. cancellata Scacchi, 1836, as indicated bv Piani, 
1979). It has been considered a Mediterranean subspe- 
cies of A. carinata bv \'an Aartsen (1982). I found no 
shells clearK" belonging to this moi"ph in the Atlantic 
material studied (mainly from the M\HN-DSE collec- 
tions). Howe\'er, in some Mediterranean areas (e.g. along 
tlie French coast in the MXHN-DSE collections), I 

loinid tlu' hpit:il iiioiph together with unkeeled shells. 
Thus I am more inclined to considei' it just a case of 
intraspecific variation, which appears well represented in 
Mediterranean populations. The keeled moqjh, charac- 
terized by having three spiral cords on sul)adult whorls 
and a veiy exq^antled body whorl (more than 3/4 of the 
shell height, see Figure 3) fits well within the concept of 
Rissoa trocltlea Michatid, 18.30, which is ceitainly a syn- 
oinni of A. cariitata. 

Alvania carinata is the type species of Galeoclina 
Monterosato, 1884, a genus created for cingulated, 
keeled, varicose rissoid shells with a wide aperture 
(Monterosato, 18S4a, p. 163). This generic division can 
appear quite artificial, not being supported bv appropri- 
ate and constant taxonomic features. Also, the consider- 
ation of Gah'odina as a well-established subgenus oi Al- 
vania, as indicated by Wenz (1938) and Van Aartsen 
(1982), appears doubtful. Ponder (1985). who sviiony- 
mized most of the available subgenera witli Alvania 
sensu stricto, expressed some uncertaintv when consid- 
ering Galeoclina as a valid taxon. He I'ecognized the ig- 
norance of anatomical features as the main gap in solving 
this systematic question. In any event, the shell charac- 
ters alone do not provide enough support for Galeoclina 
to stand as a subgenus. For example, the unkeeled 
moq^h, which usually lacks the main features of Galeo- 
clina, including the characteristic varices of the keeled 
uKJiqDh, might be compared to the Alvinia Monterosato, 
1884, species group, with which it shares a slender, tur- 
reted shell shape. It is noteworthy that Cossmann (1921) 
cited Galeoclina as a .sMionvm of Alvinia. 

\lvania cingidata (Philippi, 1836) 
Tigures 15-26) 



Rissdd eingulata I'liilippi, lS3(i: 152-1.53. 

Acinus cingiilatus (Philippi). — Monterosato, 1884b: 62. 

Cinguhi (Onoha) cingulcita (Pliilippi, 1836).— Nord,sieck, 1968: 

46, pi. VII, fig. 26.35 
Alvinia (G(ih'oclii}<i) cingulata (Philippi, 1836). — Nordsieck, 

1972; 181, pi. IWI, fig. 20. 

Description: Small, sturdy, conical-o\ate shell reach- 
ing 3.5 mm in height, 2.2 mm in width. Protoconch pau- 
cispiral, consisting of little more than 1.5 convex and veiy 
rounded whorls, sculptured bv 5-6 \'eiy fine spiral lirae. 
Several prosocirte growth scars precede protoconclV 
teleoconch transition (Figure 20). Teleoconch consists of 
about 4 cingulated convex whorls separated by slightly 
inclined sutures. Ven' early teleoconch whorls lieai- tx\'0 
main spiral cords crossed by numerous axial ridges. 
Sculpture of adult \viiorls consisting of spiral cords more 
conspicuous tlian axials, which are 5-6 and 10-11 on the 
penultimate and body whorls respectivelw Axial sculp- 
ture formed by narrowei' (half the lireatlth of a spiral 
cord) ribs, numbering about 40 on penultimate whorl, 
and which become obsolete toward shell base. Vlicro- 
sculpture consists of spiral rows of small tubercles that do 
not seem to appear on main sculpture (Figure 26). A few 
irregular scars cross these spiral rows. Last whorl ratlier 

Paee 26 

THE NAUTILUS, Vol. 122, No. 1 

\'. Garilli, 20()S 

Fuse 27 

Figiu-es 24-29. Ahania cin'^ulata (Philippi, 1836) from type localih' (Peninsula of Magnisi. Siracusa, SE Sicily, coll. Garilli), and 
a slightlv resembling Ahania sp., from the lower Pleistocene of Musala (Italy, Reggio Calabria, coll. MF ex coll. Pal, F95A), fitting 
well with the concept of A. ciiigiihita stated b\' Monterosato (18.S4b). 24-26. Ahania cingnlata. Apertural \ie\v of a subadult shell 
(24), sculpture on last whorl (25) and microscnlpture (26). 27-29. Ahtinia sp, Apertural (27), dorsal (28) vievv.s and protoconch. Scale 
bars: 500 jxm in figures 24, 27-28; 100 | in figures 25, 29; 20 jxiii in I'igure 26. Tlie white arrow in Figvu'e 29 indicates the 
protoconcli/teleoconch boundaiy. 

inflated and \vell-de\'elopecl, comprising little less than 
3/4 total height. Aperture wide and ovate, comprising 
about 4/9 and 2/3 of total height and last whorl respec- 
tivelv. Outer lip slightK" prosocline, internally smooth, 
extemallv shghtlv thickened near edge. Columellar side 
arcuate, with a thin callus forming a very narrow umbili- 
cal chink. Coloration consisting of quite large, reddish- 
brown bands on a whitish-cream background. 

Type Locality: 

eastern SiciK. 

Peninsula of Magnisi, Siracusa, south 

Type Material: One possible ,s\iit)pe, ZMB (2326) ex 
coll. Philippi was destro\ed In' B)'ne's reaction (M. 
Glanbrecht, pcrs. conmi., 2006) 

Material Examined: Italy: NE SiciK', Messina, 2 shs. 
ZMB ex coll. Monterosato, (S0914 (original label by 

Figures 15-23. Alvania cingnlata (Philippi, 18.36). 1.5-17. .Apertural (15), dorsal ( 16), profile (17) view of one shell from ZMB ex 
coll. Monterosato (80914), Messina, NE Sicily. 20, 23. Protoconch of the same shell. 18-19 and 21-22. Another shell from the same 
lot, apertural (18), dorsal (19), profile (21) view, and a detail of sculpture on the last whorl (22). Scale bars: 1 mm in figures 15-19 
and 21: 200 jjim in Figure 22; 100 |a.m in Figure 20; 50 |jim in Figin-e 23. The white arrows indicate the approximate protoconcli/ 
teleoconch boundarv. 

Page 28 

THE NAUTILUS, Vol. 122, No. 1 

Monterosato: Acinus cingidattis, 2, Messina, 1014); SE 
Sicilv, Siracusa, Peninsula of Ma2;nisi, south side, detritus 
from the beach, 1 sh., coll Garilli, 6/2006 

Habitat: Unknown. It is n()tewortli\ that sand\' and 
Posiclonia bottoms prevail in the south side ot the Majj;- 
nisi Peninsula (from where I collected the beached shell 
in Figure 24), while an almost paralic (pre-lagoon) envi- 
ronment (wth Cerastodenna. Cerhhiiim, and Gibbitki 
spp., on a finely sandy bottom) and a rocky bottom with 
algae characterize the north and east sides. 

Distribution: Its distribution appears to be limited to 

Sicily, with particular regard to the eastern coast, from 
Messina to Magnisi. It was recorded from Palei-nio h\ 
Monterosato (1872; 1875; 1878; 1884a; 1884b), who ven 
likely followed the citations of Philippi (1844) and 
Weinkauff (1885). The latter author, who indicated Mon- 
dello (the beach near Palermo), also cited Ognina (Cata- 
nia) in eastern Sicily. To ni)- knowledge, no fossil record 
of this species e>dsts. 

Remarks: This is a vei'v rare, practically unknown spe- 
cies (see Van Aartsen, 1982a). Its identification has tra- 
ditionally been quite problematic, above all after the 
misidentification b\- Piani (1979, (igs.2-3), who illus- 
trated a juvenile shell of Ahania carinata (l^a Costa, 
1778) under the name Galeodina cingidata (Philippi, 
1836). The shell figured b)' Giannuzzi-Savelli et al. ( 1996, 
fig. 499), vmder the name Alvania (Ahania) cingulata 
(Philippi, 1836), is a subadult of A. carinata. 

My identification is based on three shells (one from 
the type locality, two from ZMB ex Monterosato coll.. 
labelled as Acinus cingulafus), which agree well vvith the 
original description bv Philippi (1836; see also 1844: 
128). A few diagnostic characters allow for its correct 
identification: the primaiy cingulated sculpture \\ith 
raised cords, nmnbering 10-11 on the last whorl; the 
secondan' axial pattern formed hv \en' narrow ribs cross- 
ing the cords; the large ovate aperture; and the internally 
smooth outer lip, slightly thickened near its edge. 

As Philippi (1844) noted, Alvania cingulata may be 
compared with A. tenera, especially with the slentler 
moi-ph of the latter species (see Figure 78). Alvania te- 
nera has a smaller shell (usually not over 2.2 nnn in 
height) with more raised cords (numbering 12-14 on tlic 
last whorl), and almost lamella-like ribs, which occur just 
on the interspaces between the cords (see discussion of 
A. tenera). Furthcniiore, A. cinguliila has a paucispiial 

In addition, Noixlsieck (1972) and Van Aartsen (1982) 
considered this species as belonging to Galeodina. How- 
ever, the similarit\- between Alvania ciniiulata and A. 
carinata (type species of Galeodina) is superficial. Sev- 
eral differences separate the two: A. cingulata has a less 
inflated last whorl bearing less nunierf)us coixls and lack- 
ing varices, its outer lip lacks a varicose thickness, and its 
spiral sculpture does not mai'kedlv dominate over axial as 
in the typical /\. carinata. The lattei' species also has a 
multispiral protoconch. 

Monterosato (18S4a and lS84b) included this species 
in his genus Acinus (synonvm of Alvania according to 
Fonder, 1985), creating a link wth the t\pe species Al- 
vania cimex (Linnaeus, 1758) (which is also the type 
species of Alvania, see Ponder, 1985: 36), mainly based 
on coloration. In realitv, the similaritv between the two 
species is limited to the color pattern of the shell and the 
nodular sculptu]-e. Strong chfferences indicate that A. 
cingulata and A. cimex could be distant relatives, the 
latter having a more sturdy shell with a coarser sculpture 
(with subequal axial and spiral), and a proportionally 
smaller aperture provided with an internallv denticulate 
outer lip. The concept of A. cingulata as expressed by 
Monterosato (1884b: p. 62) seems to be applicable to an 
interesting Alvania sp. (from the lower Pleistocene of 
southern Italy, Reggio Calabria, Musala, 19 shs, F95A, 
coll.. Figures 27-29) rather than to the original descrip- 
tion b)- Philippi. The non-planktotrophic (see the pau- 
cispiral protoconch in Figure 29) Alvania sp., illustrated 
here, showing only a moderate similaritv' with the teleo- 
conch sculpture of A. cingulata, has the characteristic 
general shape of A. cimex, with which it shares the ap- 
ertural features, including the denticulations on the inner 
part of the outer lip. Furthermore Monterosato himself 
determined the shells I ha\'e illustrated in Figures 15-23 
as A. cingulata. 

In 1968, Nordsieck cited A. cingulata as Cingula 
(Onoha) cingulata. In reality, a vague similarih' with 
some Cingula Fleming, 1828 or (more specifically) 
Onoha H. and A. Adams, 1852, species exists; e.g. Onoha 
iO.) carpenteri (Weinkauff) (see Ponder, 1985: fig. 
1 14a), bears a similar spiral sculpture. Anywav, the genus 
Onoha is characterized by usually elongate shell with a 
proportionally smaller and more rounded aperture. Its 
included species, usuallv lacking the axial sculpture, may 
have a much weaker axial pattern or this may consist of 
delicate ribs, which become obsolete towards the sutures 
(see Boucliet antl Waren, 1993: figs. 1508-1509, 1514- 
1515). In addition, the protoconcli sculpture apparent on 
some non-planktotrophic Onoha species resembles that 
of A. cingulata, in having few, veiy fine spiral widely 
spaced threads. This kind of sculpture is also seen in 
Alvania (e.g. A. suhsoluta (Aradas, 1847), see Bouchet 
and Waien. 1993, fig. 1458). Cingula species appear to 
be less similar to A. cingulata in having a thick outer lip 
and lacking anv axial sculptural pattern. 

A certain similaiity exists with Alvania ivat.soni (Wat- 
son, 1873) From Madeira (see Ponder, 1985: fig. 102c-d), 
with which A. cingulata shares the general shell shape, 
characterized by an inflated last whorl provided with a 
large, ovate aperture. 

Alvaniii fnincescoi new species 
(Figures 30-43) 

Description: Shell small, sturdy, conical-ovate, reach- 
ing 4.8 mm in luight (4.3 nnn in holotvpe) and 3.1 mm 
in width (2.8 mm in holotvpe). Protoconch conical, nntl- 
lispiial with p:irliall\ ininiiTscd niick'us and convex 

\'. Garilli, 200S 

Paee 29 

whorls. Protoconcli I fDnsistinu; ot aLxnit O.S whorls 
sculptured In 5-6 line spiral lirae interspersed spirally 
and irregularly wirii microscopic granules. Protoconch II 
consisting of about 1.2 whorls sculptured 1)\ pimples, 
which are irregular in size and arrangement. The\' are 
larger and less numerous in the adapic.iJ position; fused 
into \en' short prosocline tracts in central pait. in the 
abapical part of late protoconch, groups of pimples are 
irregularK fused into tine, discontinuous spiral ridges. 
Protoconcli/teleoconch transition well-marked and sinu- 
ous. Teleoconch consisting of 3.(S— 1.6 (4.2 in holotyjie) 
rather convex whorls, with strong eaneellate sculpture 
formed b\' intersection of equally dexeloped spiral cords 
asid axial ribs. Cords are narrow and well raised, num- 
bering 2, 6-7 (6 in holotApe) and 13-15 (14 in holotype) 
in eaiiv, penultimate and bod\- \\'horl, respectively. Two 
adapical cords are usualK mori' pronounced in penulti- 
mate and last whorl. Cords are markedly stronger and 
more wideK' spaced on base. Secondan' cords may com- 
mence on late bod\" whorl, at a certiiin distance from 
outer lip. ;\.\ial sculpture consists of narrow raised ribs 
numbering 24—32 (26 in holot\pe) on the penultimate 
\\horl. On base, ribs progressively become yeiy nan-ow to 
obsolete toward columellar area. Intersection of spiral 
and axial sculpture, producing notlular small knobs, 
forms a rectangular pattern (with major side of rect- 
angles peipendicnlar to shell axis) except on last whorl, 
where a quadrangular to rhomboidal pattern occurs (Fig- 
ure 43). Microsculpture of xeiy early teleoconch whorls 
consists of micropustules, sometimes fused, forming spi- 
ralis discontinuous and irregular rows, mainly occiu'ring 
bet\\-een spiral cords (Figure 42): weak growth lines are 
present. Fine sculpture of the rest of teleoconch limited 
to weak growth lines (Figure 43). Suture slightly in- 
clined. Last W'horl well-developed, comprising about 
diree foiuths of shell height. It may bear single (paratope 
6) or double (paratope 4, Figure 37) varices, at angles of 
about 50° and 70° respectively. Aperture wide, ovate, 
and widi a rounded profile in the posterior part, com- 
prising about two thirds to three fourths of last whorl 
height. Outer lip orthocline, intemalK- smooth, extemalK" 
markedly thickened by a strong, sometimes doubled 
(Figure 31), rim, occurring very close to its edge and 
covered hv spiral cords. Inner lip moderately arcuate and 
radier thickened, with thin callus delimitating vei-y nar- 
row umbilical chink. 

T-vpe Locality: Lower Pleistocene of Cartiera Mulino 
(3'6°56'57" \.'l4°34'03" E), Vittoria, Ragusa, SE Sicily. 
The stratot\pe is the phanerogams-rich 3D1 layer of 
Costa (1989). This deposit crops out at about one hun- 
dred meters from the abandoned paper-mill known as 
Cartiera Mulino. Paleoemlronmental and stratigraphic 
information about this site are given by Costa (1989). 

T\pe Material: Holot\pe (4.3 x 2.8 mm), DGUP 
C'MRG 00.5/488. coll. \'.'Garilli. Parat\pe 1 (4.1 x 2.7 
mm), DGUP CMRG 006/489, coll. V. GariUi; parat\pe 2 
(4.8 X 3 mm), ZMA Moll. 4.07.007 (ex CM GR 007/490, 
coll. Garilli); paratope 3 (4 x 2.7 mm). ZMB MB.Ga.2479 

(ex C.MGR 008/49, coll. \'. Cianlli); parat\pe 4 (4.2 x 2.9 
mm), MNHN A25950 (t'.v CMGR 0()9/492, coll. V. 
Garilli); parat)pe 5 (4.6 x 3.1 mm), DGUP CMGR 010/ 
493, coll. v. Garilli; paratype 6 (4.5 x 3.1 mm), DGUP 
CM GR 01 1/494, coll. V. Garilli. All type material is from 
the tvpe locality, 3D1 layer. DATE? 

Etymology: The species honors the name of botli 
Francesco Garilli senior, my father and first mentor in 
my life, and ol Francesco Garilli junior, my son. 

Habitat: All the shells came from the 3D1 layer of 
C^osta (1989). The paleoenviromental reconstruction for 
this layer was characterized by Costa (1989) as compa- 
rable with the Mechterranean marine-marginal modern 
ecotone IIP (Posidonia beds)-SVMC {scnsii Peres and 
Picard, 1964). It is interesting that the 3D1 layer is quite 
rich in Posidonia remains mainly consisting of leaves. 

Distribution: This species is known from type locahty 
only, lower Pleistocene of Cartiera Muhno, Vittoria, Ra- 
gusa, SE Sicily. 

Remarks: This quite characteristic species shows a ga- 
Icodiiiiform shape, having a quite inflated and well- 
developed (also yvith double varix) last whorl and a large 
aperture provided with an internally smooth outer lip. 
However, its spiral sculpture does not markedly prevail 
over the axial, as in the tvpical Alvania caiinata. Its mi- 
crosculpture, limited to the early teleoconch whorls, is 
similar to that obsei'ved in the analogous whorls of A. 
caiinata, but in the latter this microsculpture is distrib- 
uted over the entii'e teleoconch. 

The strongly cancellated sculptural pattern of A. 
franccscoi new species, consisting of well-raised, sub- 
equal spiral and axial elements forming marked nodular 
intersections, resembles that shovvm bv A. cimicoidcs 
(Forbes, 1844). The latter .species, showing a more typi- 
cal A/ufl)i/rt shape, has a more slender shell, a less inflated 
and much less developed last whorl, and a proportionally 
smaller aperture with denticulations on the inner part of 
the outer lip. 

Alvania francescoi new species can also be compared 
with A. rosarioe new species described later in this re- 
port. Remarks on their similarities and differences will 
be dealt with in the section dedicated to the latter spe- 

Alvania lactea (Michaud, 1830) 
(Figures 44-49 and 53-61) 

Piissoa lactea Michaud. 1830: 9-10, figs. 11-12 

Rissod CMaswtid) hieica Michaud. — Bucquov et al, 1884: 298, 

pi. 25, figs. 7-13 
Massotia Dnjerleini Monterosato e.r-Schvvartz ms, 1889: 33. 
Massotia lactea (Michaud) forma Dajerleini Monterosato, 

1917: 12. 

Description: Sliell stiuxK , conical-ovate to cviiudrical- 
ovate, occasionally almost pupoid, reaching 5.2 irnn in 

Page 30 

THE NAUTILUS. Xol. 122, No. 1 

V. Garilli, 2008 

Page 31 

Figures 3S— 43. AliYinia jrinicencoi new species, protoconcli, teleocoiirh sculpture and niicrosculpture, lnwer Pleistocene of 
Cartiera Mulino, SE Sicilv, Ragnsa, Vittoria. 38. Protoconch, paratype 2. 39. Protoconcli, transition bet^veen protoconcli I and II, 
holot\pe. 40. protoconch I, holotype. 41. Sculptiu'e of early teleoconch whorls, note how the spiral niicrosculpture becomes lacking, 
pai'atxpe 2. 42. Niicrosculpture on the first whorl, paratype 2. 43. Detail of sculpture on the last whorl, note the numerous growth 
lines, paratope 4. Scale bars: 200 (xm in figures 41 and 43; 100 (jini in figures 38-39; 50 |jini in figures 40 and 42. Black and white 
arrows indicate the protoconch I/protoconch II and protoconcli/teleoconch boundaries, respectiveh'. 

height and 3.3 mm in width. Protoconch conical, consist- 
ing of about two convex whorls. Nucleus partially im- 
mersed. Protoconch I consisting of about 0.8 whorls, 
sculptured b\' -5-7 spiral lirae and intei'spei-sed with nu- 
merous closely packed granules, which are spirally and 
irregularh- arranged. Protoconch II consisting of about 
1.2 whorls sculptured by pimples irregular in size and 
arrangement. Pimples are fused into short inclined lines 
(opistliocline and prosocline. Figures 53-54) or form one 
to three discontinuous spiral ridges in abapical aspect of 
whorl. Protoconcli/teleoconch transition well-marked 
and sinuous. Large shells consist ot 4.2-5 weakly to 
rather convex whorls. These may be sculptured by a 
fineh" cancellate pattern formed by the intersection of 
spiral cords and numerous deUcate, narrow axial ribs, or 
b\' opisthochne, pronounced ribs. Ribs often start from 
earK" whorls and are crossed bv finer spiral threads. In 
cancellate shells, cords and ribs are equal to subequal 
and numbering up to nine and 40 respectively on pen- 
ultimate whorl. In ribbed shells, cords and ribs are less 

numerous, numbering up to eight and 16 respectively on 
penultimate whorl. In early teleoconch whorls micro- 
sculpture consists of fine, irregular, and discontinuous 
spiral threads formed by groups of small granules, not 
covering main spiral sculpture; remaining teleoconch 
with a finely reticulate ultrastructure covering all primary 
sculpture, formed by intersection of very narrow and 
raised lamella-like axial ridges and vew thin spiral 
threads. Suture slightl)' inclined and deeply impressed. 
Body whorl well-e.xpanded, comprising % of shell height. 
It may bear a varix, usually with angle of 180°; rarely a 
double varix may occiu', espeeiallv in slender and cancel- 
late shells. Aperture wide, ovate to almost pyriforme, 
rounded on its posterior aspect, narrowed anteriorly, 
comprising two thirds of last whorl height. Outer Up or- 
thocline, internally smooth, externalK' thickened, niainlv 
somewhat behind its edge, and co\ered by priman- spiral 
sculpture. Sometime, in ribbed nioiphs, the outer lip 
descends vei"tically, so that last whorl appears rather 
cylindrical. Inner lip weakly arcuate, with a thin to rather 

Figures 30-37. Ahania francescoi new species, holot\pe and two paratspes, lower Pleistocene of Cartiera Mulino, SE Sicily, 
Ragusa, Mttoria. 30-32. Ilolonpe, shell in apertural (.30), profile (31) and dorsal (32) view. 3,3-3.5. Parat)pe 3, shell in apertural (.33), 
profile (34: and dorsal (3.5) view; 36-37. Parat\pe 4, apertural (36) and dorsal (37) view, the last showing a double vartx on the last 
whorl. Scale bars; 1 mm. 

Page 32 

THE NAUTILUS, Vol. 122. No. 1 

\'. Garilli. 2008 


Figures 53-61. Ahaniii hictca (Michaud, 1830). protocoiich and teleoconch sculpture and microsculpturi'. 53-54. Profile and 
dorsal \ie\\s of protocoiich from the same shell as Figure 44, Italy, Tuscany, Siena, Poggibonsi, Villa Pietratitta, podere Melograni, 
lower PUocene, coll. MFev coll. PAL (F55E). 52. Protocouch illustrating variation of the abapical spiral ridges. Italy, Tuscany, Siena, 
Poggibonsi. N'iUa Pietrafitta, podere Sant'Uliviere. lower Pliocene, coll. MF ex coll. PAL (F55C). .56-57. Sculpture of protoconch I 
(56) and earh' teleoconch whorls (57) of tlie same shell as ligures 44 and 53-54. 58-61. Early (58) and ven' early (59) teleoconch 
sculpture, and niicrosculpture of first (60) and last whorls (61), from the same shell as Figure 42, Monastir-Khenis, coll, Garilli. Scale 
bars: 200 fjim in figures 57-58 and; 100 | in figures 53-55, 58, 61; 50 |xni in Figure 60; 20 |xiu in Figure 56. Black and white arrows 
indicate the protoconch I/protoconch II and protoconcli/teleoconch lioundaiies, respectiyely. 

thick narrow calhis, leaxing a veiT small uniliijical chink. 
\'er\' fresh shells show a ierruginous periostracum. Shell 
color white. 
T\pe Localit\': Michaud (1831) cited the following lo- 

calities, all belonging to the French Mediterranean coast: 
""Agde, Cette (Herault) [now Sete], Callioure, Port- 
Vendre (Pyrenees orientales)". (With this citation I con- 
sider the second edition, consulted in the MNHX-DSE 

Figures 4-1-52. Shells ol'Akania lactca (.Michaud. 1830) and Ahaiiia pruai (Fischer, 1877). 44—49. Ahaiiiu Idi-lcu. 44. ItaK, 
Tuscany, Siena, Poggibonsi, X'rUa Pietrafitta, Podere Melograni, lower PHocene, coll. MF ex coll. PAL (F55E), cancellated nioiph. 
45. Tunisia. Monastir-Klienis. eoU. Garilli. ribbed morh (yar. dajerleini Monterosato). 46. Italy, Piemonte, Asti pro\ince. Pliocene, 
MGUP coU. Doderlein (113B) shell close to the original description by Michaud (1830). 47. Italy, Piemonte, Alessandria, Tortona, 
Miocene (Tortoiuan), VIGUP coll. Doderlein (476B). 48. Profile view of the same shell as Figure 41. 49. Juvenile shell, Mediter- 
ranean France, Provence. Marseille, La Baule, coll. PAL. (2128BAU-V()08C). 50-52. Syiitvpe of Rissoa pnisi Fischer, 1877, 
Quateman- of Rliodes. MNHX ■■ DHT) coll. D'Orbisinv ' R07495), apertural (50), profile (51) and dorsal (52) views. Scale bars 1 mm. 

Page 34 

THE NAUTILUS, Vol. 122, No. 1 

libraiy, of Michaucrs work "Descriptions deplusieurs . . ." 
published for the first time in 18.30. The date 1831 may 
he doubtful, 1832 being the most commonly cited date; 

1 prefer to follow Palazzi (2003), who provided helpful 
reasons to choose the former date.) 

Material Examined: 37 shs, coll. H. Fischer, wth no 
locality. Atlantic France: Normandy, St Aubin CaK'ados, 

2 shs,'MNHN; Normandy, St.Vaast, 1 sh., MNHN; Brit- 
tany, Finistere, Anse de Dionan, under stones covered 
with sand, 75 shs, MNHN legit. S. Gofes, 197.3-78: Brit- 
tany, St. Lunaire, 40 shs, coll. MNHN coll. Dollfus, 1903; 
St. Lunaire, 15 shs, MNHN coll. Fischer; Brittany, 
Pentliierre, 4 shs, MNHN, P. Bouchet legit; Penthierre 
(Morbihan), under stones covered with sand, low tide, 10 
shs, MNHN coll. P. Bouchet; Penthierre, 10 shs, 
MNHN, 27 Apr. 1975; Aquitaine, Cote Basque, Hen- 
daye, conchiferous detritus, beach, 5 shs, MNHN coll. S. 
Gofas, 1981; Cote Basque, Hendaye, infralittoral rocks, 3 
shs, MNHN coll. S. Gofas, 1980-81; Cote Basc^ue, Ond- 
arroa, infralittoral rocks, 1 sh., MNHN coll. S. Gofas, 
1980-81; Cote Basque, St. Sebastian, infralittoral rocks, 
1 sh., MNHN coll. S. Gofas, 1980-81; Cote Basque, St 
Jean de Luz, infralittoral rocks, 57 shs, MNHN coll. S. 
Gofas, 1980-81; St Jean de Luz, outside Cape Ste Barbe, 
tide zone, stones covered with sand, 4 shs, MNHN coll. 
S. Gofas, Dec. 1988; St Jean de Luz, outside cape Ste 
Barbe, tide zone, stones covered with sand, 49 shs, 
MNHN coll. S. Gofas, 1989; St Jean de Luz, 54 shs, 
MNHN coll. H. Fischer, 1898; Aquitaine, Guethaiy, 7 
shs, MNHN coll. H. Fischer, 1898. Portugal: Algai-ve 
Sagres. Pontal dos Con-'os, {37°01.3' N, 08°58.3' W)"; foot 

of falaise, 17- 


1, 1 sh., MNHN Missione Algar\'e, 05. 

1988; Algai-ve Sagres, Baie de Baleeira, (37°00.7' N, 
08°55.0' W), tide'zone, 1 sh., MNHN Mission Algai-ve, 
May 1988. Atlantic Spain: Cantabria, Orifion prov. 
Santander, Punta de Sonabia, infralittoral rocks, 1 sh., 
MNHN coll. S. Gofas, May 1989; Asturias, Muros prov. 
Oviedo, plava de la Liana, infralittoral rocks, 2 shs, 
MNHN coll. J. Ortea-S. Gofas 08.89; Cadiz, Barbate, 
(36°10.9' N, 05°56.9' W), tide zone, infrahttoral rocks, 4 
shs, MNHN rec. S. Gofas Apr. 1994; Cadiz, Chiclaua, 
(.36°22.5' N, 06°12.5' W), tide zone, infralittoral rocks 
and sands, 12 shs, MNHN rec. S. Gofas, Apr. 1994; 
Cadiz, Barbate, conchiferous detritus, beach, .3 shs, 
MNHN coll. S. Gofas, 1976-81; Mediterranean Spain: 
Malaga, Calahonda, conchiferous detritus, beach, 9 shs 
MNHN coll. S. Gofas, 1976-81; Malaga, Port de Mar- 
bella, conchiferous detritus, beach, 3 shs, MNHN coll. S. 
Gofas, 1978-81; Malaga, Benalmadena-Costa, conchifer- 
ous detritus, beach, .3 shs, MNHN rec. S. Gofas, 1991- 
93. Atlantic Morocco: El Jachda, (.33°16' N, 08°29' W), 
large beach, rocky platform, tide zone, 16 shs, MNHN 
rec. S. Gofas, 26 Sep. 1991; Fedala, Mannesmann beach, 
conchiferous detritus, beacli, 20 shs, MNHN coll. S. Go- 
fas, 1970-72: Asilah, mouth of Oued el Heloii, conchil- 
erous detritus, beach, 21 shs, MNHN coll. S. Gofas, 
1971-72; Asilah, mo\ith of Oued el Helou, conchiferous 
detritus, beach, 30 shs MNHN coll. S. Gofas, 1972-80; 

Temara, (33°55' N, 07°00' W), Sables dOr beach, rocks 
and mud, 0-2 m, 24 shs, MNHN, MA48, rec. S. Gofas, 
17 Sep.1991; Essaouira (formerly Mogador), (SFSF N, 
09°47' W), rocky platform, tide zone, 1 sh., MNHN, 
MA48, rec. S. Gofas, 23 Sep.1991; Rabat, Lahlou, 
(34°02' N, 06°51' W), conchiferous detritus, beach, 2 shs, 
MNHN rec. S. Gofas, 28 Sep. 1991; Essaouira (formerly 
Mogador), 4 shs, MNHN; Strait of Gibraltar, Morocco: 
Tanger, Grande Plage, conchiferous detritus, beach, 8 
shs,'MNHN coll. S. Gofas, 1970-81. Strait of Gibraltar, 
Spain: Ceuta Nord, Benzu, iniralittoral rocks, 2 shs, 
MNHN coll. S. Gotas 1976-1981. Algeria: Oran, 6 shs, 
MNHN coll. Locard. Mediterranean France: Langue- 
doc, Roussilion, Banvuls snr Mer, near the beach de 
Pauhlles, iuiialittoral rocks, 1 sh., MNHN coll. Bouchet 
and Gofas, Sep. 1980; Languedoc, Roussilion, 7 shs, 
MNHN coll. Ph. Dautzenberg, (figured in Moll. Rouss. 
T. I pi. 35, figs. 7-13); Languedoc, Roussilion, Sete, 6 shs, 
MNHN coll. Locard; Provence, Toulon, 8 shs, MNHN 
coll. Petit; Provence, Cannes, 21 shs, MNHN coll. DoU- 
ius, 1903; Provence, east coast, lies Embiez, passe du 
Gaou, under stones covered with sand, 0-1 m, 5 shs, 
MNHN rec. S. Gofas, Aug. 1988; Provence, east coast, 
lies Embiez, (43°04.3' n" 5°47.4' E), passe du Gaou, 
under stones covered with sand, 0-3 m, 2 shs, MNHN 
rec. S. Gofas, Aug. 19S8; Provence, lies Embiez, 
(43°04.3' N, 5°47.4'"e) passe du Gaou, rocks, photophile 
algae under stones covered with sand, 0-3 m, 1 sh., 
MNHN rec. S. Gotas, Jun. 1995; lies Embiez, cote Nord 
et Pedt Rouveau, infrahttoral rocks, 3 shs, MNHN coll. 
S. Gofas, 1968-70; Provence, Les Embiez, cote Nord et 
Petit Rouveau, (43°05' N, 5°47' E), rocks, photophile 
algae, 0-1 m, 3 shs, MNHN rec S. Gcifas, Jun. 1995; lies 
Embiez, conchiferous detritus, beach, 15 shs, MNHN 
coll. S. Gofas, 1968-70; Provence, St Clair, (43°08.2' N, 
6°23.2' E), infralittoral rocks, 0-1 m, 2 shs, MNHN rec. 
S. Gofas, Sep. 1992; Provence, Porquerolles plage Notre 
Dame, (43°00.6' N, 6°13.8' E), rocks, 0-1 m, 1 sh., 
MNHN rec. S. Gofas, Sep. 1992; Provence, Le Dramont, 
conchiferous sand, beach, 3 shs, MNHN rec. J. Pelorce, 
1992; Provence, Marseille (Endoume, 43°16.9' N, 
05°21.0' W), littoral rocks, 1 sh., MNHN rec. S. Gofas 
Apr. 1995; Provence, St. Raphael, 5 shs, MNHN; 
Provence, Bandol, 16 shs, MNHN coll. Locard; St. 
Haphael, 2 shs, MNHN coll. Locard; Corsica, Ajaccio, 17 
shs, MNHN coll. Jousseaume; Corsica, Algajole, 35 shs, 
coll. MF, 2123. Tunisia: Monastir-Khenis, beach, 1 sh., 
coll. Garilli, legit Garilh and Galletti 4/2000; Djerba, 
beach, 1 sh., coll. MF c.v coll. C. Bogi, Jun. 1981, 
216.3JER. Italy: Piemonte, Asti, Baldichieri, "Grottino 
Monale", 20 shs, yellow sands, middle-upper Pliocene, 
coll. MF ex coll. PAL, F104A; Piemonte, Asti, 24 shs. 
Pliocene, MGUP coll. Doderlein, 113B; Piemonte, 
Alessandi-ia, Tortona, 1 sli., Miocene, MGUP coll. 
Doderlein, 476B; Alessandiia, N'illalvernia, at the Cem- 
eteiy, I sh., Astiau yellow sands, middle-upper Pliocene, 
coll. MFc.v coll. PAL, Fl 18A; Emilia Roniagna, Modena, 
Maranello, Fogliano, Gatiliardella, Rio Grizzasia sands, 1 

\'. Garilli, 2{)()S 

Pasie 35 

sh., middle Pliocene, coll. MP c.v coll. PAL, F;39A; Tus- 
cany, Siena,, X'illa Pictnititta, Podere "La 
Mgna", (SP 36, 4.9 km E side), 7 slis, lower Pliocene, 
coll. MF ex coll. PAL, F55B; Siena, Poggihonsi, \'illa 
Pietrafitta. "Sbarra", (SP 36, 5.2 km), 2 shs, sands, lower 
Pliocene, coll. MF ex coll. PAL, F55D: Siena, Castel- 
nuox'o Berardenga. Terre Rosse, (SS 73, 104 km), 1 sh., 
sands, lower Pliocene, coll. MF c.v coll. PAL, F36A; 
Siena, Poggibonsi, \'illa Pietrafitta. Pt)dere Sant'LHixiere, 
27 shs, lower Pliocene, coll. MF ex coll. PAL, F55C; 
Siena, Pogabonsi, X'illa Pietrafitta, Podere Melom-ani, 6 
shs. sands, lo\\er Pliocene, coll. MF c.v coll. PAL, F55E; 
Siena, Colle \'al d'Elsa, Bibbiano, 2 shs, \cllow sands, 
lower Pliocene, coll. MF ex coll. PAL. FIOSA: Tuscany, 
Li\onio. Tuscan Archipelago, Lsland of Elba, Procchio, 
12 m, Posidonin bed, 2 shs, coll. MF c.v coll. C. Bogi, 
2120PROC; Umbria. Terni. Ficullc, qnarn near Chiani 
i"i\"er, iSS71'l. 1 sh., Cidaris mark' sands, lower Pliocene, 
coO. MF ex coll. PAL, F6SA; Sardinia, Nuoro, San Te- 
odoro, 6 shs coll. PAL; Pngha, Taranto, MSNCS 44744 
(ex 1505), 1974, BDA legit and det.; Sicily, Messina, Mi- 
lazzo. Capo Milazzo, Gala S. Antonio, 12 shs, upper yel- 
low sands, upper Pleistocene, coll. MF c.v coll. PAL, F5A; 
SiciK", Catania, Grammichele, C.da Catallarga, 2 shs, 
coarse sands, lower Pleistocene, coll. MF c.v coll. PAL, 
F27A: Sicih-, Palermo, 1 sh., MNHN coll. Petit; Sicily, 
Palermo, 2 shs, MNHN coll. Dollfus, 1903; Palermo, 
Tommaso Natale, 13 shs, upper Pleistocene, MGUP 167/ 
2/50: Palermo, Addaura, 1 sh.. upper Pleistocene, 
MGUP 587/5/15; Sicilv, Catania, 2 shs, MSNGS 44743 
(c.v 1448), 4/1974. G. Gentile legit and det.; Sicilv. Sira- 
cusa. \'endicari. 1 sh.. coll. PAL,"2124. Adriatic Sea: 128 
shs, MNHN. Croatia: Zara, 8 shs, MNHN coll. Petit, 
1873. Greece: Ev-via Island (Euboea), Louklssia. 4—5 m. 
1 sh., coll. PAL ex coll. Bogi Cesare. 2121LUC. Israel: 
Haifa. 9 m, 1 sh., coll. PAL ex coll. C. Bogi, 1994, 
2 164 HA. 

Habitat: As indicated b\- Jeffrevs (1867), Gofas and 
Ponder (1991), and Bouchet (1978; 1992), this species 
t\picalK" li\"es buried under stones covered with sand at 
\erv shallow waters. I found ven' fresh shells collected 
from along all the upper part of the infralittoral stage. 

Distribution: .\II the .Metliterranean. probaljK' de- 
creasing abundance in eastward direction. In the eastern 
Atlantic it li\es from the British Islands (see also Jeffreys, 
1867; 1869) to Morocco. Alvania lactea has also been 
recorded from the Black Sea (Anistratenko and Star- 
obogato\". 1994). This species probablv originated in the 
Mediterranean Neogene. from where it is recorded from 
the Miocene of north Italv. It becomes more common in 
die western and central Mediterranean Phocene depos- 
its, where it is recorded from se\eral localities of north, 
central, and south Italv (material herein studied; see also 
Sacco, 1895, and Ghirli. 2006) and Spain (Estepona, Lan- 
dau et al.. 2004). In the Atlantic Pleistocene, as reported 
bv Landau et al. (2004). it is recorded from the post- 
glacial of Iceland, the North Sea Basin and the 
Isles. As subfossil, it is recorded from Sweden (Huben- 

dick and W'aren, 1969). In Mediterranean, it is common 
hi)m the lower-upper Pleistocene of SiciK' and is also 
recorded from the lower Pleistocene of Tuscany (Chirli, 

Reinark.s: This is a \eiy variable species especially 
with respect to sculpture and shell shape. The tvpical 
moi"ph, as described and figured hv Michaud (1831: fig. 
12), has a clathrate sculpture with the a\ial pattern domi- 
nating over the spiral one. The finely cancellated ty][3e, 
usually characterized b\' an elongate general shape and 
more convex whorls, corresponds to the forma iiiinor- 
teituiseulpta Monterosato, 1917. Two varieties, semiacos- 
Uitd M\d fiisnlatovaiicosa (the latter often bearing va- 
rices), were described by Sacco (1895) on similar mate- 
rial from the Pliocene of north Italy. This cancellate 
moiph (Figures 44 and 47—18), which seems to be the 
only representation of the discussed species in the Mio- 
cene, is the most common in the Pliocene collections 
studied (see also Sacco, 1895: p. 28; Chirli, 2006: figs. 
9-11 and 15-16) and becomes less common in Quater- 
naiy material. The markedK' ribbed moiph, described bv 
Sacco (1895) as var. laticostata (b'om the Pliocene of 
Italy), and better faiown as iormdclajerleini Monterosato, 
1889, bears strong ribs, starting irom the earlv teleo- 
conch whorls. According to Monterosato (1917), this rep- 
resents the Atlantic moiph oiAlvonia laetea. I agree with 
Van Aatsen et al., considering it not e.xclusively an Atlan- 
tic moiph, being present in some Mediterranean locali- 
ties (see Figure 45). It is noteworthy to remember that in 
a extensive lot (128 shells, MNHN-DSE) from the Adri- 
atic Sea, all the above mentioned morphs coexist. 

Especially when its typical moiphs are considered, Al- 
vania lactea has a veiy characteristic ribbed, cyLindiic;il- 
oxate shell and can not be confused with its congeners. 
Some problem might occur when considering the can- 
cellate moiph, which can be compared to the recently 
described Pliocene .species A. fredianii Delia Bella and 
Scaiponi, 2000 (see this article for the main chfterences 
between the two species). The same moiph of A. laetea 
shows strong similarities with the new species described 
herein, A. rosariae (see discussion below lor differential 

Ahania laetea is the t\pe species oi Ma^isotia Bucqnoy 
et al., 1884, which is considered a swonym of Alvania 
sensu stricto by Ponder (1985) on the basis of the num- 
ber of the metapodial tentacles, shown b\' tliis species, 
based on a description bv Jeffreys (1867). In my (pinion, 
a more exliaustive anatomical dataset could be proxided. 
Nevertheless, even on the basis of its shell features and 
intraspecific variation, there is no reason to consider 
Massotia as a well-supported group. 

Alvania lactea has a veiy compUcated teleoconch mi- 
crosculpture, which could certainly be considered as a 
suite of distinctive characters, not obseived in the 
closely-related taxa discussed in the present report. Early 
teleoconch whorls show a pattern similar to that ob- 
served in A. carinata, A. francescoi new species, and A. 
rosariae new species (detailed comparisons are provided 

Page 36 

THE NAUTILUS, Vol. 122, No. 1 

below), but the remainder of the shell is covered by veiy 
narrow, raised lamella-like axial ridges and veiy thin spi- 
ral threads. Also its protoconch I sculpture is rather dis- 
tinctive, showng a pattern similar to that obsei'ved in A. 
caiinata, A.francescoi, A. rosariae, and A. tcnera (tis well 
as in manv other rissoids), but ha\ing a coarser ornamen- 

Alvania pnisi (P. Fischer, 1877) 
(Figures 50-52) 

Rissoa (Alvaiiia) Pnisi P. Fischer, 1877: 80. 

Description: Shell small, stmxK, conical, partial!)' 
worn off, 4.7 mm in height, 3.2 mm in width. Protoconch 
multispiral, conical, consisting of about little more than 
two convex whorls. Teleoconch is formed by about four 
con\ex whorls, provided with a \en' delicate, cancellated 
sculpture almost giving a pitted shape to shell sui face. 
This sculpture consists of moderately pronounced spiral 
cords crossed bv apparenth' flat iLxial ribs. Spiral coi'ds 
number 7-S on penultimate whorl and 16 on body whoi'l; 
they are flatter and wider on shell base. Ribs, numbering 
about 40 on penultimate whorl, are lacking in basal part 
of last whorl. Sutures slightly inclined and deeplv im- 
pressed. Last whorl well-expanded, rather angulated at 
the base and inflated, comprising % of sliell height. 
Aperture wide, ovate to almost pyriforme, pointed adapi- 
cally, comprising Vi of the total height, % of last whorl 
height. Outer lip prosocline, internally smooth, externalK 
with an almost flat, wide thickening, apparently smooth. 
Inner lip moderately arcuate, with a modest and narrow 
reflection on columellar area, leaving a X'eiy small and 
narrow umbilical chink (filled up with sediment). 

Type Locality: Isle of Rhodes 

Type Material: One syntyi^e, R07495 in MNHN- 
DIIT, from t\pe locality. 

Material Examined: Eastern Mediterranean Sea, 
Quaternary of Rhodes, type locality, MNHN-DHT 
(R07495), 1 sh. 

Habitat: It is veiy difficult to characterize the paleo- 
ecological significance of this extinct species since no 
data about its paleoenvironment are directly available or 
deducible. Furthermore, Fischer (1877) provided no 
precise indication about the site, stratotype, and its sedi- 
mentologica! and paleontological nature. Considering all 
the other molluscan species described from Rhodes by 
the same author, a shallow water depositional environ- 
ment (linked to the luodern phanerogam infralittoral 
bottoiu) can be inferred. 

Distribution: Tlie species is known onK hom the t\pe 
locality, and its distribution appears to be limited to the 
Quaternaiy of Rliodes. 

Remarks: This taxon is practicall)- unknown. To mv 
knowledge, the most recent treatment is that of Montc- 

rosato (1917), who considered it as a separate species 
belonging to the Massotia group. 

Alvania pnisi could be confused with juvenile shells of 
Alvania lactea (the form with cancellate sculpture, see 
Figure 49), which exhibit a vei-y similar body whorl pro- 
file. Alvania pnisi is quite close to A. fredianii Delia 
Rella and Scaiponi (2000) from the Pliocene of Tuscany. 
Both species share the general shell shape and the finely 
reticulated sculpture, but the latter is of smaller size, has 
a paucispiral protoconch, deeper sutures, and a weakly 
denticulated outer lip (see Delia Bella and Scaiponi, 
2000: pis. 1 and 2). 

Alvania rosariae new species 
(Figures 62-79) 

Description: Shell small, sturdy, conical-ovate to tur- 
riforin, moderately to markedly inflated, reaching about 
5 mm in height, 3.5 mm in width (holotvpe 4 mm in 
height and 2.7 mm in width). Protoconch multispiral, 
conical, consisting of about 2-2.1 convex whorls. Proto- 
conch I of about 0.8 whorls sculptured by 5-6 very fine 
spiral lirae irregularly interspersed with microscopic 
granules. Protoconch/teleoconch transition well marked 
and sinuous adapicallv. Protoconch II sculptured by spi- 
rally arranged microscopic pimples (stronger in adapical 
portion) forming one to two spiral threads, the lower very 
close to the suture, occurring on last whorl. Teleoconch 
formed by about 4.2 usually vew convex whorls, sculp- 
tured by numerous axial ribs and slightly stronger spiral 
cords. The latter, rapidly increasing in count, number 
2-6, 6-10 (rarely ll), and 26-34 (in specimens higher 
than 3.5 mm) on the first, penultimate, and last whorl of 
the teleoconch, respectively. Some secondary, less 
marked cords may occur on last whorl, usually close to 
the outer lip. The adapical one, two, and three spiral 
cords on first, penultimate and body whorl, respectively 
are more pronounced. In specimens higher than 3.5 mm, 
axial sculpture consists of 44-60 narrow ribs (52 in ho- 
lotvpe) on the penultimate whorl. Ribs become narrower 
and lamella-like to obsolete or lacking on shell base, par- 
ticularly close to columella. The same may occur on the 
terminal portion of the body whorl (Figure 66). The in- 
tersection of spiral cords with axial ribs gives a charac- 
teristic cancellate and gently nodular shape, forming a 
rectangular (almost equilateral in the central portion of 
tlu' body whorl) pattern. The long axis of these rectangles 
is peipendicular to the shell axis on the central and 
al^apical portion of whorls and parallel on upper part. 
Microsciilptiire consists of veiy fine, sometime irregu- 
larly interspersed, spiral lirae (Figure 77). These become 
obsolete on the penultimate aiul body whorls, where nu- 
luerous, \eiy fine and narnnv lilis (possible growth lines) 
occui'. fjirae are continuous only on adapical portion of 
earlv teleoconch whorls and never cover piimaiy spiral 
sculpture. Sutures slightly inclined and rather deeply 
impressed. Last whorl well-developed, comprising about 
% to Vs of the shell height, usually with a veiy convex 
profile. It sometime bears one or two close varices which 

\". Garilli, 2008 

Page 37 

ma\- tonii angles of 10° to 210'^. Apertiiic wide, o\'ate, 
comprising Vi to % ot the total heiglit; % to % of last 
whorl height. Outer lip orthochne (slightl)' cuned), in- 
ternalh' smooth, externallv markedly thickened close to 
lip edge and co\erei! b\' spiral cords. Inner lip moder- 
ateK' arcuated and rather thickened in the cokmiellar 
area, where a \en' narrow uiiihilic'al chink occurs. 

Tj-pe Localitv: Lower Pleistocene ot Cartiera Mulino 
(3'6°56'57" N,'l4°34'03" E), Mttoria, Ragusa, southeast- 
ern Sicilv. The stratot\pe is the 3D1 layer of Costa 

Tvpe Material: Holot\pe (4.0 x 2.7 mm), DGUP 
C'MRG 12/496, coll. GarilU; Parat^pe 1 (4.6 x 3.2 mm), 
ZM.\ Moll. 4.07.014 (ex CMRG' 13/497 coll. Garilh); 
paratope 2 (not-complete shell, 3 mm width), ZMA Moll. 
4.07.08 {ex CMRG 014/498 coll. GariUi); paratype 3 (3.6 
X 2.55 mm), DGUP CMRG 15/499, coll. V. GariUi; 
paratApe 4 (4.5 x 3.3 mm), DGUP CMRG 16/500, coll. 
Garilli: parat\pe 5 (3.8 x 2.5 mm), MNHN A25951 {ex 
CMRG 17/501, coll. Garilli); parat\pe 6 (3.85 x 2.75 
mm), ZMB MB. Ga. 2480 {ex CMRG 18/502, coll. 
Garilli); pai-at\pe 7 (3.6 x 2.4 mm), ZMB MB.Ga.2481 
(e.v CMRG 19/503, coll. Garilli); parat\pe 8 (3.8 x 2.6 
mm), DGUP CMRG 20/504, coll. Garilli; paratype 9 (3.6 
x 2.6 mm), DGUP CMRG 21/505, coll. Garilli; paratype 
10 (3.95 -x 2.6 mm), DGUP CMRG 22/506, coll. Garilli; 
parat\pe 11 (4.2 x 2.75 mm), DGUP CMRG 23/507, 
coll. Garilli; paratvpe 12 (3.7 x 2.55 mm), DGUP CMRG 
24/508. coll. Gai-iih; paratype 13 (3.9 x 2.5 mm), DGUP 
CMRG 2.5/509, coll. Garilli; paratype 14 (3.7 x 2.55 mm), 
DGUP CMRG 26/510, coll. Garilli; paratxpe 15 (3.5 x 
2.6 mm), DGUP CMRG 27/511, coll. Garilli; paratype 16 
(4 X 2.6 mm), DGUP CMRG 28/512, coll. Garilh; 
parat^pe 17 (3.9 x 2.7 mm), DGUP CMRG 29/513, coll. 
Garilli; paratope 18 (3.6 x 2.45 mm), GNHM ID 30.706 
{ex KIGR .3/514 coll. Garilli); parat\pe 19 (4 x 2.7 mm), 
GXHM ID 30.707 [ex KIGR 4/515 coll. Garilh); 
parat^pe 20 (4.5 x 2.7 mm), GNHM ID 30.708 {ex KIGR 
5/516 coll. GariUi); paratype 21 (4.7 x 3.4 mm), GNHM 
ID 30.709 [ex KIGR 6/517 coll. GariUi); paratvpe 22 (4.4 
X 2.9 mm), GNHM ID 30.710 {ex KIGR 7/518 coU. 
GariUi); paraKpe 23 (4 x 2.55 mm), GNHM ID 30.711 
{ex KIGR 8/519 coll. GariUi); paratype 24 (3.35 x 2.5 
mm). GNHM ID 30.712, {ex KIGR 9/520 coU. GarilU); 
paratvpe 25 (2.9 x 1.95 mm), GNHM ID 30.713 {ex 
KIGR 10/521 coU. GarilU); paratype 26 (not-complete 
sheU), GNHM ID 30.714 {ex KIGR 11/522 coll. Garilli); 
parat\pe 27 (4.15 x 2.65 mm), GNHM ID 30.715 {ex 
KIGR 12/523 coU. GarilU); paratvpe 28 (4.0 x 2.6 mm), 
GNHM ID 30.716 {ex KIGR' 1.3/524 coll. Garilli); 
paratvpe 29 (not-complete shell, 3.85 mm), GNHM ID 
30.717 (e.v KIGR 14/525 coU. GarilU); paratvpe 30 (2.7 x 
1.9 mm), GNHM ID 30.718 {ex KIGR' 15/526 coU. 
GarilU): paratvpe 31 (3 x 2.05 mm), GNHM ID 30.719 
{ex KIGR 16/527 coU. GariUi): paratvpe 32 (3.8 x 2.5 
mm), GNHM ID 30.720 (ex KIGR 17/528 coU. GarilU); 
paratvpe 33 (3.55 x 2.5 mm), GNHM ID 30.721 {ex 

KIGR 18/529 coll. Garilli); panitvpe 34 (4.05 x 2.8 mm), 
GNHM ID 30.722 (ex KIGPi 19/530 coll. Garilli); 
paratvpe 35 (4.95 x 3.25 mm), GNHM ID 30.723 {ex 
KIGR 20/531 coU. GariUi); paratype 36 (not measured), 
GNHM ID 30.724 (ex KIGR 21/532, coll. GariUi); 
paratvpe 37 (not measured), GNHM ID 30.725 (e.v 
KIGR 22/533, coll. Garilli). Holotyi^e and Paratvpes 
4-17, from the lower Pleistocene of Cartiera Mulino, 
3D1 bed of Costa (1989), Vittoria, Ragusa, southeastern 
Sicily. Paratypes 1-3 from the same loc;ilitv, 3D2 bed of 
Costa (1989). Parat)pes 18-22 and 24-35 from the 
middle to upper Pleistocene of Kyllini, northwestern 
Peloponnesus, N2 and H6 beds of Garilli et ;il. (2005a), 
respectively. Paratypes 36 and 37, same locality, from the 
lower to early middle Pleistocene P3 layer of Garilli 
(2005b) and from a late lower Pleistocene yellowish to 
reddish sandy layer about 50 m underlying the F14 bed 
of Garilli and Galletti (2007), respectively. 

Another four, uncatalogued paratypes are housed in 
DSTC (1 sh. from 3D1, 1 sh from 3D2 and 2 shells from 
3C bed of Costa, 1989, all from the lot n° 18, as Gcdeo- 
clina carinata (Da Costa). One more uncatalogued 
paratype (e.v CMRG 030/534 coll. Garilli), from the'tvpe 
locaUtv', 3D1 layer, is in coll. MF (Prato). 

Etymology: The species is dedicated to m)- wife Ro- 

Material Examined: The tvpe material from the 
lower Pleistocene ot Cartiera Mvdino, Vittoiia, Ragusa, 
SE Sicily, 18 shs, and from the late lower (1 sh., from a 
reclcUsh to vellowish sandy bed about 50 m underlying 
the F14 bed of GariUi and Galletti, 2007), lower to 
middle (2 shs, layer P3 of GarilU et al, 2005b), and 
middle to upper Pleistocene (13 shs, layer H6; 4 shs, 
layer N2) of KylUni, Elea, NW Peloponnesus, Greece. 

Habitat: In the type locality, the species was mainly 
recovered from the layers 3D1 and 3D2 which were 
linked to the ecotone SVMC-HP (sensu Peres and Pi- 
card, 1964) by Costa (1989). In the Kyllini sites, this 
species was found in cerithids-trochids-rissoids assem- 
blage linked to the present biocenosis HP, characterized 
bv the phanerogam Posiclonia oceaniea (Linnaeus) De- 
lile, 1813 (See GarilU et al. (2005a); Garilli et al. (2005b), 
and Garilli and Galletti (2007) for more detailed infor- 
mation about the paleoecological characteristics of the 
cited Kyllini strata.) 

Distribution: The species has a lower to miildlc-upper 
Pleistocene stratigraphic range, presently limited to SE 
Sicily and NW Peloponnesus, being recorded fi-om the 
lower Pleistocene of Sicily (type locality) and from the 
lower to upper Pleistocene deposits of Kyllini, Greece 
(NW Peloponnesus). 

Remarks: Ahania rosiiriac represents a sort of inter- 
mediate form between A. carinata and A. lactca (cancel- 
late form). Compared with the former tavon, it shows 
some similarities in the t)pe of intraspecitic \ ariation, 
showing very inflated to quite elongate shells (compare 

Pase 38 

THE NAUTILUS, Vol. 122, No. 1 

\'. Garilli. 2()()-S 

Page 39 

Figures 68-69 \\'itli tin- "lonn " ccarhuita ol .A. ((ihixitci. 
Figures 1, 4, 5), in usualK' bearing \arices, aiul in the 
niicrosculpture of the earl\' teleoconcli. In addition, their 
i"especti\'e lanal shells do not show rele\ant dilTei-ences. 
Alvania rosaiiuc new species can be distingnished b\' 
absence ot a keeled siiape, having siibequal and more 
numerous spiral and axial sculptural elements. The can- 
cellate form of .A. lactea is comparable wth A. rosariac 
new species, haxing a \en' similar sculpture and occa- 
sionalK" a similar shell shape (onlv in the \en o\ate 
morph, e.g. Figm"es 44, 47 and 65-66). Howe\ei'. A. m- 
sariae new species ditiers h\- ha\ing moi'e convex whorls 
and a \'en' different niicrosculptural pattern, with only 
irregular, fine, often interrupted spiral threads covering 
the earl\" teleoconch whorls. Furthermore, the proto- 
conch I of A. hicfca bears a coarser sculptiu-e, consisting 
of more numeorns and larger pimples and iiiutli more 
elevated spiral lirae. 

Alvonia rosariac new species can also be compared 
with A. francescoi new species: both species ha\'e a ga- 
leodiniform shape, a similar niicrosculpture, and share a 
\en' similar sculptural pattern on the protoconch. Dif- 
ferences between the two species are mainly found in the 
teleoconch sculpture, which is strongly nodular and 
coarser in A. francescoi new species; furthermore the 
latter has fewer spiral cords and axial ribs. 

Alvania rosariac new species may resemble A. magis- 
tra Chirli, 2006. an interesting galeodiniform species 
from tlie Pliocene of N Italy and's Spain (Chirli, 2006: pi. 
I f , figs.13-16 and pi. 12, figs. 1-8; Landau et al, 2004: pi. 
10, figs.3a-e, as Alvania sp.), showing a quite similar 
finely cancellate sculptm-e, especially in the arrangement 
of die adapical cords, and a quite wide, ovate aperture. 
However, the latter species shows an miusual outer lip 
profile, having a wide sinus at the upper aspect, its pro- 
toconch I shows a netted Manzonia-like sculpture, while 
protoconch II appears less sculptured than in A. rosariac 
new species 

Ahania tencra (Philippi. IS44) 
(Figures 80-S9) 

Rissoa teiwra Philippi, 1S44: 128-129, pi. 2.3, fig. 15. 

Qaleochna tcnera (Philippi, 1844). — Piaui, 1979: 71, 

Description: Shell minute, conical, and subcarinate, 
to conical-oxate or elongate, moderately to markedly 
sturdy, reaching 2.2 mm in height and 1.4 mm in width. 
Protoconch multispiral, conical, with 2-2.3 convex 
wliorls. Protoconch I consists of about 0.8 whorl, sculp- 
tured by SLx \ery tlim spiral lirae and a few microscopic 
granules between them. Protoconch II is sculptured by a 

few til :iliuii(hint \vy\- small granules. These are more 
nimierons o\\ last half whorl, where the\" are spirally ar- 
ranged, forming veiy discontinuous and irregular ridges. 
Protoconcli/ teleoconch transition distinct, with a slight 
(to veiy slight) sinuosit)'. Teleoconch consists of 3-3. .5 
weakly to discretely convex whorls, quite variable in 
width. These are sculptured by well-raised, narrow, spi- 
ral, occasionally almost k'eel-like cords (in quite conical 
shells), which overide the axial sculpture. They number 
.3-4, 4-.5, and 12-13 on first, penultimate, and body 
whorl, respectively. Usualh', weaker cords are present on 
shell base or close to upper suture. Axial sculpture 
formed by veiy nai'i'ow, prosocline, rows ol sliort seg- 
ments, forming discontinous ribs, occurring between spi- 
ral cords and becoming xen' thin to lacking toward shell 
base. Ribs number 40-60 on last whorl. At the intersec- 
tions with axial elements, spiral cords generally appear 
very finely nodular (Figure 86). Last whorl well- 
expanded, with a rather rounded profile, sometimes in- 
flated, comprising about % to '/t (in subcarinate and 
conical shells) of total sliell height. Aperture ovate, an- 
teriorly rounded, posteriorly angulated, comprising 
about Vi and % of total shell and last whorl height, re- 
spectively. Outer lip roimded, markedly prosocline, thin, 
internalK' smooth, externally with no thickening. Inner 
lip weakly arcuate, witli a thin and narrow (Figuj'e 82) to 
stronger and wider (Figure 80) callirs, leaving a \'ei-\' nar- 
row umbilical chink. Coloration usually consists of red- 
dish to brown spots on a crcaiii-wliitish or (rarely) 
brownish background. 

Type Locality: Peuiusida of Magnisi (originally indi- 
cated as "Peninsula Thapsum ' from the old Cireek name 
of Magnisi), Siracusa, southeastern Sicily. 

Material Exaininecl: Atlantic: Canaiy Islands, Tene- 
rife, Pal-Mar. 6-8 m, 1 sh., MNHN coll. P. Bouchet, 15 
Jul. 1980; Morocco: Asilah, mouth of Oued el Helou, 
concliiferous detritirs, beach, 9 shs, MNHN coll. S. Go- 
fas, 1971-72; EI jadida, (33°16' N, 08°29' W), large 
beach, conchiferous detritus, beach, 5 shs, MNHN rec S. 
Gofas, 26 Sep. 1991; El Jadida, (.33°16' N, 08°29' W), 
large beach, tide zone, 4 shs, MNHN rec S. Gofas, 26 
Sep. 1991. Strait of Gibraltar: Spain, Cadiz, conchiferous 
detritus, beach, 2 shs, MNHN coll. S. Gofas, 1976-81; 
Cadiz, Tarifa, beach, 4 shs, coll. PAL ex coll. C. Bogi, 
Jun. 1986, 2030TAR; Cadiz, Taiifa, Torre de la Pefla, 
conchiferous detritus, beach, 3 shs, MNHN coll. S. Go- 
fas, Aug. 1981; Vlorocco, Tanger, Grande Plage, conchif- 
erous detritus, beach, 1 sh., MNHN coll. S. Gofas, 1970- 
81. Mediterranean: Spain, Andalusia, Punta della Mona, 
43 m, 5 shs, coll. PAL ev coll. C. Bogi, 2032PMO; Spam, 
Malaga, industrial dredging, 20^0 m, 1 sh., MNHN rec. 

Figures 62-69. Ahania rosariac new species, holotvpe and parat\pes 16, 21, and 3.5. 62-63. Apeitural (62), dorsal (63) and prolile 
(64 1 \ie\vs of holoUpe. lower Pleistocene of Cartiera Mulino, bed 3D1 of Costa (1989). SE Sicily, Ragusa, Vittoria. 65-66. Parahpe 
16, dorsal (65), illustrating \arices on the last wliorl, and apertural (66) \ie\v, same site and localit)-. 67. Faratvpe 21, a veiy inflated 
last wliorl and a strong varix opposite the outer tip, middle to upper Pleistocene of Kyllini, NW Peloponnesus, N2 bed of Garilli et 
al. (2005a). 68-69. Apertural (68) and dorsal (69) view' of paratvpe 35, showing a quite .slender shell, niiddl' to upper Pleistocene 
of K\-Uim. noitliwestem Peloponnesus, H6 bed of Garilli et al (2005a). Scale bars 100 jj-ni. 

Page 40 

THE NAUTILUS, Vol. 122, No. 1 

V. Garilli, 20()S 

Page 41 

S. et C. Goias, May 1991; Malaga, t:alaliuiKla, conchif- 
erous detritus, beach, 1 sh., MNHN coll.S. Gofas, 1976- 
81; Malaga, Benalniadena, conchirerous detritu.s, beach, 
6 shs, NfNHN rec S. Gofas, 1991-93; Malaga, Mijas, 
detritus, 10 ni, 4 shs, coll. Stetauo Ruhui; Malaga, Gabo 
Pino, detritus, 10 m, 2 shs, coll. SR, (41.S0g); Morocco, 
Gabo Negro, beach, 1 sh., MNHN coll. S. Gofas, det. W. 
Pouder, 19S6; France, Provence, Marseille, Curry, 
beach, .3 m, 2 shs, coll. PAL ex coll. C. Bogi, Tun. 1986, 
2037M.\R; FroNence, Marseille, La Baule, small beach at 
25 km w^est from Marseille, 9 shs, coll. PAL ex coll. G. 
Bogi, Oct. 1986, 2035BAU; Provence, Le Dramont, 
(43'^°24.7' N, 6°51.7 E), 22-30 m, 26 .shs, MNHN rec. J. 
Pelorce, 1992; Pro\-ence, Les Enibiez, cote Nord et Petit 
Rom-eau, (43°05' N, 5°47' E), rocks, algae, 0-1 m, 1 1 
shs, MNHN rec S. Gofas, Jun. 1995; Provence, 
Marseille, Gap Morgiou, "caique de la Triperie," 
(43°12.2' N, 05°26.9' E). mudcK- sand , inside cavit)', 22 
m, 4 shs, MNHN rec H. Zibrowius, }tm. 1996; Provence, 
Marseille, Grand Gongloue, {43°10.6' N, 05°24.2' E), .33 
m, 50 shs, MNHN^■ec. H. Zibrowius, Jun. 1996; 
Provence. Les Embiez, passe du Gaou, rocks, photophile 
algae, (43°04.3' N 5°47.4' E), 0-3 m, 1 sh., MNHN rec 
S. Gofas, Jun. 1995; Provence, St. Glair, infralittoral 
rocks, (43°08.2' N 6°23.2' E), 0-1 m, 1 sh., MNHN rec. 
S. Gofas, Sep. 1992; Tunisia, Sfax, 4 shs, MNHN coll. 
Staadt, 1969; Italy, Liguria, Portofino, 1 sh., coll. PAL, 
2038; Italy, Tuscanv, Livomo, Antignano, 0.5 m, lirowii 
algae on rocks, 1 sh., coll. PAL legit Bogi, Apr. 1999, 
2029; Livomo, under littoral rocks, 0.5-1.0 m, 5 shs, coll. 
PAL ex coll. G. Bogi, 2039LIV; Livorno, Meloria, 10-30 
m, 3 shs, coll. PAL ex coll. G. Bogi, 1995, 2031MEL; 
Li\omo, Tuscan Archipelago, Island of Elba, Gapoliveri, 
32 m, 2 shs, coll. PAL ex coll. G. Bogi, Aug. 1994, 
2036CAPOL; Tuscan Archipelago, Gemini Islanti, 
(southern side of Island of Elba), 11 m, 3 shs, coll. PAL, 
2034; Tuscan .Archipelago, Isola del Giglio. Punta 
Fenaia, 32 m, Ish., coll. c.v coll. G. Rug- 
gieri; Tuscany, Grosseto, Argentario, 25 m, 1 sh., coll. ex coll. G. Ruggieri, Jul. 1988; Italy, 
Lazio, Ostia, Tor Patemo, 33 m, 5 shs, coll. medshells. 
com ex coU. G. Ruggieri; Lazio, Roma, Santa Marinella, 

on ' 

ex reti, 2 shs, coU. ex coll. G. Ruggieri; 
Italv, Sardinia, Oristano, Santa Gaterina di Pittinurri, 5 
m, '3 shs, coll. PAL ex coll. G. Bogi, 30 Aug. 1986, 
2040SGP; Sardinia, Sassari, Island ot Maddalena, beach, 
3 shs coll. PAL ex coll. C. Bogi, 2028IMA; Sardinia, 
Nuoro, Sant'Antioco, Gala Lunga, 20 shs, coll. med- 
shells. com ex coll. G. Ruggieri, 03.1989; Italw Gam- 

pania, i'ciiinsula ol Sorrento, Punta Penna, 2 shs, coll. ex coll. G. Ruggieri, 1988; Italy, Sicily, 
Palermo, 2 shs, MNHN coll' Locard; Sicily, 1 sh., 
MNHN coll. Petit, 1873; Palermo, 3 shs, ZMB ex coll. 
Monterosato, 81013 (originally labelled b)' Monterosato 
as Cingida tenera, 3, 1125, Palermo); Palermo, Arenella, 
2 shs, ZMB c.v coll. Monterosato, 81014, (originally la- 
belled by Monterosato as "Cingida teitera var ... ,2, 
Arenella, Palermo'"); Sicily, Siracusa, Capo Passero, 16 
m, 3 shs, coll. mc< c.v coll. G. Ruggieri, 05 Sep. 

Habitat: This species is clearlv limitetl to infrtdittoral 
depths. In the upper part of its distribution, it seems to 
live in veiy shallow waters, on algae. It likely lives also in 
the cavities occurring in infralittoral muddv sandv bot- 

Distribution: In the western and central Mediterra- 
nean tile species seems to be well distributed; Adriatic 
and eastern Mediterranean occurrences should be veri- 
fied. In the Atlantic it li\'es along the Moroccan coasts 
and in the Canan Islands. To my knowledge, there is no 
fossil record of this species. 

Remarks: This small species is characterized by having 
a variable shell shape and sculpture. The conical shells, 
bearing a strongly cingulated sculptiu-e, which markedly 
prcN'ails over the axial, are comparable with t\pical 
keeled moi-ph of Alvauia carinata. In addition, Cingida 
species provided with a strong spiral sculpture are com- 
parable to A. tenera. The ovate, slender shells of this last 
species, with a finely cingulated sculptural pattern, might 
vaguely resemble some species of Setia H. and A. Adams, 
1854. As a consequence, Piani (1979) and Van Aartsen 
(1982) included A. tenera in Galeodina, whereas Nords- 
ieck (1968; 1972) placed the same species in Setia and 
Cingida i-espectiveK. The last two views should not be 
accepted. In fact, the species of Setia have a smooth or 
veiy slightly sculptured shell with more convex whorls 
than A. tenera, and Cingida species are characterized by 
shells usually lacking axial sculpture and ha\ing a very 
thick outer lip. 

The shells from eastern Atlantic (Ganaiy Islands and 
Morocco) do not show meaningful differences from the 
Mediterranean ones studied. With regard to the resem- 
blance between A. tenera and A. carinata, it is manifest 
in the keeled, conical-inflated shell shape shown by their 
typical respective moi"phs, and in having a quite wide and 
de\elnped apeiinie. However A. tenera never shows \'a- 

Figures "0-79. Ahiiiiid rosariae new species 70. Protoconch of the liolot\pe, lower Pleistocene of Caitiera Mulino, bed .3D1 of 
Costa (1989), SE Sicilv, Ragusa, \ittoria. 71. Protoconch of paratype 16, showing variation of the abapical ridges on protoconch II, 
same locality and bed. 72. Dorsal view of protoconcli of paratype 21, showing protoconch I sculpture and the afiapical ridges on 
protoconch II, middle to upper Pleistocene of Kyllini, NW Peloponnesus, N2 bed of Garilli et al. (200.5a). 73-74. Holotype, detail 
of earl\- protoconch (73) showing sculpture of protoconch I and protoconch I/protoconch II boundary, and dorsal view of protoconch 
(74). 75. Sculpture of protoconch I, paratype 21. 76-79. Holotype, detail of teleoconch sculpture; early whorls (76), first to second 
whorl (771, showing; microsculpture, penultimate to last whorl (78), showing the microscopic incremental scars, and last whorl (79). 
Scale bars: 100 (jim in Figures 70-72, 74, 76 and 78-79; .50 [x,m in Figure 77; 20 (xm in Figure 75. Black and white arrows indicate 
the protoconch I/protoeonch II and protoconclVteleoconch l^oundaries, respectively. 

Page 42 

THE NAUTILUS, Vol. 122. No, 1 

V. Garilli, 2008 

Page 43 

rices, bears a \'en ilclicatr axial sfiilpdirr, its outer lip is 
alwa\'s \ei"\ tliin, aiul iisualK has more eoloied shells. 

C^.enus GaIc(>(liiiopsi.s Saeco, 1(S95 

T^pe Species: Rissoa tihrriiiiiti ("oppi, ISTtt h\ origi- 
nal designation. 

Galeodinopsis tibcriana (Coppi, 1876) 
(Figin-es 81-99) 

1862 Rissoci tuba Doderlein, 1862: 17 {nomen nudum) 

1876 Rissod Tibcriana Coppi, 1876: 201-202. 
Manzoniafariai Roldii and Fernandes. 1990: 64-65, pi. 1, figs. 

Alvaninfariai iRolan and Ft-rnandes, 1990).— Gotas, 1999: 88- 

89, figs. 39-42. 
Alvania fariae (Rolan and Feniandes, 1990). — I.andan et al., 

2004: 41, pi. 7, figs. 3^. 

Description: Shell eonieal, sturdy, reaching 3.75 nnri 
in height and 2.75 mm in width. Protoconch multispiral, 
conical, with 2-2.2 convex whorls and a rather immersed 
nucleus. Protoconch I consists of about 0.7-0.8 whorls, 
wdtli a netted sculpture, consisting of 7-S veiy thin spiral 
lirae and numerous, irregular, short and ven' nai'row imal 
segments occurring in interspaces beUveen lirae. Proto- 
conch II is sculptured by very small, sparse granules, 
fused into 2—1 discontinuous ridges on central and abapi- 
c;il portions of latter part of last whorl. Groups of gran- 
ules form \'erv short, prosocliiie segments on central 
area, mainly close to protoconcli/teleoconch transition, 
wliich is marked by a quite sinous and thin lip. Teleo- 
conch formed hx 3-3.5 convex whorls, sculptured by a 
priman- pattern of strong, slightly sinuous and opistho- 
chne ribs, numbering 12-14 on penultimate whorl, be- 
coming \-er\' weak to lacking toward shell base. Ribs are 
cox'ered bv a secondary spiral sculptrn-e, formed b\' flat 
naiTower cords, numbering 4-5 on pemdtimate whorl, 
becoming more marked on shell \yAse. Each cord bears a 
pitted microsculpture, consisting of microscopic subcir- 
cular pores forming 8-10 spiral alignments (Figures 99). 
Between cords, numerous, closely spaced, veiy fine spi- 
ral ridges appear. They are formed by rough prismatic, 
tooth-like elements, extending perpendicularK- from 
shell surface. At their base, ridges have a lamella-like 
e.xpansion which covers the interspaces occurring be- 
tween them. Last whorl inflated, well-expanded, com- 
prising about % of total shell height, often bearing 1-2 
strong \arices, most frequentK" just before the outer lip. 
Aperture ovate, \"ery rounded anteriorly, weakly angu- 
lated posteriorly, comprising about little less tlian Vi and 

% of total shell and last wlioii height, respectively. Outer 
lip sinuous, weakly opisthocline; internally smooth, with 
a thin rim on its edge; externally with a veiy marked 
varicose swelling, covered by spiral sculpture, and having 
a narrow ridge on its base, toward aperture, so that it 
appears double-rimmed. Inner lip weakly cui^ved, with a 
veiy narrow columellar thicknening, forming an obsolete 
umbilical chink. Shell color white in Recent material. 

Type Lociililv: (^oppi (1876) cited "La Tagliata", an 
unknown! name in toponymy (veiy likely referring to a 
recently deforested woodland) corresponding to tlie lo- 
calit)' Gagliardella {Maranello, Modena, Emilia Roma- 
giia. North Italy) (S. Palazzi personal comm., 2006). 

T^pe Material: A lot (IFUM 13721), from Coppi coll, 
tspe localit)', with more than 100 possible svnt\pes (not 
seen), is housed in the MPOB, Modena. 

Material E.vaniinetl: Senegal: Region de Dakar, 250 
m S.W. Cap Manuel, 12 shs, MNHN coll. Marche- 
Marchad, det. S. Gofas [The Nautilus 113: 88-89, figs. 
40, 42]; Region de Dakar, S.W. Goree large Cap Vert, 
250-150 m, 1 sh., MNHN coll. Marche-Marchad, det. S. 
Ciofas [The Nautilus 113: 88-89, figs. 40, 42]: Region de 
Dakar, St. 56-1-lOA Goree 150-200 m, 1 sh., MNHN 
coll. Marche-Marchad, det. S. Gofas [The Nautilus 113: 
88-89, figs. 40, 42]; Angola: Luanda, Ilha de Luanda, 
Circalittoral, 120 m, 3 shs, MNHN coll. S. Goflrs, det. S. 
Gofas [The Nautilus 113(3): 88-89, figs. 40, 42]; Luanda, 
Ilha de Luanda, circalittoral, 40-60"m, 1 sh., MNHN 
coll. S. Gofas 1981-82, det. S. Gofas [The Nautilus 113: 
88-89, figs. 40, 42]; Luanda, Au large de Mussulo 
(Mococo), dredging 50-70 m, 2 shs, MNHN coll. S. Go- 
fas 1981-1987, det. S. Gofas [The Nautilus 113: 88-89, 
figs. 40, 42]; Au large de Mussulo, circalittoral, 90-100 
m, 2 shs, MNHN coll. S. Gofas, det. S. Gofas [The Nau- 
tilus 113: 88-89, figs. 40, 42]; Ambrizete, dredging, 80 m, 
5 shs, MNHN colt S. Gofas, det. S. Gofas [The Nautilus 
113: 88-89, figs. 40, 42]; Ambrizete, (07°00' S, 12°20' E) 
sediment, 60 m, 3 shs, MNHN coll. S. Gofas, 1983, det. 
S. Gofas [The Nautilus 113: 88-89, figs. 40, 42); Am- 
brizete, (06°57' S, 12°23' E), sediment, 45 m, 1 sh., 
MNHN coll.S. Gofas, 1983, det. S. Gofas [The Nautilus 
113: 88-89, figs. 40, 42]. Italy: Piemonte, Asti, Baldich- 
ieri, "Grottino Monale", 2 shs, yellow sands, middle- 
upper Pliocene, coll. MF c.v coll. PAL, F104A; Asti prov- 
ince, 9 shs. Pliocene, MGUP coll. Doderiein, UlA; Lig- 
una, Savona, Rio Torsero, between Ceiiale and Peagna, 
5 shs, clays, lower Pliocene, coll. MF ex coll. PAL, F58A; 

Figures 80-89. Alvania tcncra (Philippi, 1844). 80. Shell of a keeled and conical iiioipli. Strait of Gibraltar, Spain, Cadiz, Tarifa, 
coll. P.\L (2030T.'\R). 81-82. Conical-ovate, not-keeled moiphs, illustrating the variable sculpture, France, Provence, Marseille, 
Cum-, coll. PAL (2037NL\R). 83. Profile view of the same shell as Figure 80. 84. Profile view of an unkeeled moiph, France, 
Provence, Marseille, La Baule, small beach at 25 km west from .Vlarseille, coll. PAL (2035BAU). 85-86. Sculpture of early teleoconcli 
whoris (85) and detail of sculpture (86), Italy, Tuscany, Livorno, Antignano, coU. PAL (2029). 87-88. Protoconch (87) and detail of 
protoconch I (88) of die same shell. 89. Protoconch of the same shell as Figure 84, showing sculpture variation. Scale bars: 0.5 mm 
in Figures 80-84: 200 )xm in Figure 85; 100 [xm in Figures 86-87 and 89; 20 p.m in Figure 88. Black and white arrows indicate the 
protoconch Fprotoconch II and protoconcli/teleoconch boundaries, respectively. 

Pase 44 

THE NAUTILUS, Vol. 122, No. 1 

\'. Garilli. 2008 

Page 45 

Emilia Romasna, M 

t)tlena pnnince. 

.35 slis, PI 

coll. Doderlein, lllB; Emilia Roiiiagiia, Modeiia pro\- 
ince, 1 sh., Miocene, MGUP coll. Doderlein, 474; Emilia 
Romagiia, Piacenza, Lugagnano ^'al d'Arda, 2 shs, "calan- 
chi di valle" (marls), middle-upper Pliocene, coll. MF c.v 
coll. PAL, Fl;3A; Piacenza, CastelFArquato, Monte Pa- 
do\a, 1 sh.. blue cla\s, middle Pliocene, coll. MF ex ct)ll. 
PAL, F16A; Emilia Roniagna, Parma, San Nicomede, 
Stirone river, 50 shs and fragments, lower clays middle- 
upper Pliocene, coll. MF ex coll. PAL, F14B; Emilia 
Romagna, Modena, Marano on the Panaro, Panaro river, 
1 sh., cla\s storni\' la\ers, middle-upper Pliocene, coll. 
MF ex coll. PAL, FSO.'V; Emilia Roma2;na, Modena, Ma- 
ranello, Fogliano, Gagliardella (type locality), Rio Griz- 
zaga, 60 shs, sands, middle Phocene, coll. MF ex coll. 
PAL, F39A; Tuscany, Siena, San Donato, Ciuciano, 
Prison, 1 sh., clavs and sands, lower Pliocene, coll. MF ex 
coll. PAL, F112A; Siena, Castiglioncello del Trinoro, 
Poggio Rotondo, 3 shs, marls, lower Pliocene, coll. MF ex 
coll. PAL, F54A: Lazio, Rome, Vlagliano Sabina, Cla- 
docora >ellow sands, 23 shs, lower Pliocene, coll. MF e.v 
coll. PAL. F15A; SiciK', Palermo, Altavilla Milicia, rigdi 
side of Milicia river, 12 shs, sands, lower-middle 
Phocene, coll. MF ex coll. PAL, F2A; Palermo, Partitico, 
Trappeto, Lido Ciammarito to Nocella river mouth, 11 
shs, clavs. lower Phocene, coll. MF ex coll. PAL, F72A. 

Habitat: In the .'\tlantic Ocean, the species has a lower 
sheh-upper slope distribution (see Goias, 1999), is indi- 
cated bv the fossil Mediterranean occurrences. A shal- 
lower and more restricted distribution, hmited to shelf 
paleoenvironments, with sandy to muddy bottoms. 

Distribution: The species lives in the eastern Atlantic, 
from Senegal to northern .Angola (see also Gofas, 1999). 
It was also collected from the coasts of Mauritania (S. 
Palazzi, pers. comm., 2006). The species has a Mediter- 
ranean .Miocene to Phocene paleoditribution, being re- 
corded from the Miocene of northern Aperuiines 
(Modena). the Pliocene of northern (Piemonte, Liguria, 
Toscana, Emilia Romagna), central (Lazio) and insular 
(nortlaw'estem Sicily) Italv, south Spain (Estepona) Lan- 
dau et al. (2004, as Alvania faiiae). and Algeria (Coss- 
mann, 1921). In Atlantic, it is recorded from the Portu- 
guese middle Pliocene of Mondego Basin (Landau et al., 
2004). The citation of Wenz (1938: 616), according to 
\^'hich the species hved in the Oligocene (up to Pliocene 
of Europe, \orth Africa and North America), should be 
verified. The species was not found in the veiy rich mol- 

luscan assemblages from the Oligo-Miocene of south- 
western France ^Lozouet, 1998: 1999). 

Remarks: Gofas (1999) and Landau et al. (2004). dis- 
cussing and re-describing this species, originally de- 
scribed as Mauzonia faruii h\ Rolan and Fernandes 
( 1990) from West Africa, did not recognize its identity as 
Rissoa tiberiaua Coppi, 1876, the latter being a common 
species from the Mediterranean Neogene, particularly 
from the Pliocene. This species, originallv not illustrated 
by Coppi (1876), was figured by Sacco (1895: hgs. 67, 
a-bis and 68, a-b), who designated it as the type species 
ot the subgenus Galeodiiiopsis Sacco, 1895. More re- 
cently, this species was illustrated by Gossmann (1921: 
pi. 1, figs. 5.5-56) and Wenz (1938: fig. 1715). Compari- 
son between fossil nuiterial of R. tibenana (Figures 90— 
94, also from topot^'pe material; see also Landau et al., 
(2004; pi. 7, figs. .'3-4), to that of Rolan and Femandes 
(1990: pi. 1, figs. 4-6), and of Gofas (1999: figs. 39-42) 
strongly confirms the abo\e mentioned synonymy. The 
rather conical, ribbed shell with an inflated last whorl, 
the frequent presence of varices on the last whorl, and 
the double-rimmed outer lip are the most characteristic 
features of this species, which shows a modest variability 
in the number and strength of ribs and in the spire el- 
evation (see Figures 90-92). 

Gofas (1999) moved this species from Manzonia Bru- 
sina, 1870 [type species Manzonia crassa (Kanmacher, 
1798), see Figures 104-107] to Alvania based on the lack 
of the characteristic punctate spiral sculpture of the 
former taxon. This does not appear appropriate. In fact, 
in reasonably well-presei"ved shells, the primary spiral 
cords clearly bears a microsculpture consisting of regu- 
lar, spirally arranged pits, quite like M. crassa (compare 
Figure 99 with Figure 106). This spiral pitted micro- 
sculpture, considered a typical Manzonia character by 
Moolenbeek and Faber (1987), was indicated by 
Bouchet and Waren (1993) as not restricted to this genus 
((jccurring in Alvania, Gofasia Bouchet and Waren, 
1993, and with a rough similaritv, in Rissoininae species, 
see Gofas, 1999, figs. 79-80, 85, and 89]). The same 
authors inteipreted it as a s)anplesiomoi-phy retained in 
Manzonia and in other rissoid genera. However, the par- 
ticular structure of the secondary, very fine spiral 
tiireads, formed by roughly prismatic elements growing 
pei"pendicularly to the shell surface, is a character shared 
bv the Manzonia species, never obsei-ved in Alvania, and 
retained only in tlie recentK* described genus Gofasia 
(see Bouchet and Waren, 1993, fig. 1557). The combi- 

Figures 90-99. Qaleodinopsis tiberiana (Coppi. 1.S76). 90-93. Shells from the t\pe localitw sh(miiig\anabilit\ and \arices, middle 
Pliocene of Italy, Emilia Romagna, Modena, Maranello, Fogliano, Gagliardella, Rio Grizzaga sands, coll. MF ex coll. PAL (F39A). 
94. Profile \iew of a shell from the middle-upper Pliocene of Italv, Emilia Roniagna, Paniia, San Nicomede, Stirone River, coll. MF 
ex coU. P..\L iF14B,L 9.5—96. Protoconchs from the same locality, note variation of the abapical sculptiu"e just behind the transiction 
to teleoconch. 97. Detail ot protoconch I, showing die netted microsculpture and the partialK' immersed nucleus, same shell as 
Figure 9.5. 98. Detail of teleoconch sculpture from the shell as Figure 90. 99. Detail of teleoconch microsculpture from the shell as 
Figures 9.5-96: note the pitted pattern on the spiral cord and the stiiicture of the fine spiral ridges. Scale bars: 1 mm in Figures 90-94; 
100 p,m in Figures 9.5-96; 50 |xm in Figure 98; 20 (jim in Figures 97, 99. Black and white arrows indicate the protoconch I/protoconch 
II and protoconcli/teleoconch boundaries, respectively. 

Paee 46 

THE NAUTILUS, Vol. 122, No. 1 

y. Garilli. 200S 

Paee 47 

nation ot these characters ipittecl sculpture and structure 
of line spiriil tiu'eads) represents a t[uite sin;j;ular antl 
original feature, which should be regarded as limiteil to 
Manzonia-reVdted species.: e.g. M. clarwiiii Moolenbeek 
and Faber, 1987 (pi. 1, fig. IS), M. crispa (Watson, 1873) 
(see Moolenbeek and Faber, 19S7. pi. 3, fig. 54), M. 
boogi kinzarottii Moolenbeek and Faber, 19S7 (pi. 2, fig. 
39), M. spreta (NA'atson, 1873) (see Mook'nbeek and 

Faber, 1987, pi. 3, fig. 57 j 

M. vigoensis (Rolan, 1983) 

(see Bouchet and ^^'aren, 1993, p. 656, fig. 1499) and 
several others. The double rimnied outer lip and the 
netted sculpture ot protoconch I are also characters 
shared hv Manzonia species. However, it must be con- 
sidered that these tv\"0 features, considered h\ Ponder 
(1985: 46) as t\pical oi'Manzonia sensu stricto, should be 
regarded \\ith suspicion, being as they are shared b\' 
well-recognized Alvania species. In regard to the proto- 
conch, species such as A. testae (Aradas and Vlaggiore. 
1844), A. zetlandica (Montagu, 1815) (see Bouchet and 
Waren. 1993, figs. 1386-1387 and 1502; Landau et ai. 
2004. pi. 9. figs. lb-Id), the Pliocene Alvania magistia 
Chirli, 2006 (pi, 11, fig. 16 and pi. 12, figs. 1-3), have the 
same sculptural pattern on protoconch I. A. tomentosa 
(PaUan", 1920), which has a paucispiral protoconch, also 
e.xliibits diis sculpture (see Bouchet and Waren. 1993, 
fig. 1388). 

Rissoa tiberiana could be regarded as one (probably 
the sole) of tlie few sui"\ivors of a group of species close 
to Manzonia sensu lato, which very likely originated in 
tlie upper Paleogene. The European Oligocene Rissoa 
duboisi Nvst, 1843 (Figures 100-103) certainlv belongs 
to tliis group. Both diese species share several intersting 
characters: a Cjuite conical Alvania-Vike shell shape, often 
pro\ided with \arices on the last whorl, the arrangement 
of the basal cords (not so strong as in Manzonia sensu 
stricto, where keels occur on the shell base), the above 
mentioned combination of the microsculptin"aI pattern, 
and the kind of a.\ial sculpture (with slightK^ sinuous, less 
pronounced ribs than those shown b\' Manzonia), and a 
double, weaklv opisthochne outer lip. Ml this leads me to 
re\'alue Galeodinopsis as the useful generic placement 
for such Manzonia-Te]a.ted species. 

In o\erall appearance, G. duboisi (Nyst, 1843) strongly 
resambles G. tiberiana, from which it differs principally 
m ha\ing less numerous cords and ribs and less con\ex 
whorls (see also Ponder, 1985, fig. 100c). Another com- 
parable species is the Recent Macaronesian M. spreta 
(Watson. 1873), wliich has a similar shell shape but dif- 
fers from G. tiberiana in ha\ing a more delicate axial 
sculpture (the spiral cords being large and almost flat) a 

more rounded and smaller aperture, and a paucispiral 
protoconch (see Moolenbeek and Faber, 1987, figs. 
47a-b and 55-57). With some significant resenations, it 
could lie regarded as belonging to Galeodinopsis. The 
species M. foraniinata (Lozouet, 1998), originally de- 
scribed as Alvania (from the upper Oligocene of south- 
western France, see Lozouet, 1998, fig. 9f-h), M. nioii- 
linsi (dOrbigny, 1852) (see Lozouet, 1998, fig. 9i-j, from 
the French upper Oligocene), M. scalaris (Dubois, 1831) 
(Kowalke and Harzhauser, 2004, fig. Sd, from the middle 
Miocene, Badenian, of Austria, Hungan-, Poland and Ro- 
mania and the Miocene of Russia) and the Recent Man- 
zonia crispa (Watson, 1875) (of which I studied 3 shells 
from Madeira, ZMA Moll. 101.0, ex coll. R.G. Moolen- 
beek; see also Ponder. 1985, fig. lOOA and Moolenbeek 
and Faber, 1987, te.\t-fig. 46, and pi. 3, fig. 52-54) show 
less affinities, having a more slender shell with more 
ciu-\"ed libs (protruding over die suture in M. scalaris), 
which become stronger on the base of the shell. AU these 
lour species appear more related to Manzonia than to 

Sacco (1895) and Cossmann (1921) indicated Rissoa 
multicostata Speyer 1864 (pi. 41, figs. 3-5, from the Oli- 
gocene of Germany) as a possible Galeodinopsis species. 
I did not see anv shells of this Alvinia sensu lato-like 
species, which more closely resembles the group of Al- 
vania zetlandica (Montagu, 1815) and A. weinkaiiffi 
(Weinkauff, 1868 e.v Schwartz ms.). The original illustra- 
tions show a turreted shell with a more finelv cancellate 
(not-ribbed) sculpture, bearing almost orthocline axial 
ribs, characters which militate against placement in Ga- 

The material of Rissoa tuba Doderlein, 1862, housed 
in the MGUP Doderlein's collection (from the Miocene 
and Pliocene of North ItaK), belongs to this species. 
Anvwa)-, Doderlein (1862: 17) just listed this taxon with- 
out providing a description or a valid reference, so that R. 
ttdia must be considered a mnnen nndnm. 


As indicated bv Ponder (1985), the s\"stematic grouping 
oi Alvania species at the subgeneric level is quite diffi- 
cult and putative groups usually fold into synonymy with 
Alvania sensu stricto. This \iewpoint appears be appli- 
cable to the species studied in this i-eport (except for 
Rissoa tiberiana Coppi, 1876). The subgeneric di\ision 
into Galeodina and Massotia lacks anv \alid and convinc- 
ing basis in shell features (especialK' on consideration of 

Figures 100-107. Galeodinopsis duboisi (Xvst, lS4.3j (100-103). Irom the upper Oligocene ol Hcssen (Germany, SE oi Kasscl, 
Hessiscli-Lichten. GUnimerode. coll. MF ex coll. Pal. FE15A) and Manzonia crassa (Kanmacher, 179S) (104-107). type .species of 
Manzonia Brusina. 1870, from tlie middle-upper Pleistocene of Kvllini (NW Peloponnesus, Greece, N2 bed of Garilli et al, 2005a). 
100-102. .\pertural ( 100 and 102) and profile (101) views. 10.3-104. Microsculptures, showing the typical pitted surface on the flat 
cords and the microstructure of the narrow spiral threads in G. duboisi ( 10.3. same shell as Figure 101) and M. crassa (104, same shell 
as Figure 10.5). 105-106. .^pertural (105) and profile (106) view of shell 107. Detail of protoconch I, showing the netted sculpture. 
Scale bars: 1 nun in Figures 97-98; 20 (j.m in Figure 100: 10 |jLm in Figiu-e 99. The black arrow indicates the protoconch I/protoconch 
II boundarv. 

Pasre 48 

THE NAUTILUS, Vol. 122, No. 1 

the often extensive intraspecific variation). In elfect, a 
link between the markedly keeled and inflated shells 
(e.g. A. caiiiiatti), and the more tvpical A/rrt/n'rt-shape 
taxa could be hvpothesized. It could be articulated and 
summarized in the transition A. carinata-A. lacfea (via A. 
francescoi new species-A. rosariae new species) to more 
typical Alvania secies. A parallel trend could be con- 
stnicted: A. tenera and A. cingnlata, showing a progres- 
sive weakening of the spiral sculpture, the proportional 
reduction in the expansion of the body whorl and the 
formation of the outer lip thickening. Furthermore, 
other interspecific trends, involving various nominal 
groups oi Alvania, could be constioicted when consider- 
ing the widely variable shells of most of the species dis- 
cussed here. Nowstanding this, I cannot deny that all 
these hypothetical links appears tenuous and that these 
taxa, sharing a quite inflated and well-expanded body 
whorls, a wide, internally smooth aperture, seem some- 
what distant from Alvania sensu stricto This point of view 
could lead to regard them as belonging to an inclusive 
taxonomic group (preferablv at subgeneric level). Galco- 
dina could sei-ve as the appropriate taxonomic unit. Nev- 
ertheless, since I beheve that appropriate anatomical 
studies (of which there is no e.xliaustive data-set) should 
be used to solve this question, I have prefeiTed a more 
open-ended option, placing the discussed species here, 
from carinata to tenera, in Alvania sensu lato 

The re-analysis of the Neogene-Recent R. tibcriana, 
type species of Galeoclinop.sis, has lead me to reevaluate 
the latter tiixon as the appropriate genus lor grouping 
species characterized bv a particular teleoconch micro- 
sculpture (consisting of veiy fine spiral threads, formed 
by roughly prismatic elements, extending pei"pendicu- 
larly respect from the shell surface, plus the pitted sur- 
face on the primary spiral cords, as seen in Manzonia) 
and by having a conical A/uffMirt-like shell shape. The 
oldest representative of this genus is the European upper 
Oligocene R. chiboisi. The upper Oligocene M. foranii- 
nata and M. tnonlinsi are probabK' more linked to A/(/)(- 
zonia sensu lato than to Galeodiiinpsis while the Recent 
Macaronesian M. spreta could be doubtfullv I'egarded as 
belonging to Guh'odinopsis . 

Because of its affinity with Alvania, Galeodinopsis 
probably derived from some Oligocene group (jf that 
taxon and could be regarded as a transitional link to 
Manzonia. It is interesting that, as indicated b\' Lozouet 
(1998), the genus Alvania had a suiprising radiation in 
the upper Oligocene, creating a strong diversification. 
Furthermore, the Manzonia sensu stricto species, char- 
acterized by the quite slender shell with strongly opis- 
thocline ribs and veiy marked spiral cords on the shell 
base, seem to be well-established from the European 
Neogene, where they are represented b\ the following 
species; M. crassa, M. fahiniea, M. pontileviensis, and M. 

All the discussed species have a multispiral proto- 
conch, indicating a planktotrophic lai-val development, 
with the exception of ,\. cin'^iiJala, whicli is the most 

geographical]) restricted species, limited to Sicilian wa- 
ters. The reason lor its very limited geographical distri- 
bution is not certainlv solely attributable to its non- 
planktotrophic larval development. Our knowledge 
about this species is not encouraging; I just know that; 

• There is no knowm fossil record ... is it a veiy recent 
Mediterranean acquisition? 

• From a large amount of bulk samples (about 50 liters 
from Magnisi and 20 liters from Mondello), I picked 
only a single eroded shell! ... Is it still a living species? 

Among species with planktotrophic development dis- 
cussed in this report, three show an eastern Atlantic- 
Mechterranean distribution; A. carinata. A. lactea, and 
A. tenera. Among these, die Hrst two species have a Neo- 
cene to Recent distribution, beinsj known from the 
Pliocene and the Miocene of Mediterranean basin, re- 
spectively, and show a quite extensive east Atlantic dis- 
tribution, being commonly recorded from the English 
Channel to Morocco. Regarding the third species, living 
in Mediterranean and along the Atlantic Morocco and 
die Canaiy Islands coasts, I did not find any Mediterra- 
nean or Atlantic fossil records. The extinct species A. 
francescoi, A. pnisi. and A. rosariae are limited to the 
Mediterranean Pleistocene, while Galeodinopsis tibcri- 
ana has a Miocene-Pliocene MediteiTanean distribution 
ami lives along the W African coasts from Mauritania to 
northern Angola. 

In general, the protoconchs ot the species reported 
here do not indicate any distinguishing taxonomical char- 
acters at the supraspecific, and, in most cases, at the 
species level, being characterized by sculptural patterns 
shown by several rissoid species. Among the plank- 
totrophic species, the connnonest sculptural pattern on 
protoconch I (obsei^ved in A. carinata, A. francescoi, A. 
lactea, A. rosariae, and A. tenera), consisting of fine spiral 
lirae and scarce to abundant pimples between them, is 
sho\\ni b\- A. cancellata (Da Costa, 1778) (see Giannuzzi- 
Savelli et al., 1996, fig. 408b), A. beani (Hanley in 
Thoi"pe, 1844) (see Giannuzzi-Savelli et al., 1996, fig. 
412), A. cimex (Linneus, 1758) (see Ponder, 1985, 
figs.S6C-B), A. cimicoides (Forbes, 1844) (see Bouchet 
and Waren, 1993, fig. 1385 and Giannuzzi-Savelli et al., 
1996, fig. 408d), A. dingdensis (Janssen, 1967) (see Pon- 
der, 1985, figs. 88F-G), A. genjonia (Nardo, 1847) (see 
Giannuzzi-Savelh et al., 1996, fig. 395c), A. hispidula, 
Monterosato, 1884 (see Gofas, 1999, fig. 26), A. punctura 
(Montagu, 1803) (see Giannuzzi-Savetli et al., 1996, fig. 
436), A. stephanensis Lozouet, 1998 (fig. 9D), Cri.silla 
semistriata (Bouchet and Waren, 1993, fig. 1535), Rissoa 
parva (Da Costa, 1778) (see Ponder, 1985, figs. 79B), 
and several other species. The sculptural pattern on pro- 
toconch II (also obsened in G. tiberiana), consisting of 
1-3 more or less discontinuous, abapical to central, spiral 
threads (also formed by short, fused, prosocline seg- 
ments close to the beginning of the teleoconch) and tu- 
bercles on the remaining surface, is also showm b\' most 
of the aboNc niciitioned species and several others. Al- 

\'. Garilli, 20()S 

Paee 49 

Laniii lactca could represent a slight exception, iiaxing a 
quite chai'acteristic, easil\' distinguishalile protoconch I 
with a coarser sculpture than that shouni b\' the other 
studied species. A. tcncra shows a similar protoconch II 
sculptural pattern, with few to abvmdant \en' small gran- 
ules spir;ill\' arranged, which ma\' form \en' discontinu- 
ous and irregular ridges. Likewise this kind of sculpture 
is shared b\' other rissoids (e.g. A. tarwdvs (Watson, 
1SS6) (see Bouchet and Waren, 1993, fig. 1450) and 
Chsilla semistriata). The sculpture of the pancispiral 
protoconch of A. cingulata, which is almost a replica of 
protoconch I of the first group discussed abo\e, repre- 
sents a quite common pattern shown h\ several non- 
planktotrophic rissoids: e.g. A. argillcnsis Lozouet, 1998, 
A. macandrewi (Manzoni, 1868), Lironoba inidtilirafa 
(T. \\'oods, 1878), Onoha gianninii (Nordsieck, 1974) 
and, with a moderate similarity bv A. subsohita (Aradas, 
1847), Onoba seinicostata (Montagu, 1803), and A. Liro- 
duncnsis Lozouet, 1998 (see Ponder, 1985, fis 89A, 109E 
and 126C; Bouchet and Waren, 1993, figs. 1458, 1525; 
Lozouet, 1998, figs. lOF and lOK). 

There is similarit\' between the West African rissoul 
assemblages, including the Macaronesian province, and 
die European ones, with particular regard to those from 
tire Mediterranean Neogene. This similaritv is perhaps 
more marked than it has been indicated by Gofas (1999). 
The case of G. tiberiana is a further (see Monegatti and 
Raffi, 2001. and Garilli and Galletti, 2007) interesdng 
case of a moUuscan species that lived in the Mediterra- 
nean Neogene and today occurrs along the West African 
coasts. In this xiew, it is noteworth\- to remark that, as a 
whole, most of the species here included in GalcodUwp- 
sis li\ed in the European Oligo-Phocene while its living 
representings occurs along the West Africa and Macaro- 
nesian Pro\inces. In addition, A. tcncra, living in the 
Mediterranean, Atlantic Morocco, and in the Canaiy Is- 
lands (Tenerife), should be regarded as a new^ record, 
further suppoiting the discussed similarit)". The rissoid 
Rissoina d'Orbignv, 1840, species from the Mediterra- 
nean Plio-Pleistocene (see the good illustrations of 
Greco, 1974, figs. 11, 13, 15, 17 and Chidi, 2006, pi. 23, 
figs. 7-12), usuallv cited as R. dccussata (Montagu, 
1803), is \'er\" likeh" the same taxon as living along the W 
African coasts, Sao Tome and Cape Verde Islands [see 
Gofas. 1999; 97, figs. 69-73, and treated as R. piinc- 
fostriata (Talavera, 1975)]. 

The radrer common presence of varices on the last 
whorl (a rare character in rissoids) of A. caiinatcL A. 
francescoi new species, A. Jactea. A. rosariac new spe- 
cies, and G. tiberiana .should not be regarded as a sahent 
taxonomic character at the supraspecific level, being 
present in quite unrelated species (e.g. A. carinata and 
G. tiberiana). 


This article would not have been possible without the 
generous support of friends, private collectors, antl re- 

searchers. I particularK thank Maurizio Forli (Prato), 
Stefano Palazzi (Modena), Nino Adorna Sbrana (Gros- 
seto), Stefano Rufini (Anguillara) for lending a large 
amount of precious and indispensable material. Many 
other people kindly made their collections available. 
R.G. Moolenbeek (ZMA, Amsterdam) donated three 
shells of Manzonia crispa from ZMA Moll, collection, 
l^ossana Messina (Palermo) provided a special support, 
measiuiug most of the investigated shells and assisted me 
during the bibliographical research in the DGUP library. 
Luca Galletti (Monreale), a companion for all the field- 
works in Kvllini and Cartiera Mulino, also help me in 
providing SEM images. Mv special thanks to M. 
Glaubrecht (ZMB, Berlin), who veiy kindly lent precious 
material (with particular regard to the two Alvania cin- 
gulata shells from Vlonterosato collection), to D. Merle, 
j. M. Pacaud (MNHN-DHT, Paris), V. Heros, and P. 
Lozouet (MNHN-DSE, Paris), who provided helpful 
and kind assistance during my visits in MNHN. A. Rosso 
and I. Di Geronimo (DSTC, Catania) kindly allowed the 
\isit to tlie malacological collections of the DSTC. Kathie 
W'av (British Museum of Natural lliston', London), H. J. 
Niederhofer (Staatliches Museum fur Naturkunde, 
Stuttgart) and P. Seivend (MPOB, Modena) provided 
information on t\pe material. Thanks to M. Fiore, P. 
lacopelli, V. P. Li Vigni (MSNCS, Terrasini), who al- 
lowed the visit to the conchological collections of the 
MSNCS. C. D'Aipa (MGUP, Palermo) lent some fossil 
material housed in the MGUP. Even when the library 
was moving, G. Barranca (DGUP, Palermo) very kindly 
allowed me the access to some papers. Riccardo Gian- 
nuzzi-Savelli (Palermo) and R. C. Moolenbeek helped 
me in some bibliographic researches. R. G. Moolenbeek 
and S. Palazzi also provided verv useful and coustnictive 
comments on an early manuscript. 

I am grateful to D. Merle, R. G. Moolenbeek, H. 
Scholz (ZMB) and E. Theodorou-\'ardala (GNHM, Ki- 
fissia), who quickly provided the catalog numbers for 
type material of A. francescoi new species and A. rosariac 
new species. I am also grateful to H. G. Lee (Jackson- 
ville, Florida) and P. Lozouet (MNHN, Paris) who re- 
viewed the manuscript and provided valuable comments. 
H. G. Lee also kindly helped improve mv shaky English. 
This work also benefited from oitical readings by S. Go- 
fas (Departamento de Biologia Animal, Universidad de 
Malaga) and Jose H. Leal (the Bailev-VIatthews Shell 
.Vluseum, Sanibel, Florida). 


Aartsen, J. J. Van. 1982a. Synoptic Tables of Medit. & Europ. 

Conchology. Gen. Alvania (Subg. Ahinia & Galendina). 

La Conchiglia 14 (162-163): 8-9. 
Aartsen, J. J. Van. 1982b. Synoptic Tables of MecUt. & Europ. 

Conchology. Genus Alvania (Sbg. Arseuia & Alvaniella). 

La Conchiglia 14 (164-165): 4-6. 
Aartsen, J. J. Van, H. P. M. G. Menkhorst. and E. Gittenijerger. 

19S4. The marine Mollusca of the Bay ol Algeciras, Spain, 

v\ith general notes on Mitrclla. MarginelUdae and Tur- 

ridae. Basteria Supplement 2: 1-1.35. 

Paee 50 

THE NAUTILUS, Vol. 122, No. 1 

Anistratenko, V. V. and Ya. I. Starobogatov. 1994. MoIlu.scs of 
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16,39 pp. 

THE NAUTILUS 122(l):52-55, 2008 

Page 52 

Reallocation of Cijamiocardium crassilahrum Dell, 1964, into 
Perrierina Bernard, 1897 (Bivalvia: Cyamiidae) 

Diego G. Zelaya 

Departaniento Biodiversidad y Biologia 


Facultad de Ciencias Exactas y Naturales, 



Division Zoologia Invertehrados 

Museo de La Plata, Paseo del Bosque s/n, 




Re-examination of the holots-pe ol Cijaiiiiocarditnn crassihi- 
l)nim Dell. 1964. allowed confirmation that the .species was 
wrongly allocated to the genus Cijamiocardium. This material, 
together \A'ith additional specimens from the western Malvinas 
(Falklands) Islands allowed for a re-description of that species. 
The species is also properly illustrated through scanning elec- 
tron microscopy, and its updated generic placement discussed. 
The presence of a "taxodont" hinge plate is the main moipho- 
logical character supporting the re-allocation of this species in 
the genus Perrierina Bernard, 1897. Infonriation on the brood- 
ing condition of the species is provided. 

Additional Keywords: Perrierina. Cvamiidae. Magellanic Region 


Study of the collected bv the RA' William 
SCORESBY in the Magellan Region and adjacent Antarctic 
waters resulted in the discoveiy of several species new to 
this region, among them, a small bivalve described by 
Dell (1964) as Ciiamiocardiinn cra.s.siliibriini. The only 
subsequent record of this species was repoited bv Cas- 
tellanos (1980) fnjm tlie western Malvinas (Falkland) Is- 

The genus Cijamiocardium was introduced by Soot- 
Ryen (1951) to allcjcate Cyamium denticuJaium E. A. 
Smith, 1907 (the tj^De species by original designation), a 
species widely distributed throughout the Antarctic Re- 
gion (Laniy, J910, 1911; Powell, 1958; Dell, 1990). Other 
Antarctic species assigned to this genus by Soot-Ryen 
(1951) were Cijamiornrdium nrtiindaluin (Tliiele, 1912), 
C. dahli Soot-Ryen, 1957, and Cijamiocardium cra.s.sila- 
bnim Dell, 1964, from the Magellan Region. In the con- 
text ot a .systematic revision of these species (ciu'rentlv in 

progress), the type material and additional specimens of 
"Cijamiocardium" crassilahrum were examined. As a re- 
sult, I liave concluded in that the placement of this spe- 
cies in Cijamiocardium is incorrect. In the present paper 
the generic placement of this species is revised, and the 
taxon is re-described and properly illustrated by first 


This study is based on dried preseived specimens col- 
lected during the IW Shinkai Maru expechtion (1978- 
79). Voucher specimens have been deposited in the col- 
lections of the Museo de La Plata (MLP). Photographs of 
the holot}'pe of Ci/amiocardium crassilahrum were re- 
ceived from The Natural Histon' .Museum (NHM), Lon- 
don. Eor comparative puiposes, specimens of Cijamio- 
cardium dciiticuhiliiiiL C. rolninltilinn, and C. daldi were 
also examined. 

The specimens reported by Castellanos (1980) as Ci/a- 
miocardium crassilahrum could not be located either at 
the MLP or the Museo Argentino de Ciencias Naturales 
"Bernardino Rivadavia" (MACN). The material here 
studied, originally at the Instituto Nacional de Investiga- 
ciones y Desarrollo Pesquero (INIDEP), comes from the 
same Shinkai Maru sampling station than the material 
previously reported by Castellanos (1980). 

Shell measurements were made under a stereoscopic 
microscope, according to the following criteria: L: maxi- 
mum antero-posterior distance; H: maxiimuvi dorso- 
ventral distance, perpendicular to L; W: maximuTU dis- 
tance across valves, peipendicuhu" to H. Shell niorpliol- 
ogy was stu(hed under scanning electron microscope; for 
this, Philips XL3() TMP and JEOL JSN-6360 LV scan- 
ning electron mio'oscopes were used. 

D. Zelava, 200S 

Page 53 


C\ainii(lat' G. O. Sars, 1S7S 
Perricriita Bernard, 1897 

Perricrina crassilahnini (Dell. 19fi4) new combination 

I, Figures 1-12) 

Ci/ainiocardium ciYissildhniiii ^^•ll. U)(i4: 204, Cig. 4, 

nuniber.s 1, 2, and pi. 6, figs. 1, 2; Castellano.s, I9S(): 135. 

T^-pe Locality: 5()°17' S, 60°06' W, station 211. \\A' 
W'lLLI.VM S(.:OKKSBY. lttl-174 m 

Material Examined: Photographs of the Iiolot\pe 
(NHM 1962S63); 7 dried specimens and 1 valve, 51°'29' 
S. 61" 50' ^^■. Nhihinas (Falkland) Islands, 192 m (MLP 

Literature Records: 50°17' S, 60°06' W, station 210, 
RA' WiLLi.\M SCORE.SBY, 161 m; 50°35' S, 57°20' W, 
station 229, RA' Scoresby, 210-271 m (Dell, 
1964); 51°29' S, 6r50' \V, 192 m (Castellanos, 1980). 

Distribution: OnK kno\\n Ironi the Atlantic sector of 
the Magellan Region, in tlie vicinitv' ot the Malvinas 
iFalkland^ Islands. 161-271 ni. 

Description: Shell solid, small (maximum L: 6.3 mm), 
outline nearl\- circular (H/L = 0.99 ± 0.01, n = 6), slightK' 
inequilateral', not inflated (W/H = 0.65 ± 0.01, n = 5), 
whitish. gloss\'. .Anterior margin short, eun-ed, continu- 
ous witli \entral margin, \\'hich is evenly arcuate; poste- 
rior end rounded, shghtlv expanded (Figures 1, 2). Juve- 
nile shell slightly ovate in outline, elongated antero- 
posteriorlv (Figure 3). Beaks full, subcentral, directed 
sUghtK' anteriorlw Protoconch ovate, smooth, about 590 
jjLui in diameter (Figure 5). Shell surface sculptured with 
rounded, strongly marked, and regularly spaced radial 
cords. 50-60 in larger specimens (Figure 6); cords sepa- 
rated b\" interspaces wider than ribs. RegularK" sepa- 
rated, microscopic commarginal threads also present. 
Radial scidpture also e\ident on inner margins, where 
thev produce prominent crenulations (Figures 7-9). 
Hinge plate narrow, somewhat broader anterior to beaks, 
where the carchnal teeth are inserted. Right valve with 
large, hooked cardinal 3: 3a liigh, solid, triangular, en- 
larged at base, which is frequently bifid; 3b delicate, 
narrow, nearlv straight, one-third of size of 3a (Figures 7, 
10). Left \al\-e \\-ith prominent triangular cardinal 2, thin 
and st\liform cardinal 4b behind resilifer, and slender 
but solid, nearK" straight anterior tooth (referred to as 
"cardinal 4a" bv Bernard (1897) and as "anterior lateral 
11" by Lamy (1917)) (Figures 8. 9, 11). Both valves with 
two series of tubercles anterior and posterior to beaks, 
producing the appearance of a taxodont hinge plate (Fig- 
ures 10. 11); each series composed oi four tubercles; 
posterior series stronger than anterior in juveniles (Fig- 
ure 12). In both series, tubercles diminish in size and 
degree of de\elopment from beaks to anterior and pos- 
terior ends, where thev are followed by marginal crenu- 
lations. Internal hgament somewhat solid, located in 

shdil, ()lili(|U(' rcsiliter posterior to cardinal teeth. E.xter- 
ual ligament short. 

Biological Obsei'Nations: One ot the specimens e.x- 
aniinetl contained numerous embryos between the as- 
cending and descending lamellae of inner demibranchs. 
Embryos were in different stages of development, rang- 
ing from incipient (unshelled) to well-developed .speci- 
mens, the latter mnnbering 32, and reaching 700 |xm in 
diameter (Figure 4). These obsen'ations are consistent 
with that by Dell (1964), who reported 44 "developing 
voung" contained in one of the specimens he studied. 

Remarks: Pcrricrino was proposed bv Bernard (1897) 
loi" P. Iiixiuloiitii (t\pe species by monotxpv; illustrated by 
Bernard, 1897; tig. 3) a species he described from lie 
Stewart (New Zealand). The description oi the genus was 
merged with that of the type species, in which Bernard 
(1897) described the presence of several "lamellae" an- 
terior and posterior to the cardinal teeth, resembling a 
taxodont hinge. This character, infrequent among Cya- 
mioidea, is also present in Legrandina Tate and May, 
1901 (ty[3e species: L. bemardi Tate and May, 1901, by 
original designation), an Austrahan genus regarded by 
Ponder (1971) and Powell (1979) as a subgenus oi Per- 

Perrieiina and Cijamiocardiiim have as conunon char- 
acters the number, moqiholog\', and arrangement of 
hinge teeth, i.e., the presence of a hooked anterior car- 
dinal 3 in the right vah'e and two prominent cardinal 
teeth (cardinals 2 and 4) in the left valve. The specimens 
studied here generally agree \\ith these eharaeteristies, 
but, additionallv, the hinge e.xamined showed several an- 
terior and posterior tubercles, which, as a group produce 
the appearance of a taxodont hinge, a diagnostic charac- 
ter for Perrierina that is absent in Cyamiocardiiim. Fur- 
thermore, the studied specimens lack the posterior 
"pseudo-lateral tooth" described by Smith (1907) for the 
i)1je species of Cyamiocardiiim. Anotlier character that 
is useful in separating the genera Cyamiocardiiim and 
Perrierina is the presence in the former of mantle margin 
papillae (Soot-Ryen, 1951). Accorchng to Ponder (1971), 
mantle margin papillae are absent in Perrierina. Unfor- 
tunately, the poor preseivation of the specimens on 
which this paper is based made it impossible to confirm 
the state of this character. Nevertheless, the differences 
in hinge moi"phology call for the reallocation of "Cija- 
miocardiiim" era.ssico.itaUim into Perrierina. 

Perrierina craasilahnnn shows some characteristics in- 
termediate bet\veen the subgenera Perrierina and Leg- 
randina. The presence of prominent beaks is a charac- 
teristic shared with Perrierina, whereas a hinge plate 
broader at the base of the cardinals is characteristic of 
Legrandina. Furthermore, there are some other features 
of P. era.s.silahnnn not prexiously knowii for other species 
of Perrierina or Legrandina, such as the relatively lai'ge 
adult size, the nearly circular shell outline of adults, and 
the strong radial ornamentation. 

Page 54 

THE NAUTILUS, Vol. 122, No. 1 

Figures 1-12. Ferrierina crnfssilaJinim. 1, 9. Holotype (NIIM 1962863). 1. E,xternal vit-w n\ Ic II \,i1m . !). Iiilfnial view of left valve. 
2-8, 10-12. Specimens from 51" 29' S, 61° 50' W, SEM micrographs. 2. External view of an adult specimen, right vaKe. 3. E.xternal 
view ot a juvenile. 4. External views of lan'ae removed from an adult. 5. Dorsal view, showing protocoiich and hinge plate. 6. Detail 
of shell microsculpture. 7. Inner view of right valve. 8. Internal view of left valve. 10, 11. Detail of adult hinge platt's. 10. Right \aK(>: 
3a, 3b = cardinal teeth, L = ligament. 11. Left valve. 2, 4b = cardinal teeth, at = anterior tooth. 12. Detail ol hinge plate ol a jnxcnile 
of 2.3 mm length. Scale bars: 1-3, 7-9 = 1 nun; 4 = 100 fxm; 5 = 200 n,m; 6 = 50 |xm; 10-12 = 500 |xiii. 

D. Zela\'a, 2008 

Page 55 

The taniilial plat'einent of Prii'ici'iitd is soiiicwliat con- 
tusing: the genns was suceessiveK' placed within the 
Mactridae tBernard, 1897), Leptoni'dae (D;ill, 1899), and 
CrassateUidae (Suter, 1913; Lamy, 1917). Later, Mar- 
wick (1928) regarded the "taxodont" lameUae of the 
hinge as a chai"acter sufficient to warrant proposition of 
the famil\- Perrierinidae, but this faniiK' was subse- 
quentK' regarded a,s a s\iion\ni of Cvaniiidae (e.g. Cha- 
van (1969), Thiele, (1934)', Fleming (1948), Ponder 
(1971), and Powell (1979)). The Cyamiidae is a family 
relativeK' well-diversified in the Magellanic Region, 
where a total of 12 species belonging to Ci/ainimn, Ci/a- 
iniocariliinn, Ga'unardia, and Kiddcnu have been re- 
ported and recognized as \alicl by Zelaya (2005). The 
present paper represents the first record in Magellanic 
waters of a species o{ Pcnicrina, a genus thus far known 
onK' from Australia, Tasmania, and New Zealand. 


The author is grateful to A. Roux (INIDEP) who kindly 
made available the material collected during the RA^ 
Shixkai M.\ru cruise: K. Wav and A. MacLellan (NHM) 
IdndK' sent photographs of the holotspe of Ci/amiocar- 
diitin crassilabmm; C. Digiani (MLP) helped with older 
literature: and C. Ituarte provided valuable criticism and 
suggestions on an earh^ \^ersion of the manuscript. 

The author is member of the National Research Coun- 
cil for Science and Technolog)' (CONICET), Argentina. 
This stud\' was partially supported bv a grant from the 
Western Societ\" of Malacologists, made possible by the 
Santa Barbara Malacological Societv, the Southwestern 
Malacological Societ)-, the San Diego Shell Club, and the 
Nortliem California Malacological Club, and by grant 
PICT 282 from "Agencia Nacional de Promocion Cienti- 
fica V Tecnologica," Argentina. 


Bernard, F. 1897. Sur quckjues coq\iilles des LaiiRliibraiiches 

de 1' lie Stewart. Bulletin du Museum d" Histoire Na- 

turelle 7: 309-314. 
Castellanos, Z. A. de. 1980. Micromoluscos pdco conocidos del 

sur argentino-chileno. Xeotropica 25 (74): 133-140. 
Chavan, A. 1969. Superfamilv Cyamiacea. Pp. 537-543. In: 

Leslie Reginald Cox et al.. Part. N [Bivalvia], part. 6, vols. 

1 and 2: xxx^ii + 952 pp. Treatise on invertebrate paleon- 

tologw Lawrence, Kansas. Geological Societx' of America 

& Uni\'ersitv of Kansas. 
Coan. E. ^^. P. \. Scott y F. R. Bernard. 2000, Bivalve seashells 

of Western North America. Santa Barbara Museum of 

Natural Historv. Monographs number 2, Studies in Biodi- 

\ersit\' n°2, 764 pp. 

D;ill, W. H. 1899. Synopsis ol the Recent and Tertiaiy Lep- 
tonacea of North America and the West Indies. Proceed- 
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pis. 87-S8. 

Dell, R. K. 1964. Antarctic and sub-Antarctic MolKisca: Am- 
phineura. Scaphopii(l:i :in(l Hi\:iKi,i. Disco\en' Reports 33: 
93-250, pis. 2-7. 

Dell. R. K. 1990. Antarctic Mollusca with special reference to 
the fauna of the Ross Sea. Bulletin of the Royal Societ)' of 
New Zealand 27: 1-311. 

Egorova, E. N. 1982. Biological results of the Soviet Antarctic 
E.xpeditions, 7. Explorations (jf the fauna of the seas 26 
(34): 1-143. 

Fleming, C. A. 1948. New .species and genera of marine Mol- 
lusca from the Southland Fiords. Transactions ol the Ro\al 
Society of New Ze;il;md 77 (1): 72-92. 

Lamv, E. 1910. Mission dans I'Antarctique dirigee par M.Le Dr 
Charcot (1908-1910), collections recueillies par M. le Dr. 
Jacques Liouville. Bulletin du Museum National 
d'Histoire Naturelle 7: 388-394, 

Lamy, E. 1911. Gasteropodes pro.sobranclies, scaphopodes et 
pelecypodes. Deuxieme Expedition Antarctique Fran^aise 
(1908-1910) eonnnmandee par le Dr. J. Charcot. Sciences 
Naturelles: Documents Scientifiques. 32 pp., pi. 1. 

Lamy, E, 1917. Re\ision des CrassateUidae \i\'ants du Museum 
dHistoire Naturelle de Paris, lonnuil tic Gonchyliologie 62 
(4): 197-270 + pi. 6. 

Mai-wick, J. 1928. The Tertiar)- Mollusca, Catham Islands. 
Transactions and Proceedings of the New Zealand Insti- 
tute 58 (4): 432-506. 

Fonder, W. F, I97I. Some New Zealand and Subantarctic bi- 
\al\es of the Cvamiacea and Leptonacea with descriptions 
of new taxa. Records of the Dominion Museum 7 (13): 

Powell, A. W. B. 1951. Antarctic and Subantarctic Mollusca: 
Pelecypoda and Gastropoda collected by the sliips of the 
Discoveiy Committee during the years 1926-1937. Dis- 
coveiy Reports 26; 49-196, pis. 5-10. 

Powell, A. W. B. 1958. Mollusca from the Victoria-Ross quad- 
rants of Antarctica. B.A.N.Z.A.R. Expedition B. 6: 16.5- 

Powell, A. W, B. 1979. New Zealand Nkjllusca. Marine, land 
and freshwatei' shells. Auckland, William Collins Publish- 
ers, 500 pp. 

Smith, E. A. 1907. Mollusca. 5. Lamcllibraucliiata. NatiouiU 
Antarctic Expedition 1901-1904. Natural Histoiy 2, Zool- 
og)' (Vertebrata, Mollusca, Crustacea): 1-6. 

Soot-R\'en, T. 1951. Antarctic Pelecxpods. Scientific Results of 
the Nonvegian Antarctic Expeditions I927-I928, 32: 1 — i6 
+ I pi. 

Suter, II, 1913. Manual of the New Zealand Mollusca, |()lm 
MacKay, Government Printer, Wellington. 1120 pp. 

Thiele, J. 1934. Handbuch der svstematischen Weichtierkunde. 
Gustav Fischer Verlag, Jena. Bd 3, Dritter Teil, pp. 1193- 

Zelaya, D. G. 2005. The bivalves from the Scotia Arc islands: 
.species richness and faunistic affinities. Scientia Marina 69 
(,suppl. 2): 11,3-122. 

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Volume 122, Number 2 
June 23, 2008 
ISSN 0028-1344 

A cjuai-tedii devoted 
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Vohimc 122. Number 2 

June 23. 2008 

ISSN 0028-1344 

Luiz Ricardo L. Sinione 
Carlo M. Cunha 

Jonathan Hendricks 
Roger W. Portell 

Janine O. AiTuda 
Jose ^^'. Thome 

Roland Hoiiart 

Guido Pastorino 
Fabrizio Scarabino 

Rexision of the genus Spinosipella (BivaKia: \''erticordiidae). with 

dfscriptions of two new species from Brazil 57 

Late Eocene Conits (Neogastropoda: Conidae) ironi Florida, USA 79 

S\nonvinization ot Ncolii/dliiiKix Siniroth, 1S96, and Oiiuilonijx d'Orhigny, 
1837, witli a redescription of Oiiuilniii/x lirasilicnsia (Siniroth, 1S96) 
(Gastropoda: Succineidae) 94 

Rehabilitation of Ergalatax inai-tcnsi (Schepnian, 1892) (Gastropoda: 
Minicidae), senior SMionvm oi Ergalafax obscura Houart, 1996, and description 
of Eriifilatax jtiukmac, new name for Moriihi indiiensi Dall, 1923 99 

Two ne\\' deep-sea niuricids (Gastropoda) from Argentina 107 

MBLWHOI Library 

JUN 2 Ij ?on8 

Massachusetts 02543 

THE NAUTILUS 122i2):57-78, 2008 

Page 57 

Re\ision of the genus SpinosipcUci (Bivalvia: Verticorcliidae), 
\\ith descriptions of two new species from Brazil 

Luiz Ricardo L. Sinione 
Carlo M. Ciinha^ 

Miiseu tie Zoologia da Universidade de 

Sao Paulo 

Ciiixa PostiJ 42494 

04299-970 Sao Paulo. BRAZIL 



.■\ re\ision of die deep-water \erticordiid genus Spinosi))cUa is 
proxided. based on conchological and anatomical characters. 
The genus is considered distinct from Veiiiconlia {of w'hich it 
was considered a subgenus) based on the strong ribs, prickly 
surface, reduction of lunula, relative large size, weakly spiral 
\al\e shape, and odier characters. The following species are 
considered in die genus: (1) Spinosipella agnes new species, 
ranging from Florida, USA, to Rio de Janeiro, Brazil, and also 
including die Porcupine Abvssal Phiin in the North Atlantic; (2) 
S. tingti new species, occurring troni Rio de Janeiro to Rio 
Grande do Sul, Brazil; (3) S. acuticostata (Philippi, 1844). a 
Phocene fossil from southern Itah'; (4) S, deshaijesiana (Fis- 
cher. 1S62). from soudi and central Indo-Pacific (S. ericia Hed- 
le\". 1911, die t)pe species of die genus, was revealed to be a 
new s\Tion\nn of S. deshmjesiana); and (5) S. costeminens 
(Poutiers, 1981), from die tropical west Pacific. The five species 
differ mainK" in conchological details of the number and size of 
ribs, of die prickl\' sculpture, shape of die shell, of the hinge 
and die degree of com-e.xitw Anatomical description is also 
pro\ided for die two Pacific species, which differ among them- 
selves mainl\- b\" die size of the pair of renal folds. From the 
standpoint of anatomical characters, the more significant are; 
die wide Hdiodesma; die elongation of the auricles, crossing the 
roof of pallial ca\-it\': a taD digital fold in posterior region of 
supraseptal chamber; die low but wide palps; the muscular, 
gizzard-Uke stomach; die complete separation of both constitu- 
ents of the heniiaphrocUtic gonad (a ventro-posterior testicle 
and a centro-dorsal o\arv), and a complete fusion of the visceral 

Additional Kei/uords. MoUusca, .\nomalodesmata. Septibran- 


The \'erticordudae is a famiK' of septibranch bi\'alves 
comprised of carnivorous and inostK^ deep-water species. 
The\' are tvpicaUy small (less than 10 mm) but some 
species reach 30—40 mm. The\- are mostI\- radialK' sculp- 
tured and usuallv have nacreous inner surface. 

The genus Spinosipella Iredale, 1930 (tv-^je Veriicordia 
ericia Hedley, 1911, bv original designation) is usually 
considered a subgenus of Verticordia Sowerby, 1844 
(e.g., Thiele, 1934; Moore, 1969; Abbott and 'Dance, 
1983). The genus encompasses species with shell having 
pricklv outer surface, lunule veiy reduced, thick walls, 
and generally larger size (up to 30 mm). In addition to 
the type species, S. ericia, three other species are cur- 
rently included in this genus, S. acuticostata (Pliilippi, 
1844), from Atlantic and Mediterranean (middle Tertiaiy 
to Recent); S. deshaijesiana (P. Fischer, 1862a) and S. 
costeminens (Poutiers, 1981), from Indo-Pacilic. Some 
authors hax'e considered S. deshaijesiana as an Indo- 
Pacific occurrence of S. acuticostata (e.g., Nobre, 1936; 
Crozier, 1966; Rosenberg, 2005). 

E.xamination oi worldwide samples, with an emphasis 
on the Western Atlantic, showed that two species actually 
e.xist in the Atlantic. Both are separate from the fossil S. 
acuticostata. In addition, it was possible to reorganize the 
Indo-Pacific species, mainly because oi the abundant 
material deposited at the .Museum national d'Histoire 
naturelle, Paris (MNHN), which results from several e.x- 
peditions. A revision of the taxonomy and a necessaiy 
re-definition of taxa are provided in this paper, as part of 
a larger project revising Western Atlantic mollusk tax- 
onomv, based on moipliolog)'. 


A detailed list of the material examined follows each 
species description. Specimens generallv belong to mu- 
seum collections. Most material consists of shells e.xam- 
ined under a stereomicroscope. Some few Pacific 
samples have presei^ved soft parts in 70% ETOH. They 
were dissected b\' standard techniques, under stereo- 
microscope, with specimen immerse in alcohol. All dis- 
secting steps were digitally photographed; all drawings 
were made with the aid of a camera lucida. In the case of 
the material examined of Spinosipella deshaijesiana and 

Pacre 58 

THE NAUTILUS, Vol. 122, No. 2 

S. costeniinens, as the quantity of examined lots is veiy 
large, mainly thorugh courtesy of staff at MNHN (Paris), 
the list only contains the comitn' and the quantitv' of 
specimens. The full list oi examined lots oi these speci- 
mens is being published elsewhere, in a complementan" 
paper (Sinione and Cunha, in press). 

Abbreviations used in figures are: am, anterior adduc- 
tor muscle: an, anus; au, auricle: bs, byssus: by, byssal 
gland or fiu^row: cc, cerebral commissure; ce, cerebral 
ganglion; cj, connective tissue; cm, circular muscle layer; 
CO, cerebro-\isceral connective; cr, crustacean inside 
stomach; cv, ctenidial (efferent) vein; dd, ducts to diges- 
tive ch\'erticulae: dg, digesti\^e di\'erticula: es, esophagus; 
fa, foot aperture ot mantle; fm, posterior toot retractor 
muscle; fr, anterior foot i-etractor muscle; ft, foot; ga, 
genital aperture; ge, gastric epithelium; gi, gill: he, 
hemocoel; ic, intra-septal chamber; in, intestine: ki, kid- 
ney; Im, lateral muscle; lo, longitudinal muscle layer; It, 
lithodesma; nib, mantle border; mf, fiised mantle edge; 
nig, radial mantle gland; nio, mouth; nip, mantle ten- 
tacle; ms, mantle muscles of iucurrent siphon; nit, 
mantle; mu, muscular tissue; iie, uephropore; nv, nene; 
oy, ovan'; pa, posterior adductt)r mirscle; pc, pericar- 
divmi; pg, pedal ganglia: pi, papilla oi e.xcurrent chamber 
roof; pni, pallial muscles: pp, palp; rs, renal fold; rt, 
rectimi; se, e.xcurrent siphon; sh, shell; si, iucurrent si- 
phon; sm, septum muscle; sp, septum; ss, style sac; st, 
stomach; su, supra-septal chamber; sy, ciystalline style; 
ts, testis; um, shell umbo; ve, \entricle; vg, \isceral gan- 
glia; vin, \isceral mass. 

Abbreviations of institutions: AMS, Australian Mu- 
seum at Sydney, Australia; EGC, Emilio Garcia collec- 
tion; FMNH, Florida Museum of Natural History, 
Florida, USA: HGLC, Harrv G. Lee collection; 
INVEMAR-MHNMC, project of Museo de Historia 
Natural Marina de Colombia; MHNMC, Museo de His- 
toria Natural Marina de Colombia (Programa de Bi(5di- 
versidad y Ecosistemas Marinos); MNHN, Museum na- 
tional d'Histoire naturelle, Paris, France; MZSP, Museu 
de Zoologia da Universidade de Sao Paulo, Brazil; 
RLPC, Rafael La Perna collection (Universita di Bali, 

Specimens irom other verticordiid species were also 
examined for comparative pui"poses. This material in- 

Hiiliris fischeiiaiiii Dall, 18S1: 

UNITED STATES OF AMERICA. Florida, Fowey Light, 130 
m depth, MZSP 199.34, 2 valves (R.V. EoLis sta. 184). 
BRAZIL. Rio (le Janeiro; 22°.34' S, 40°29' W, 213 m 
depth, MZSP 1S751, 1 valve (on Laminarias, VV. Besnard 
col, est. IX). Rio Grande do Sul; 30°42' S, 49°03' W, 
182-186 m depth, MZSP 187,50. 5 valves (21 Aug. 1972); 
.32°55' S, 50°.34' W, 99 m depth, UFRG 1688, 1 shell and 
10 valves (sta. 4.5, 68.39 dredge, 04/iv/1998). 

Euciroa sp.: 

BRAZIL. Hio Cniiide do Sul, off Tramandai, .30°42' S, 49°03' 
VV, 182-186 ni, MZSP 18750, 5 valves (RA' W. Besnard, 

GEDIP sta. 1856, 21 Ang. 1972), MADAGASCAR. 600 m 
depth, EGC 23588. 1 shell. MOZAMBIQUE, off Moron- 
dava, Channel Madagascar, 600-800 m depth, MZSP 
61816, 3 shells (Trawled by local fisherman. May 2002). 

Euciroa clcis,aiitissinia (Dall, 1881). 

84°58.99' W, 457 m depth, EGC 13005, 6 valves (dredged, 
RA' Pelican), 24°09' N, 82°31' W, about 64.3 km off 
Southwest of Key West, 549 m depth, EGC 23688, 1 shell 
(IW Oregon II col, cruise #45, sta. 13.362): Monroe Co. 
Straits of Florida, 549 m depth, FMNH 209892, 1 shell 
(Frank Lyman col), 24°15.1' N, 82°11.71' VV, 525 m 
depth, FMNH 164794, 1 valve (G.H. Burgess, et. al. 
GHB-90-8, 23 Apr. 1990). COLOMBIA. Santa Marta, 
Cen-o de Punta Betin, A.A. 1016, (MHNMC INVEMAR), 
MHNMC 2782, 3 valves, MHNMC 2781, 1 valve. 


Gemis Sj)iii(}.\ipill/i Iredale, 19.3() 

lphi>^enia Costa, 1850: 398 (type species by original designation 
Hippci'^us aciiticostatus Philippi. 1844) (pre-occupied) 
(non Schumacher, 1817). 

SpiuDsipella Iredale, 1930: 388 (t\pe species by original desig- 
nation Vciiicordia ericia Hedley. 1911); Poutier and Ber- 
nard, 1995: 142. 

Verticordia (SpinosipcUa): Thiele, 1934: 1428; Moore, 1969: 

Diagnosi.s: Shell relati\el\' large, obese, with spiral 
\alves. Surface prickly including on radial I'ibs. Radial 
ribs tall, extending beyond shell margin. Liunda veiy re- 

Description: SniiLL: From small to medium size (up 
to .30 mm). Widtli/length ratio usually about 1. Color 
opaque-whitish. Sculptured by strong and tall radial, 
weakly cuived ribs, triangular in section, bulging weakly 
be\'ond shell edge, alternating in both \alves. Surface 
spuiy, constituted by uniform sized, veiy small bulbs, 
covering almost entire outer surface. Lunula veiy re- 
duced. Umbo projected, weakly spiral. Right valve witli 
single tall, pointed and broad cardinal tooth. Left valve 
with low, liroad tooth (posterior to tooth of right valve), 
and plane cardijial eoncavit)' as socket of tooth of right 
valve. Ligament just anterior to anterior hinge tooth, in- 
serted at some distance from median hne, in appro.xi- 
matelv middle way bet\veen hinge mechal edge and lun- 
bonal caviU'- (Figures 10, 12). Lithodesma wide, cune, 
occupying about 0.25 of hinge length, possessing a pah" of 
lateral ligamental articulations (Figvu-es 93, 96-105). 

List of Included Taxa: S. acuticostata (Philippi, 
1844); S. (ignes new species; S. costeniinens (Poutiers, 
1981),- S. cicshni/csiann (P. Fischer, 1862a) [=S. ericia 
(Henley, 1911)]; ,S. fingn new species. 

Spinosipella agues new species 
(Figures 1-18, 27-29, .31, .55) 

Ycrlirfrihii (initicoslata. ^\ohre. 1^)36: :l(),3-.304; 1938: 769- 
770; Abbott, 1974: .563: Abholl ;in(! Dance, 1983: 375 

L. R. L. Simone and C. M. Cunha, 2008 

Paee 59 

Figures 1-14. SpinosipcUa agnes new species. Shells. 1-10. Holotvpe (length 23.2 mm) shell. 1. Left valve, outer view. 2. Right 
\al\e. .3. Right \'al\e, inner view; 4. Left valve, inner view. 5. Dorsal view. 6. Posterior view. 7. Anterior view. 8. Detail of shell surface 
in SEM, middle region of right valve. 9. Detail of inter-umbonal region, dorsal view. 10. Hinge, ventral-inner view. 11-14. Paratv'jies. 
11. HGLC. from Florida, left valve, outer view; 11 mm. 12. Same, venti'al view, valves opened for showing whole view of hinge. 
1.3-14. ECtC 17419- from Colombia, outer view of right and left valves; 18 rrmi. 

Page 60 

THE NAUTILUS, Vol. 122, No. 2 

Figures 15-29. Sjiiuosi^jella new .species. Sliills. 15-18. S, agues parat^-pe AMNH 16280.3, Florida, .specimen vwth remains of soft 
parts; 14.8 mm. 19-26. S. tiiiga t^pe specimens. 19-21. Holotxpe, left valve, outer, inner and dorsal views; 16,9 mm. 22-24. 
Paratype MORG 18085, right valve, dorsal, outer and inner views; 10.1 mm. 25. Holot)'pe, dettiil oi hinge, left valve. 26. Parat)pe 
MORG 18085. right valve, detail of hinge. 27-29. S. digiics. 27. Holohpie, left valve, detail of hinge; arrow indicating well-developed 
posterior cardinal tooth. 28-29. Paratype INV-MOL 294.3. froiri Colombia, a specimen of equivalent size of main types of S. tinga 
lor showing major differences (compare with Figures 19-20); inner and outer views; 17.0 nun: note de\eloped posterior cardinal loot 
o( hinge (arrow), fewer, tallei' and more .spaced ribs and more projected ribs at edges. 

L. R. L. Simone and C. M. Cunha, 2008 

Page 61 

(fig.); van Aartsen, 1992: 45; Poppe and Goto, 1993; 139; 
PMcLean and Geiger, 199S: 27, 109 (fig.); Salas, 1996: 46; 
Rosenberg, 2005 (part) (non Philippi, 1844). 
Veriicoirlia deshaijcsiana. — Rosenberg. 2005 (part) (in s\ii- 
onvnu') (non Fi.scher, lS62a). 

T'vpe Material: I lOLOTYPE, MZSP 36917; BRAZIL. 
Rio de Janeiro, off Cabo Frio, 23°41' S, 41°03' ^^', 750- 
800 m 'depth (o.t.). PAR.ATYPES. UNITED STATES 
OF AMERICA. Florida: Off' Cape Canaveral, 903 m 
depth. USNM 64039, 1 ri2;ht val\e; SE of Sand Key, 
AMNH 24S45S. 4 shells (Jan. 1970, J. M. Bijur Collec- 
tion). AMNH 248459, 2 shells (}un. 1970, J. M. Bijur 
Collection); Monroe Countv, S.E. Sand Kev, 270 in 
depth, HGLC, 2 shells, FMNH 154594, 1 'specimen 
(dredged, Jern Phelps col., Jun. 1970); 120.6 km east of 
DaNtona. 29°i7' N, 79°27'" ^^^ 878 m depth, USNM 
810590, 1 shell and 1 left valve (RA'. Oregon, sta. 6690, 

9 Max- 1967); Marquesas Key, 24°I5' N, 82°13' W, 278- 
419 m depth. 1 left, USNM' 810889, 1 right \al\-e (RA'. 
Blake, A. Agassiz 1877-1878). COLOMBIA, off Carta- 
gena, 10°28'' N, 75°42' W, 280 m, MHNMC 2203, 1 
valve (E-47), 10°31' N. 75°37' \\\ 309 m. MHNMC 
2775, 1 \alve (E-141), Palomino, Dibulla, ir29' N, 
73°27' W, 476 m, MHNMC 3104, 4 valves, (E-21), Gua- 
jira, Bahi'a Honda, 12°31' N, 72°8' W, 452 m, MHNMC 
2943. 1 shell (E-12), Guajira Peninsula, 12°30' N, 72°08' 
\^', 470 m depth, EGG 17419, 1 shell; Cabo de la Vela, 
12°19' N, 72°42' \\\ 464 m, MHNMC 3087, 2 valves 
(E-19), Islas del Rosario, 10°10' N, 76°01' W, 510 m, 
MHNMC 2208, 2 vaKes (E-78). BRAZIL. Rio Grande 
do Norte; 206 m depth, MZSP 84627, 1 shell (Sta. D-22, 

10 Nov. 2001); Pernambuco; 690 m depth, MZSP 84628, 
1 shell iSta. D-11). 

Diagnosis: Shell with 15-17 radial ribs; smooth pre- 
umbonaJ region wide (about 0.25 of shell length); prickh' 
sculpture chaoticalK' organized. Widtli/length ratio in 
each \alve approximately 0.57. Posterior carchnal tooth ot 
left val\'e liinge well developed; main cardinal tooth of 
left \-alve relati\"el\- low and cylindrical; main cardinal 
tooth of right \al\'e tall (about 0.2 of vaKe width) and 

Description: Shell; Up to 22 mm, equivalve, inflated, 
each \;d\"e s\'mmetricallv and \veaklv spiral (1 whorl) 
(Figures 7. 9). Color white. Degree of convexit)- (widtli/ 
lengtli) in each \al\e appro.ximately 0.57. Outer surface 
spiny, opaque forming an irregular mosaic (Figures 8, 9). 
Umbones located in middle region of dorsal surface, spi- 
ral, liigh. di\ergent. separated from each other at about 
Vi of shell width (Figures 1—4, 1.3-18, 55). Sculptured bv 
strong, uniform, arched, radial ribs, from 15 to 17 in each 
\"al\e. Posterior edge about twice as wide as anterior 
edge. Between umbo and anterior edge a conca\itv bear- 
ing trans\ersal ribs, slightly wider than ribs of remaining 
region (Figures 5, 7, 9). Pre-umbonal region smooth, 
narrow, 0.2 of shell length (Fisures 5, 7, 9). Anterior, 
ventral and posterior edges forming zigzag (Figures 3, 4); 
tips of tliis zigzag coinciding with tips of each rib, tips 
encasing in concavit\- of opposite valve (Figures 1, 2, 11, 

1.3-16), Inner surface iridescent, whitish, glossv; includ- 
ing hinge (Figures 3, 4, 10, 12, 17, 18). Hinge with a large 
cardinal tooth in right valve, stubby, tall (about 20% of 
\aK'e width), broadly pointed, weakly ciin'ed fonvards 
(Figures 3, 10, 12, 18), circular in section; correspondent 
socket in left valve shallow, restrict to dorsal surface; tliis 
socket flanked by small tooth in each side, anterior 
smaller and lower than posterior (Figures 4, 10, 12, 17, 
27-28 [arrow]). Ligament just anterior to anterior hinge 
tooth, inserted at some distance from median line (Fig- 
ures 10, 12), approximately midway between hinge me- 
dial edge and umbonal cavitv'. Scars of adductor muscles 
shallow (Figures 3, 4, 17, 18, 28); anterior scar elliptical 
(longer dorso-ventraUy), located close to anterior edge, 
area about Vrs of inner surface of valve; posterior scar 
circular, about 0.33 larger than anterior scar, located 
close to posterior shell edge. Palhal line continuous, lo- 
cated at wide distance from shell edge, about 0.33 of 
distance betsveen ventral and umbonal height. 

Measurements (Length, height, width, in mm): 

Holot\.pe: 20.1 by 23.2 by 22.2; EGC 17419: 15.6 by 15.5 
by 15^5. 

Geographic Distribution: Florida, USA, to Rio de 
Janeiro, Brazil. 

Habitat: Muddv bottoms, 270-900 m. 

Material E.xamined: T\|ies. BARBADOS. USNM 
6.3200, 3 vaKes (Blake Coll., sta. 100). CUBA. Havana; 
Gulf of Mexico, 419 m depth, USNM 63201, 3 right, 4 
left valves (Blake Coll., sta. 5). PORTUGAL. Porcupine 
Bank; USNM 63204, 2 right valves (Jeffreys Coll.. Por- 
cupine Exp. 1870). UNITED STATES OF AMERICA. 
Florida; Gulf of Mexico, off Cape San Bias, 309 m depth, 
USNM 323871, 1 left, 1 right valve (sta. 2400); 120.6 km 
east of Davtona, 29°17' N, 79°27' W, 87S m depth, 
USNM 810590, 1 shell and 1 left valve. BRAZIL. Es- 
pirito Santo (IW Marion-Dufresne MD55, May 1987); 
off Concei^ao da Barra, 18°59' S, 37°50' W, 637 m 
depth, MNHN, 10 valves (sta. CB76); off Pontal da 
Regencia, 19°34' S, 38°55' W, 340-360 m depth, 
MNHN 1 valve (sta. CB92). 

Etymology: The specific epithet refers from the 
Greek agues, meaning pure, an allusion to the whitish 
color of the shell. 

Remarks: The above listed examined material that was 
not designed as t\q3es are normally lots with eroded 
specimens, or sometimes they liave aberrant characters. 
This is the case of die MNHN material collected off 
north coast of Espiiito Santo, Brazil. They actually are 
free valves that I'esemble the Pacific species Spinisopella 
costeininens. in having a weakly larger radi;d thread be- 
tvveen middle and posterior thirds, and in lacking ante- 
rior tooth in hinge. As they can represent another spe- 
cies, they are not designed as types; on the other hand, 
the material is not sufficiently well-presen'ed for further 
analysis. Because of they can only represent an extreme 

Page 62 

THE NAUTILUS, Vol. 122, No. 2 

of \ariation of the S. agnes. thev are listed as additional 
examined material of this species. 

Spinosipella finga new species 
(Figures 19-26, 30, 32) 

Vei'ticnrcUa acuticostata: Mariiii. 1974: 242, figs. 5, 6 (noii Phil- 

ippi, 1844). 
Veiiicordia (Htiliris) acuticostata: Rios, 1975: 262, pi. 85., fig. 

1261; 1985: 282, pi. 99, fig. 1.391; 1994: 304, pi. 104, figs. 

1489 (noil Philippi, 1844)" 
Vei-ticordia deshai/esiana: Marini, 1974: 242 (in SNiionvmy); 

Rios, 1975: 262; 1985: 282; 1994: 304 (in synonymy); 

Rosenberg. 2005 (part) (in synonymy) (non Fischer, 


Diagnosis: Shell with 17-18 radial ribs; smooth pre- 
umhonal region ven' narrow (less that Vs ot shell length). 
W'iddi/length ratio in each vahe approximatel)- 0.47. Pos- 
terior cardinal tooth of left valve hinge absent; main car- 
dinal tooth of left \alve low and c\lindrical; main cardinal 
tooth of right \alve lower (about 10% oi valve width) and 

Description: Shei.l: Up to 11 mm, equi\ahe, in- 
flated, both \al\es weakly spiral (1 whorl) (Figures 21, 
22). Color white. Degree of convexity (widtMength) in 
each valve approximatelv 0.47. Umbones located in 
middle region of dorsal surface (Figures 19, 20, 23, 24); 
umbones weakK' spiraled, somewhat high, divergent, 
separated from each odier. Sculpture of strong, uniform, 
arched, radial ribs, 17-18 in each valve. Outer surface 
opatjue, covered by a mosaic of small, blunt, loosely 
aligned spines parallel to radial ribs (Figure 32). Anterior 
edge almost same size as posterior edge. A concaxlt)' 
bearing trans\'ersal ribs oi same width as remaining ribs 
between umbo and posterior edge (Figures 21, 22, 30). 
Anterior, ventral, and posterior edges rounded, tips of 
ribs prominent (Figures 19, 20, 23, 24), fitting with con- 
ca\it\^ in opposite \'al\'e. Inner surface iridescent, whitish, 
glossy, including hinge. Hinge with a somewhat large 
cardinal tooth in right vaKe (Figiu'es 20, 25); cardinal 
tooth stubb\-, tip rounded, flat in ci'oss-section, tooth 
length about 10% ot valve width; correspondent socket in 
left \al\'e shallow, restricted to dorsal surface; this socket 
flanked by small, low, posterior tooth (no anterior tooth) 
(Figures 24, 26). Ligament just anterior to anterior hinge 
tooth, inserted at some distance from midline, approxi- 
mately midway bet\\'een hinge medial edge and umlional 
cavih-. Scars of adductor muscles shallow (Figures 20, 
24); anterior scar elliptical (longer dorso-ventrally), lo- 
cated close to anterior edge, area about '/15 of inner sur- 
face of valve; posterior scar circular, about '/; larger than 
anterior scar, located close to posterior shell edge. Pallial 
hne with a veiy weak pallial sinus, located at wide dis- 
tance from shell edge, about Vs of distance between ven- 
tral and umbonal height. 

Measurements (respectively length, height, width, 
in mm): IloIot\pe: 16.9 x 1.5.3 x 9.6 (single valve); 
MZSP 18752: Parat>pe #1, S.6 x 8.6 x 4.l' (1 valve); 

Parat\pe #2, 11.9 x 11.4 x 5.4 (1 valve); MZSP 18753: 9.5 
X 9.6 X 4.8 (1 valve). 

Tyjje Material: Holot\-pe, MZSP 19345, 1 \-alve, from 
type l(jcalit)^ (RA' \\'. Besnard, GEPID Est. 458, 9 Dec. 
1968. Paratypes, Rio de Janeiro, Cabo de Sao Tome, 
31°08' S, 49°31' W, 182-253 m, I valve, MZSP 18752 
(RAHV. Besn.ard. GEDIP st. 1858, 6 Aug. 1972); 22°34' 
S, 40°29' W, 213 m. 1 \aKc. MZSP 18753; (RA' W. 
Besnard, St. IX, 11 Feb. 1969), 100 m, 2 valves, MORG 
18085 (RA^ Al-MIRANTE Saldanha, Mar. 1972), off 
Solidao, 240 m, 2 valves, MORG 31888 (RA' .A.tlantico 
Sue, Exp. Goltro, 14 Oct.1993). 

Type Locality: BRAZIL. Rio Grande do Sul, off Al- 
bardao, .33°29' S, 50°44' \\'. 200 m, nuiddy bottom. 

Geographic Distribution: Brazil, from Rio de Janei- 
ro to Rio Gi'ande do SuI. 

Etymology: The specific epithet refers to the color 
wliite of the shell, from the Tup\' language: tinga. 

Spinosipella acuticostata (Philippi, 1844) 
(Figures 33-40) 

Hippagiis aciiticostatus Pliilippi, 1844: 42 (pL 14. fig. 19) [fossil 

in Lamati valley, Calabria, Italy]. 
Verticordin acuticostata: Vlicali and Mllari, 1991: .353. 
Spinosipella acuticostata: Pouticis and Bernard, 1995: 143, 


Diagnosis: Shell with 12-13 radial ribs; pre-umbonal 
region narrow, smootli; posterior cardinal tooth of left 
val\'e hinge shallow or absent; main cardinal tooth of left 
\^alve longer and flat (Figures 35. 40); main cardinal tooth 
oi right vaKe shallower (Figui"es 34, 37) (about 10% of 
valve widdi). 

Description: Shell: Up to 24 mm; widtMength ratio 
approximatek' I (Figures 36-38) to 1.5 (Figures 39, 40). 
Degree oi con\e.xit\' (\\idtli/length) in each valve approxi- 
matel\- 0.55. Outer surface spiny, opaque, spines forming 
rachalh' aligned mosaic parallel to ribs (Figures 36, 38, 
39). Scidptiu-e of strong, uniform, arched, radial ribs, 
12-13 in each \alve. Posterior edge about t\\ice as broad 
as anterior edge. A concavitv' bearing trans\ersal ribs 
weakK' broader than ribs of remaining region between 
imibo and anterior edge (Figures 35, 39); pre-umbonal 
region narrow, smoooth (Figure 33), about 10% of shell 
length. Anterior, ventral, and posterior edges forming 
zigzag (Figiu-es 35, 37, 40). Hinge \\ith a large cardinal 
tooth in right valve, stubby, tall (about 10% of valve 
width), broadly pointed, weakly cuned anteriorly (Fig- 
ures 34, 37), circular in section; correspondent socket in 
left valve shallow, restricted to dorsal surface; this socket 
sometimes flanked by small tooth in each side, anterior 
absent or N-erv weak (Figures 35, 40). 

Measurements (respecti\'ely length, height, vridth, 
in mm): RLPG #1: 11.4 x 14.5 x 6.6 (\-alve); #2: 10.0 x 
9.8 X 4.7 (\al\e). 

Geographic Distribution: .\leditcn-ancan. Pliocene 
fossil from soutli Italv (Calabria and Sicily). 

L. R. L. Simone and C. M. Ciinha, 20()S 

Page 63 

Figures 30^7. Spi}io.'iipcllci species. Shells. 30. S, tini:,ii. Holohpe, left valve, dorsal view. 31. S. agues, Parahi^e INV-MOL 2943, 
from Colombia, a specimen of equivalent size of Holotspe of S. tingu for showing major differences (compare with Figiu'e 3(.)); dorsal 
\iew; 17.0 mm; note higher comexits', fewer, tallei' and more spaced ribs. .32. S. tinga, left valve, SEM of Holotvpe, sliowing prickly 
sculpture. 33—40. S. acuticostata. 33-38. Pliocene fossil IVom Messina, Italy, USNM 63202. 33, Specimen 2, doisal-slightK anterior 
\iew. .34. Specimen 3, right valve, detail of hinge. .35-36. Specimen 1, left valve, inner and outer views; 13.5 nnn. 37-.38. Specimen 
2, right \"al\'e, inner and outer views, hinge broken; 12.2 mm. 39-40. RLPC, from Rometta, Italy, left valve, specimen with long 
shape: 19.0 mm. 41-47. S. deshayesiana. 41-43. Paratype 1 of S. ericia AMS 032068, left valve, inner, outer and dorsal views; 3.5 
mm. 44. Paratope 2, right valve, inner view; 2.6 mm. 45, Type specimen of S. japonica ANSP 49639, right valve: 5.2 mm 46-47. 
ANSP 292986 (from India), right valve, outer and inner views; 10.5 mm. 

Page 64 

THE NAUTILUS, Vol. 122, No. 2 

Paleohabitat: Middle and upper Pliocene beds ot 
bathyal environments. 

Material Examined: ITALY. Sicily, Messina, 38°11' 
N, L5°34' E, Segnenza, USNM 6.3202, 2 left, 2 right 
valves. Middle Pliocene outcrops at Rometta, 4 valves, 

Spinosipella deshaijesiana (Fischer, 1862) 
(Figures 41-54, 66, 67, 72-82, 9.3-102) 

Verticordia Dcshat/esiniw Fischer, lS62a: .3.5-36 (pi. .5, fig. 10- 

11) [China Sea]. 
Verticordia japonica A. Adams, 1862: 224. 
Verticordia ericia Hedley, 1911: 96; Prezant, 1998: 421 (fig. 

Spinosipella deshaijesiana. — Poutiers and Bernard, 1995: 110- 

112, 143, 1.59, 161 (figs. 7-9). 
Spinisopella ericia. — Poutiers and Bernard, 1995: 143, 1.59. 
Verticordia acuticostata. — McLean and Geiger, 1998: 109 (non 

Philippi, 1844). 

Diagnosis: Shell with 16-19 radial ribs unilornih' dis- 
tributed, closely packed; pre-umbonal region narrow, 
smooth. Each rib bearing well-developed crests wth 
small, prickl\' granules. Posterior cardinal tooth oi leit 
valve hinge absent; main cardinal tooth of left valve low 
and flat, \\ith insertion of anterior valve edge approxi- 
mately in middle region ot this tooth; main cardinal tooth 
of right valve high (about 10% of vah'e width) and 

Description: SHELL: Up to IS mm. Color white. De- 
gree of convexity (widtli/length) in each valve approxi- 
mately 0.57. Outer surface spiny, spines organized some- 
what radially, parallel to ribs; each rib with well- 
developed crests with small, pricklv granules (Figures 42, 
45, 46, 48-54). Sculpture of strong, uniform, arched, ra- 
dial ribs, 16-19 in each valve (Figures 42, 45, 46, .50), 
somewhat closely packed. Posterior edge about twice 
broader than anterior edge. A concavity bearing trans- 
versal ribs similar to ribs of remaining region present 
between umbo and anterior edge (Figures 43, 53); pre- 
umbonal region narrow, smooth, about 10% of shell 
length (Figures 43, 53). Anterior, ventral, and posterior 
edges forming zigzag (Figures 42, 44, 47, 50, 51, 59, 67, 
100), with tips projected, longer, and narrower. Hinge 
with a large cardinal tooth in right valve, stubby, tall 
(about 10% of valve width), broadly pointed, somewhat 
flat (Figures 44, 47, 51, .59, 67); correspondent socket in 
left valve shallow, restrict to dorsal surface; this socket 
flanked by small posterior tooth, with insertion of ante- 
rior valve edge approximatelv in middle region of this 
tooth (Figures 41, 50), anterior tooth absent (Figures 41, 

LiTHODESMA (Figures 93, 96-99): Saddle-shaped, 
hemi-cylindrical. Dorsal surface concave (Figures 97- 
98), flanking ventral surface of hinge, along '/» of hinge 
length; located just posterior to teeth. Left and right 
edges straight, turned upwards and medially, connected 
with valves by dark-brown Bgament inside umbonal cav- 
ity closer to hinge inner edge (Figures 100-102). Outer 

surface convex, covering dorsal-middle, inter-umbonal 
region of visceral mass (Figure 75). Anterior and poste- 
rior edges concave; anterior edge slightlv deeper and 
with tenuous slope. Both edges covered by opaque, yel- 
lowish periostracum (Figures 96, 101, 102). Lithodesma 
thickness equivalent to that of shell. 

M.^oH Muscles (Figures 72-75, 77, 78, 80, 91, 92): 
Both adductor muscles similar in size and position (Fig- 
ures 72-75), near valve edges; insertion size equivalent to 
1/20 of valves inner surface each; approximately two 
times taller than wide; outer length about half of inner 
length, with insertion in valves greatl)- oblique (Figures 
75, 91, 92). Anterior adductor muscle with anterior re- 
gion about 3 times narrower than posterior region, di- 
vided transversally (dorsoventral) in two similar halves 
(quick and slow components). Posterior adductor muscle 
similar to, but inverted arrangement in comparison to 
anterior adductor muscle; components different, how- 
ever, one of them horseshoe-shaped, occup^dng ventral 
and posterior sides (Figure 77); another component fill- 
ing internal region of muscle, only exposed in posterior 
and dorsal sides (Figm"e 77). Pair of anterior foot retrac- 
tor muscles long and narrow (Figure 80); originating just 
dorsal to anterior adductor muscle in area equivalent to 
1/10 of adductor (Figures 75, 80); nmning ventrallv and 
posteriorly; spreading after insertion in anterior and lat- 
eral regions of foot base. Pair of posterior foot retractor 
muscles similar to anterior pair, but about half narrower 
(Figures 75, 78, 80); originating just dorsal to posterior 
adductor muscle in area equi\'alent to 1/20 of that ad- 
ductor; Rinning \entral and anteriorh'; inserting in pos- 
terior and lateral regions of foot base. Pair of palp 
muscles, septal and pallial muscles described below. Pair 
of foot protractor nmscles absent. 

Foot and Byssus (Figures 73, 74, 80): Foot conical, 
pointed; estimated volume equivalent to 1/6 of that of 
chamber of valves; base located in middle region of ven- 
tral surface of \isceral sac. Bwssal furrow shallow and very 
narrow, length about half ot that ot toot, offset ventrally 
and distally, lying along posterior surface and micUine, 
ending at short distance from foot apex. Byssus found in 
a single specimen, bnjwii, \\ith single filament, narrow; 
proximal end attached to distal region of byssal furrow. 

Mantle (Figures 72-74): Dorsal fusion of mantle 
lobes about Vs of flieir edges, along entire hinge length 
and about Vfe of valves height toward \entral, in both 
sides. Edges of mantle lobes with t^vo folds. Inner fold 
fused between two lobes along entire posterior half (ex- 
cept for siphonal apertures) (Figiu'e 74). Both lobes free 
from each other along anterior half, up to dorsal level ot 
anterior adductor muscle: in this region both folds are of 
similar size, with height equivalent to 'As of valves height. 
Mantle edges thick, muscular, insertion relatively thick in 
pallial line (Figure 72, pm). Pallial muscles originating in 
pallial line in location about '/i from \entro-dorsal dis- 
tance; no clear pallial sinus. Incurrent siphon as aperture 
ot a septum formed by fusion of inner mantle edge folds; 
aperture about Vs of posterior fused region ot mantle. 

L. R. L. Simone and C. M. Cunha, 2008 

Page 65 

Figures 48-63. Spimmpclla species. Shells. 48-54, 56-59. .V. (kshaijcsiinui. adult specimens. 48-53. M\HN (Sta. CP147,5, Fiji): 
12.5 mm. 48. Left valve, outer view. 49. Right valve, outer \'iew. 50. Lett valve, inner view. 51. Right valve, inner view. 52. Posterior 
\iew. 53. Anterior \-iew. 54. Dorsal view, HGLC, from Philippines; 1L8 mm. 55. S. agues parat\pe, BMNH, LS.2 mm. .56-59. 
S\nt\pes of S. dcshaijcsiana MNHN. 56. Outerview, specimen 1, left valve. 57. Outer view, specimen 2 (possihK figured by Fischer, 
1862ai, right valve. .58. Inner view, specimen 1. 59. Inner view, specimen 2; length = 8 nmi. 60-6.3. S. costvmincns Holotype MNHN. 
60. Outer \iew. left \al\e. 61. Outer view, right vaKe. 62. Inner \iew, left valve. 6.3. Inner \iew, right \alve; 17 mm. 

Pase 66 

THE NAUTILUS, Vol. 122, No. 2 

Figures 64-71. Spiiio.siju'llti species. Sliells. 64-65. .S, cnxlcinincns HoloU'jx' MNHN. 64. Right \';il\e, autenui" \ie\v. 65. Left 
valve, anterior wew; 17.0 mm. 66-67. S. ciicia Holotvpe AMS, right valve. 66. Outer \ie\v. 67. Inner \'iew: .5.8 nnn. 68-71. S. 
costemitiens lacking projections on ribs, MNHN (Sta. CP 992, Vanuatu), right valve. 68. Outer-right \'ie\v. 69. Anterior N-ievv. 70. 
Posterior \ie\v. 71. Inner view; 29.0 mm. 

longer dorso-veiitrall)' (Figure 76); walls thick, muscular; 
outer surface flanked by 9 tentacles surrounding siphonal 
aperture; all tentacles of similar size, tiu-ned inwards, 
somewhat conical, tip blunt and rounded, length e(|uiva- 
lent to that of siphonal aperture; single impaired tentacle 
located ventrally; five secondaiy smaller tentacles located 
externally, midway between siphonal aperture and 
mantle edge, of similar size, about Vs of size of major 
tentacles, well separated irom each other, one of them 
located in ventral region of siphonal aperture, other four 
located laterally, in ventral half of siphon (Figure 76). 
Incurrent siplion a small pore located in small elevation, 
approximatelv midway between excuirent siphon and 
hinge; a pair of small tentacles similar to secondaiy ten- 
tacles of incurrent siphons, located laterally, in dorsal 
region of siphon base (Figures 74, 76, 77). Rachal mantle 
gland present along mantle edges outer fold (Figures 7.3, 
92, mg), occupying about half of outer fold volume, situ- 
ated closer to inner surface of this fold. 

Pallial Cavity (Figures 73-75, 94, 95): Occupying 
about 70% of volume of valves. Transversal, horizontal 
septum located approximately midway in animal, i.e., su- 
pra- and infra-septal chambers of equivalent length (Fig- 
ure 72). Paired palps low, wide, bilobed folds (Figures 
73, 81, 95, pp) that occupy anterior third of dorsal sur- 

face oi inira-septal chamber, permanently open as a fun- 
nel. Pair of palp muscles (Figures 72, 74, 79, Im) located 
laterally; originating in anterior region of umbonal cavit)', 
in a distance from origin of anterior foot retractor equiva- 
lent to '/i of anterior adductor muscle height; located in 
same horizontal level of origin of anterior foot retractor; 
size equivalent to 'A of that of anterior foot retractor; 
running ventrally attached to mantle for a distance 
equivalent to Va of \al\'e height; spreading after insertion 
in lateral region liet\veen iimer and outer hemipalps. 
Palp muscles also connect anterior end of septum. Sep- 
tum with two constituents: external one produced by a 
fold of mantle (about % of septum area); internal pro- 
duced by gill (Figures 7.3, 9.5). External septum element 
thick, muscular; posterior muscles originating as a pair, 
just dorsal to posterior adductor nuiscle (Figure 77, ms); 
running ventralK' inunersed in mantle, at some distance 
from each other (equivalent to half of their width) and 
from midline, gradually becoming wider and thicker, in 
anteiior sini'ounding posterior surface of posterior ad- 
ductor nuiscle and lateral edges of excurrent siphon; 
some secondaiy muscular bundles originating from cen- 
troposterior region of posterior adductor muscle uniting 
with main, vertical bundles (Figure 77); muscles spread- 
ing within septum in region between incurrent and ex- 

L. R. L. Simone ;md C. M. Cunha. 200S 

Paee 67 

sp mf by 

ft 'Cfa 

gi au 

Figures 72-75. Spinosipella deshaijesiana . A]iatoni\. 72. W'liole specimen just extracted tioin sliell, right view. 73. Same, nglit 
mantle lobe in its intra-septal region removed, right-slightlv ventral view, left shell valve also shovvai. 74. Same, right mantle lobe 
almost completelv removed, right poition of septum also removed. 75. whole specimen, dorsal-slightly right view, most ol mantle and 
dorsal integument artificiallv shown as transparent, hthodesma (It) shown in its in situ topolog)-. Scale bars = 2 mm. 

Page 68 

THE NAUTILUS, Vol. 122, No. 2 

Figures 76-79. Spinas^ifUa ilcslifii/c.siinici. Anatoniv, 76. Detail ot region ol .siphons, posterior-slightK right view. 77. Peri-anal 
chamber, right view, adjacent region of right mantle lobe sectioned and deflected to show inner surface and muscles, inferior region 
or right mantle lobe removed along median line. 78. Reno-pericardial region and adjacent structrn"es, right xdew, right wall of 
pericardium removed. 79. Whole right view, slio\\'ing topoIo_g>' of genital system, rciio-pericaidial structures, palps, main ganglia and 
inuscles, most structures artificially shown as transparent. Scale liars = 2 mm. 

L. R. L. Siinone and C. M. Cunha, 200S 

Page 69 

Figures 80-82. Spiii(>sii)cllt! desluii/csiaiui. Anatomw SO. Whole right \ievv, emphasizing digestive structures, main musculature 
and main nenous gangUa; topolog)' ot some adjacent stmctures also shown, evei^-thing else represented by transparency. 81. Same, 
anterior region of digestive structiu'es opened longitudinallv, some objects inside stomach preseiA'ed, topology of some adjacent 
structures also sliowii. 82. \'isceral ganglia (left), ventral view, and pedal ganglia (right), postero-doisal \iew. Scale bars = 1 mm. 

current siphons. Outer component of septal muscles in- 
serted in shell just ventral to posterior adductor muscle 
(Figure 72, sm), in area equivalent to Vw ot that of ad- 
ductor muscle insertion. Internal element of septiun 
constituted bv gills. Gill with both demibranchs narrow, 
of similar size, flattened, in same plane of remaining 
septum; both gills surrounding posterior and lateral re- 
gions of foot base (Figures 73, 75, 95). Gill attached to 
remaining septum \ia tissue; gill attachment to foot by 
cfha. Connection between gill filaments of 6-7 longitu- 
dinal, equidistant bridges of similar width of filaments. 
Papilla situated in posterior region of roof of supraseptal 
chamber (Figures 78-80, 94, pi), positioned just ventral 
to \isceral gangUa, internally solid; length about Vw of 
posterior adductor muscle length and about '/s of it in 
width; tip broadly pointed, normally turned to anterior. 
VisCER,\L Mass (Figures 72, 75, 79): Strongly bilobed, 
as internal mould of well-separated umbos (Figures 72, 
7.5). Most dorsal structures, just inside valve apexes, 
formed bv sponge-like connective tissue. Pair of ovaries 
cream in color, occupying central and dorsal regions sur- 
rounding stomach and digestive diverticula, reaching 
dorsal areas up to dorsal sponge-Bke connective when 
fuUv dexeloped. Testes brr:)\\ii, consistence harder, lo- 
cated ventraily and laterallv, totally separated from ova- 
ries: anterior region irregularly digitiform (Figures 75, 

79, ts). Digestive di\'eiticula situated compressed be- 
tween stomach and gonads, color greenish-lieige; occu- 
pying about Vd of visceral volume. Stomach and intestine 
lying in central region, occupying about '/t of \isceral 
volume (Figures 80). Reno-pericardial stn.ictiu-es located 
just anterior to posterior adductor muscle and posterior 
foot retractor muscles, with volume approximateK' Vf, of 
visceral volume (Figures 75, 79). 

Circulatory and Excretory Systems (Figures 75, 
78): Pericardimn located at short distance anterior to 
posterior adductor muscle; with about half of reno- 
pericardial volume, and with a pair of e.xpansions toward 
anterior, surrounding roof of pallial cavity where lies pair 
of auricles. Auricles connecting to anterior end of gills, in 
short isolated ctenidial vein (Figure 75, cv); abruptly 
cuning towards posterior and dorsal; after this curve, 
auricles increasing gradually, surrounding obliquely pe- 
riphery of visceral mass in roof of pallial cavity" (Figures 
72, 75), walls thin, translucent; close to midline auricles 
abruptlv narrowing and connecting to ventricle (Figures 
75, 78); posterior region relatively lobed. Ventricle lo- 
cated in center of pericardium, surrounding intestine; 
relatively narrow. Kidney mostly solid, color dark pui-ple- 
almost black; most of renal gland located just anterior to 
posterior adductor muscle, ventral to pericardium (Fig- 
ures 75, ki); a pair of folds originating from this region, 

Page 70 

THE NAUTILUS, Vol. 122, No. 2 

loinning long roof of pallial, supraseptal cavitw just \en- 
tral and external to auricles (Figures 74, rf), this pair of 
folds with about Vs of supraseptal chamber height, nni- 
ning posteriorh' in middle region of roof of this chamber, 
gradually approaching visceral mass towards anterior, 
fusing to viscera] mass after rimning about '/^ of chamlier 
length (Figure 74). Pair of nephropores as small slits 
located in posterior region of supraseptal chamber, co\- 
ered by posterior end of renal fold, just dorsal to paii' of 
posterior retractor muscles of foot (Figure 74, ne). 

DiGESTi\'E System (Figures 80, 81): Palps partialK- 
described above (pallial cavit}'), widely fused as pair of 
folds along midUne (Figures 95. pp). Mouth central (Fig- 
ure 95, mo), with sphincter relati\ely well de\'eloped. 
Esophagus with about % of \isceral mass length, not 
attached to anterior adductoi' muscle, width about '/i of 
that of anterior adductor nuiscles; wall relati\el\" thick, 
muscular; inner surface with about 20 longitudinal, nar- 
row, low folds as continuation from those of palps (Fig- 
ure 81). Stomach main chamber with about V-i of visceral 
mass volume, elliptical, anteroposterior!)- longer; walls 
thick, muscular (Figure 91, st). Gastric inner surface 
smooth; two pairs of ducts to digestive diverticula 
present, each one located in ventro-lateral region just 
posterior to esophageal insertion. Stomach normallv con- 
taining 3— t isopod crustaceans (Figure 81, cr). Sbide sac 
with about Vi of gastric main chamber \'olume, located in 
middle of gastric ventral wall, somewhat elliptical (longer 
dorso-\entrall\'); cnstalline st}ie occupying entire st)ie 
sac (Figure 81, sy); inner surface of style sac smooth, 
lacking anv fold separating it from intestine; gastric 
shield lacking. Intestine a single sigmoid loop with about 
half of st\ie sac width. Inner surface simple, smooth. 
Intestinal portion crossing through pericardium in some- 
wiiat anteroposterior direction. Rectum attached to dor- 
sal and posterior surface of posterior adductor muscle, 
with about % of remaining intestinal width. Anus simple, 
sessile, located in ventral third oi posterior surface of 
posterior adductor muscle (Figure 77). 

Gexit.\l Syste.m (Partially described above under \'is- 
CERAL Mass); Pair of testes and ovaries converging to a 
single common, short duct, of about '/i5 of \isceral mass 
length. Genital pores small slits located at short distance 
from nephropores (Figures 74, 78, 94, ga). 

Central Ner\'ous System (Figures 80, 82): Cerebral 
ganglia somew'hat triangular, each ganglion with \ nlume 
equivalent to Vis of that of anterior adductor muscle; 
anterior end narrow, possessing thick pair of nenes run- 
ning to pallial region dorsal to palps; pair of ventral 
nei-ves also thick, originated in middle region of ganglia, 
running ventrallv to palps; Posterior end originating 
cerebro-visceral connective (Figures 80, 106); cerebral 
conunissure length about Vs of posterior smiace of an- 
terior adductor muscle. Pair of cerebro-visceral commis- 
svn-es relativel}- thick, running though visceral mass be- 
tween stomach and testes. Pair of pedal ganglia located 
in ventral third of anterior pair of pedal retractor 
nmscles, touching these muscles, both totallv fused with 

each otiier along midline, almost forming a sphere, vol- 
ume of both equivalent to that of each cerebral ganglion; 
pedal nen'es and cerebropedal connectives originating 
subterniinally in posterior surface of gangha. Pair of vis- 
ceral gangha located anterior to ventral surface of poste- 
rior adductor muscle; both also totally fused with each 
other along median line, being somewhat squared in \'en- 
tral view; size equivalent to that of pedal pair of ganglia; 
cerebrovisceral connectives and siphonal nenes located 
in vertices. 

Measurements (re.spectively length, height, width, 
in mm): HGEC: 11.5 by 12.2 by 12.0; MNHN (Sta. 
DWll): 15.7 bv 17.7 bv 9.2 (valve); MNHN (Sta. 
CP889): 19.7 bv'l7.0 bv 9.1 (vaKe). 

Geographic Distribution: South and Central Indo- 
Pacific in 140-805 ni tlepth. 

Material Examined: Paratvpes of S. ericia: AUSTRA- 
LIA; South Cape Wiles. 174-183 m, 35°39' S, 136°40' E, 
AMS 032068, 1 left, 1 right valves (Zoological Results of 
the F.I.S. Endeavour, 28 Aug. 1909). 

Other Material Examined: Holotvpe of S. japonica: 
JAPAN. ANSP 49639, 1 shell. MNHN. SW PACIFIC. 
Lovaute Islands, 16 lots [122 v]. TONGA IS. 12 lots [59 
V ]. GUAM. Marianas Islands, 3 lots [15 v]. AUSTR.ALIA. 
South Cape Wiles, 1 lot [6 v]. NEW CALEDONIA. 
South, 3 lots [7 specimens]; Banc Esponge, 2 lots [3 
specimens]; Chesterfield Plateau, 1 specimen. PHILIP- 
PINES. Aliguri Is. 2 lots [1 specimen and 3 v]; Bohol 
Sea, Off Balicasag Island, 1 lot [1 v]. FIJI. 1 specimen. 
MYANMAR (BURMA). 1 lot [5 v] Preparis North Chan- 
nel, 1 lot [4 v]; N.W. of Tavov I., 1 lot [11 v]. ANDA- 
MANS SEA. 1 lot [1 v]. THAILAND. Phuket I., 1 lot [11 
v]; Andaman Sea, 1 lot [1 v] (Details in Simone and 
Cunha, in press.) 

Spiiiosipclla costcniinciis (Poutiers. 1981) 
(Figures 60-65, 68-71. 83-92, 103-108) 

Vciiicordia (Spinosipclhi) costeinincns Poutiers. 1981: .351 (pi 

4. figs 1-4, te.\t fig 5). 
Spinosipella costeminens. — Poutiers and Bernard, 1995: 110, 

14.3. 1.58 (figs. 1-2). 

Diagnosis: Shell with 16-17 tall radial ribs, those 
more posterior to iiiiildle surface veiy taller, normallv 
possessing blade-like projectit)ns along tip; 3-4 more 
posterior abruptly lower, preceded bv a ven- tall, carina- 
like rib. 

Description: SHELL: Up to 30 mm. Color white. De- 
gree ol eonvexitv'- (widtli/length) in each valve 
matelv 0.50. Outer surface prickly, with somewhat cha- 
otic organization (Figures 60, 61, 68-70). Sculptured bv 
strong, uniform, arched, radial ribs, from 16 to 17 in each 
valve (Figures 60, 61); ribs increasing from region ante- 
rior to umbo to region betw^een middle and posterior 
thirds, last ribs in this region taller and more separated 
from each other, last one on a weak carina (Figure 70); 
larger ribs normalK' possessing blade-like, projection 

L. R. L. Sinione and C. M. Cunha, 2008 

Page 71 

Im au oy 

mg mb by 

mb ^^ 9' ft pm 

Figures S3-86. Spiiuisipclla costemint'iis. AnatuiiiN. 83. Whole specimen willi right \al\e extracted, light \ie\v. 84. Specimen 
extracted from shell, posterior view, showing siphonal area. 85. Whole right \riew, some portions of right mantle lobe exti'acted, 
particularh- regions \entral to septum, and ventral and dorsal to I'enal fold (rf) to expose inner surface; cerebral ganglion (ce) seen 
bv transparencv. 86. Same, \entral-slighdy right view. Scale bar = 5 mm. 

along tip; posterior third as a slope, haxdng 3-4 ribs simi- 
lar to those of anterior region; blade like projection ab- 
sent in some specimens (Figures 68-71). Posterior edge 
about t\\"ice broader than anterior edge. Between umbos 
and anterior edge a concavity bearing transversal I'ibs 
similar to ribs of remaining region (Figures 64, 65); prc- 

iiinbonal region nairow, smooth about 10% of shell 
length (Figures 64. 65, 69). Anterior, \entral, and poste- 
rior edges forming zigzag (Figures 62, 63, 71, 103), vvith 
tips longer and narrower projected in those middle and 
larger ribs. Hinge with a large cardinal tooth in right 
valve, stubby, tall (about 10% of valve \wdth), broadly 

Page 72 

THE NAUTILUS, Vol. 122, No. 2 

Figures 87-90. Spiiio.sipeUa costeminens. Anatomy. 87. Whole liglit view, mainly showing digestive tubes and main gangha, 
topolog\' of some stmctures also showai. Scale bar = 5 mm. 88. Scheme of layers of tissue in indicated region of stomach. Scale bar 
= 0.5 mm. 89. Fore- and midgut opened longitudinalh' for exposing inner surface (same scale of Figure 87). 90. Foot, \entral-slightlv 
posterior view, sectioned transversally in two levels to sliow inner layer of tissues. Scale bar = 1 mm. 

pointed, somewhat flat (Figures 63, 71, 103): correspon- 
dent socket in leit valve shallow, restrict to dorsal sur- 
face; this socket flanked by small posterior tooth, with 
insertion of anterior valve edge approximateK" in middle 
region of this tooth (Figure 62), anterior tooth absent 
(Figure 62). 

Additional details for this species see Poutiers (1981), 
Poutiers and Bernard (1995). 

LiTHODESM./v (Figures 103-105): Characters similar 
to those in preceding species, differing in being propor- 

tionallv shorter and wider (Figures 104-105). Length 
about Vs to Ve of hinge length, and about 1.5 times wider 
and long. 

Main Muscle System (Figures 83-87): Characters 
similar to those in preceding species. Anterior adductor 
muscle about 20% dorso-ventrally longer (Figures 83, 85). 

Foot .'Vnd Byssus (Figures 85, 84, 90,"l07): Shape 
and disposition similar to those in S. deshaijesiana. Byssal 
gland relatively deep, lanming immersed in ventral re- 
gion of pedal musculature at about half of byssal furrow 

Figures 91-108. Spiiuisipclhi species. Anutomv. 91. S". costeiniucris, middle liorizontal, longitudmal section through visceral mass 
at same level as pericardium (MNHN sta. CP767, Mallory, 5 |J.m). Scale bar = 2 mm. 92. Same, detail of posterior region of mantle 
border. Scale bar = 1 mm. 93-102. S. deshaijesiana . 93. Detail of hinge region of left valve vvidi lithodesma (It) still attached, right 
view. Scale bar = 2 mm. 94. Detail of posterior region of supraseptal chamber, right view, right mantle lobe removed (MNHN sta. 
DW1499). Scale bar = 1mm. 95. Infraseptal chamber roof, ventral view, right mantle lobe removed (MNHN sta. CP767). Scale bar 
= 2 mm. 96-99. Lithodesma (MNHN sta. DW739). Scale bar = 1 mm. 96. Ventral view. 97. Dorsal view. 98. Posterior-slightlv dorsal 
view. 99. Posterior view. 100. Same specimen, emptv shell, ventral view, valves slightly open, lithodesma still in situ. Scale bar = 2 
mm. 101. Same, detail of hinge and lithodesma. 102. Same, ventral-slightly anterior view. 103-108. S. costeuuncns. 103. Shell, 
ventral view, valves open, lithodesma still attached to left valve (VINHN sta. CP1460). Scale bar = 2 iiiiii. 104-105. Ijithodesma, same 
lot (other specimen), dorsal and ventral views respectively. Scale bar = 1 mm. 106. Detail of anterior region, right view, integument 
removed, mainly showing right cerebral ganglion (ce) (same lot). Scale bar = 1 mm. 107. Infraseptal chamber roof, ventral view, right 
mantle lobe removed (MNHN CP767). Scale bar = 1 mm. 108. Detail ol posterior (siphonal) region, po.sterior view (MNHN 

CP1460), Scale bar = 2 mm. 

Page 74 

THE NAUTILUS, Vol. 122. No. 

length towards dorsal (Figure 90, by). Thick muscular 
layer surrounding a nucleus ot conective tissue (Figure 
90, cj). 

Mantle (Figures 84-86, 92, 108): Characters similar 
to those in preceding species, with following distinctive 
characters. Pair of secondary tentacles positioned be- 
tween incurrent and excurrent siphons (Figures 84, 108); 
remaining tentacles similar in size and position. Ventral 
pair of tentacles of incrnrent siphon generally symmetri- 
cal. Zigzag formed by mantle edge haxdng secondaiy 
folds positi(jiied in more distal tips, possibly elated to 
taller radial shell ribs (Figure 108). Radial mantle gland 
(Figure 92) similar to S. deshai/esiana. 

Pallial Cavity (Figures iS5-86, 107): Characters 
similar to those in preceding species, except for wider 
platform bet^veen posterior region of gills as part ol sep- 
tum (Figure 107). 

Visceral Mass (Figures 85-87): Characters similar 
to those in preceding species, differing mainly by wider 
region separating pair of renal folds in supraseptal cham- 
ber (Figure 85). 

Circulatory and E.xcretory Systems (Figures 85, 
91): Pericardium and heart with characters similai' to 
those in S. dcshmjcsiana (Figure 91). Kidnevs of similar 
features, differing niainK' bv enlargement of pair of renal 
f(jlds (Figures 85-86, rf), taller and wider, almost divid- 
ing supraseptal chamber in hvo — internal and external — 
halves. Height of renal fold about 80% of that of su- 
praseptal chamber heiglit. In addition to an enlargement, 
both lenal lolds still haw postei'ior end in more anterior 
position and wider sepaiiition between folds and \ isceral 
mass (Figure 85). 

Digestive System (Figures 87-89): Characters simi- 
lar to those in preceding species. Esophagus with about 
■/•! of visceral mass length, rimning horizontally, peipen- 
dicular to pf)sterior surface of anterior adductor muscle 
(Figure 87, es). Stomach main chamber with longer re- 
gion as a l:)lind-sac projected posteriorly. Castric wall 
constituted In external la\er of weak connecti\e tissue 

(Figure 88, cj), two thick muscular layers of similar size, 
with outer layer of longitudinal muscle and inner layer of 
circular muscle (Figure 88, lo and cm). Inner surface of 
stomach (Figure 89) with posterior end of esopliageal 
folds clearly more evident that together form a flat fold. 
Another ventral fold surrounding apertures to digestive 
diverticula. Gastric sh'le narrower (about Vfe of gastric 
width); internally a pair of tall folds separating intestinal 
from stv'le sac components (Figure 88, ss, in). 

Genital, System: Characters similar to those in pre- 
ceding species. Separated masculine and feminine com- 
ponents of gonad showai through histological sections in 
Figures 91(ts, ov). 

Central Nervous Syste.m (Figures 87, 106): Three 
ganglia with similar localization and size to those of pre- 
ceding species. 

Mca.surcmcnts (respectively length, height, width 
in mm): MNHN (Sta. 1.3(Vl): 22.0 bv 28.1 bv 12.5 
(valve); MNHN (Sta. CC996): 20.0 by 24.3 by 14.3 
(valve); MNHN (Sta. CP992): 19.6 bv 23.3 by 12.6 

Geographic Di.stribution: 'I'lopical West Pacific. 

Depth Range: 750-925 m. 

Material Examined: Holotype; Additional material 
(MNHN): SW PACIFIC. 4 lots [32 v, 11 specimens]; 
Wallis Is., 6 lots [15 v]; Banc Combe. 5 Lots [28 v]; 
Fortuna Is., 5 lots [18 v]; Banc Watenvitch, 2 lots [3 v]; 
Banc Tuscarora, 29 lots [63 v]; South Vanuatu - Monts 
Gemini, 4 lots [4 v, 1 specimen]; TONGA IS. 8 lots [52 
v]; Eua Is. 6 lots [12 v]; Seamount, 6 lots [29 v]; South of 
Nomuka group, 1 lot [25 v]; Ha'apai Group, 2 lots [4 v]; 
N Ha'apai group, 3 lots [ 6 v]; NW Tongatapu, 3 lots [16 
v]; SW Tongatapu, 5 lots [22 v]; Tongatapu, 6 lots [8 v]; 
S. Nomuka group, 2 lots [6 v]; Vava' group, 1 lot [2 v]; 
NEW CALEDONIA. 5 lots [5 v, 5 specimens]; Lord 
Howe, 1 lot [1 v]; Banc Nova, 2 lot [8 v, 1 specimen]; 
North New Caledonia, 10 lots [tota 20 v]; South New 

+ .S', aciiliciisiulu * •''' li'ifit-t • .f. co.sWminens 

■* S dcshaycsiuna ^ S. agues 

Figure 109. (.;c(j(j;rapliic distrihiitioii ul Sf)iiu)sii)cll(i .spp. 

L. H. L. Siinone and C. M. Cunha, 2008 


Caledonia, 1.3 lots [46 \, 1 specimen]; off Norfolk, IS lots 
[98 \]: Banc Esponge, 11 lots [144 \']; Banc Kainion- 
MaiTi, 9 lots [38 \]; Banc Antigonia, 1 lot [1 \]; Banc 
JunieauA\"est, 4 lots [17 \]; Banc Intron\-able, 7 lots | l(i 
v]; BiUic St\'laster, 1 lot [1 \] ; X'olcans Hunter and Mal- 
tliew, 2 lots [2 v]; S.E. New Caledonia, 2 lots [2 \]; East 
New Caledonia, 6 lots [30 \] Banc Capel, 1 lot [lota 12 \]; 
Banc Kelso, 1 lot [6 v]: 1. Lo\aute, 22 lots [44 \]. FIJI. 
South of \'iti Le\ai, 42 lots '[328 v]; Southeast of \''iti 
Le\u. 17 lots [.57 \]; Bohol/Sulu Seas, 2 lots [.5 v]; Bohol 
Sea - Balicasag Island, 3 lots [5 v]; Bordau, 1 specimen; 
T.AIWAX. B;i.shi channel, 2 lots [3 \]; South China Sea, 

1 lot 

East Taiwan, 2 lots [.5 v]. (Details in Simone 

and Cunh;i, in press.) 


The Genius Spixosipella \'\'ithin the Verticordiidae. 
Despite their larger size, the prickl\' outer surface of the 
shell, and the reduction of the lunule, which differenti- 
ates SpinosipcIIa from the remaining verticordiids, this 
taxon has traditionally been considered a subgenus of tlic 
genus Veiiicordia. This set of characters is sufficient in 
m\" opinion to idlocate SpinosipcIIa as a separate genus. 
This \ie\\- \\as previously defended by the author of the 
genus (Iredale. 1930) and by Poutiers and Bernard 
(199.5). Other cUstinctive characters are the spiral uni- 
bones (Figiu-es 5, 7, 21, 22, 33, 54, .53), the tall, some- 
what uniform radial sculpture, triangidar in section; and 
the obesitv of die valves. The spiial umliones and the 
obesit\' of Spinosipella are quite similar to those in the 
fossil genus Pcccliiolia Savi and Meneghini in Vlurchison, 
1850 [t\j)e-species (b)' monot\'"py): Pecchiolia argentca 
Sa\i and Meneghini in Murchison, 1850 (= Chama ari- 
etina Brocchi, 1814) middle Tertian', Europe] (Keen, 
1969: 857), from \\"hich SpinosipcIIa differs in ha\"ing 
well-developed ribs and zigzag edges. 

The fuU genus status of SpinosipcIIa is based on the 
differences with the t)pical Verticordia sensu stricto 
[t\j)e species (bv monot\|3v) Veriicordia cardiifonnis 
Sowerbw 1844]. such as the higher size and obesit)' of the 
\'al\es: the additional development of the prickly surface 
(wliich also covers the radial ribs, whereas in Vciiicordia, 
when a pricklv surface is present, it does not cover the 

ladial iil)s). the absence ol lunule; tlie spiral fashion ol 
both vahes; and the similarity among the radial ribs (rep- 
icsentatives of Vci-ticordia usually have an unusually 
huger rib or space between ribs). The same set of char- 
acters also differentiates Spinosipella from TiigonuUna 
d'Orbigny, 1842 [type species (by monotyj^y) T. ornata 
tl'Odiigny, 1842] in the sense of Jung (1996: 46-47). 

Representatives ot SpinosipcIIa also resemble those of 
the genera Haliris Dall, 1886, and Euciroa Dall, 1881, by 
their larger size, conve->it)', and prickly shell surface. Spi- 
nosipella difters from those two genera, however, in the 
higher degree of convexitv, reflected in more obese 
shells in its species; in the much more developed and 
taller radial ribs; higher degree of spiralization of die 
valves; and in the expansion of the ribs bevond the shell 

Further analvsis on the verticordiid svstematics and 
phyiogeny can be found in the literature (e.g., Pelseneer, 
1888; Salvini-Plawen and Haszprunar, 1982; Bieler and 
Mikkelsen, 1992). 


The diiferentiation between the five species of Spino- 
sipella is sununarized in the respective chagnoses and in 
Table 1. The degi'ee of differentiation in the samples of 
each species examined allows for specific separations. 
The number of radial ribs is tlie most notable feature; 
despite certain a small amount of intraspecific variation, 
the number of radial ribs is somewhat constant in each 
species, at least in specimens of larger size. The fossil S. 
aciiticostata is the species with fewest ribs, 12-13 (Fig- 
ures 36, 38, 39), while S. dcsliai/esiana has the largest 
number of ribs, 16-19 (Figures 46, 48, 49, 54, 53). The 
other species possess an interniedian' number of ribs. 
The species of Spinosipella usually have radial ribs of 
relatively uniform size; the single exception is S. costcm- 
inens, which has ribs clearly increasing posteriorly; in the 
posterior shell slope, however, the ribs abruptly reduce 
in size, although in some specimens, particularlv in the 
young ones, this cliaracter is not so clear, i.e., the ribs are 
somewhat uniform-sized. The shell inflation is well de- 
veloped in most Sj)inosij)ella species, but this is clearer in 

Table 1. Comparison ot cliaracters between tlie five studied species of Siiinosii)cIla. 



Sjiinosipella agues 




Distribution Mediterranean Tropical W'. Atlantic; S-SE Brazil 

Caribbean; to SE Brazil 

Shell Inflated Strongly HigliK \\'eakl\- 

Sculptured between Radial Disorganized Radial 

radial ribs 

Pricklv ribs outer Rough Rough 


Number of Ribs 12-1.3 1.5-17 17-18 

Size (mm: 20 20,2 10.4 

South and Central Tropical \\'est Pacific 


Strongly HighK- 

Radial Disorganized 

^^'eakl\■ pnckl\ Strongly pnckK Rough 



Page 76 

THE NAUTILUS, Vol. 122, No. 2 

the larger specimens; \\iiile the \ouiig specimens are 
considerably flatter (Figures 41-45). The prickly outer 
shell surface is an outstanding character of the Sj)infl- 
sipella species; however, this character is consenative 
among the five species; the single exception is the rela- 
tively chaotic arrangement in S. agncs (Figiu'e 8) and S. 
costeminens . while in the remaining species a radial ar- 
rangement is apparent (parallel to the radial ribs) (Figure 
32). The Pacific species S. cle.sliai/esiana has much larger, 
crispy prickles along the tip of the ribs (Figures 42, 45, 
46, 48, 49). This is lacking in the remaining species, 
e.xcept in some very young specimens (e.g., USNM 
810889, S. agues, 6 mm), where the prickles, however, 
are not fully developed. The prickly surface is strongly 
damaged in eroded specimens (Figure 55), becoming 
almost completely smooth. Spinosipella dcshaijcsiana, 
perhaps because of this character, has the distal tips of 
the zigzag edges of the shell even longer and more pi'o- 
jected (Figures 41, 44, 47, 50, 51, 59, 67). The series of 
radial ribs is interrupted in the region between the um- 
bos, where a triangular smooth area appears. This area is 
particularlv large in S. agncs (Figures 7, 9), but is prac- 
tically absent in S. tinga (Figures 21, 22); it is narrow in 
the remaining three species. The size of the specimens 
appears to be another distinctive feature, as S. tinga is 
small (around 10 nun), whereas the remaining species 
are larger (20-30 mm). The hinge does not vaiy much 
between the Spinosipella species; however, some par- 
ticularities exist. The posterior tooth of the left valve is 
well developed in S. agncs [Figures 4, 10, 12, 27, 28 
(arrow)], ven" low in S. acuticostata (Figures 35, 39), and 
practically absent in remaining species (Figures 20, 25, 
50). The tall and pointed cardinal tooth of the right valve 
is more de\'eloped in S. agncs, in such it is also shaipK' 
pointed and cui"ved (Figures 3, 10, 12). In the remaining 
species this tooth is weakly shorter and more rounded 
(Figures 26, 34, 47, 51). 

The geographic and stratigraphic distribution are 
somewhat mutually exclusive for most of the species 
(Fig. 72); Spinosipella actiticnstata is the onlv Mediter- 
ranean species, S. agncs occurs from Florida to Rio de 
Janeiro, S. tinga is found from Rio de Janeiro to Rio 
Grande do Sul, along the Brazilian coast. The fine- 
resolution distribution of the Indo-Pacific species is still 
unclear, but S. cleshaijesiana and S. costeminens, appear 
to be sympatric. Spinosipella acuticostata is a fossil spe- 
cies, occurring in Pliocene strata, while the remaining 
species are found in the Recent. Apparently no Recent 
Spinosipella occur in the MediteiM'anean. 

All samples of Spiuosipella from the Atlantic and 
.VIediterranean have previously been accepted as belong- 
ing to the single species S. acuticostata (e.g., Abbott, 
1974; Abbott and Dance, 1983; Rios, 1994). However, 
analyses of the concholf)gical, geographic, and strati- 
graphic differences, show that the separation into three 
species is warranted. As the shape changes considerably 
during ontogeny, a specimen of S. agues at same size as 
the holotype of S. tinga was chosen to show the differ- 

ences between those species. Figures 28-31 illustrate 
these differences. Spinosipella agues has fewer, taller, 
and more widelv spaced ribs than S. tinga (Figures 19, 
29). The shape of the shell edge is much more uniform 
in S. tinga than in S. agues, in that the tips of die ribs are 
more expanded, extending longer beyond the shell mar- 
gin (Figures 20, 24, 28). The posterior cardinal tooth in 
the hinge of the left valve is present in S. agues, in ani- 
mals larger than 5-6 mm, while this tooth is never 
present in S. tinga (Figures 20, 25, 27-28, arrow). The 
degree of convexity is higher in S. agues and in S. tinga 
(Figures 30, 31); S. agues has a degree of convexity 
(widtli/length) in each \"al\'e of about 0.57, while S. tinga 
it is 0.47. " 

The comparison of the previously valid species Spiuo- 
sipella ericia. including paratopes (Figures 41-44), and S. 
cleshaijesiana, does not reveal any distinction between 
them. Normally, specimens of smaller size were identi- 
fied as S. ericia, and the large ones as S. cleshaijesiana. 
But examination of shell features along a growth series 
show a complete gradient linking the two taxa. The same 
lack of distinction is found in the literature for both spe- 
cies, including the original descriptions. For these rea- 
sons, despite the fact that S. ericia is the type species of 
the genus, the older name S. dcshaijesiaua should be 
used. Furthermore, a type specimen of S. japouica was 
also examined (Figure 45), confirming the synonymy of 
this species with S. deshai/esiaua. 

The distinction between the Pacific species Spiuo- 
sipella cleshaijesiana and S. costeiniueus is not always 
easv. With the large quantity of specimens kindlv pro- 
vided by the MNHN (Paris), it \vas possible to analyze 
the degree of variation of both species. Spiuosipella 
costeminens mostly has samples with shell possessing the 
outstandingly large, carina-like spiral ridge between the 
middle and posterior thirds of the shell, but sometimes 
this ridge is not so different from the others, and the 
animal become more rounded, similar to S. cleshaijesi- 
ana. The distinction is based mainly on the pi'esence of at 
least a weak carina in the region between middle and 
posterior thirds, and also by the more robust ridges of S. 
costeminens specimens (Figures 60, 61), while those of S. 
deshaijcsiana lack any clear radial carina and tlie ridges 
are more delicate, uniform and apparently close from 
each other (Figures 46, 48). 

The lot USNM 63200 includes 3 valves (2 left and 1 
right), collected in Barbados, the knowai geographic dis- 
tribution of Spiuosipella agues. However, die right valve 
has the characters of S. cleshaijesiana, instead of those of 
S. agues. In addition, is looks different in the state of 
consenation, color anil associated sediment, from the 
other 2 \al\es of the same sample. 


More in-depth anatomical descriptions and discussions 
on verticordiids are provided by Alk'u and Turner 

L. R. L. Simone and C. M. Cainlia, 2008 

Page 77 

(,1974), wlu) stiulit'd 19 species of several <i;enera. How- 
e\"er, no iniorniation on the anatonn- of tlie genns S/)/- 
nosipcllo is ioniul in the Uterature. Although anatomical 
information is available here onK lor two of the five 
species of the genus (ot course one ol them is a Pliocene 
fossil), some SNstematie inferences can be made based on 
the scenario gixen in the literature the N'erticortliidae and 
related families (Allen and Turner, 1974, and others, e.g., 
Fisher, 1S60, lS62b; Pelseneer, 1888; Nakazima, 1967; 
Allen and Morgan, 1981). Besides the conchological 
characters discussed alio\-e, some anatomical features are 
possibK' restricted to Spinosij)clla, such as: the \\ide litli- 
odesma (Figin-es 93, 96-104, It); the simplified siphonal 
tentacles (Figin-e 108), which normally have secondaiy 
papillae; the papilla on the root of tlie e.xcurrent chamber 
(Figures 78-80, 85, 94: pi); the absence of incnrrent 
\al\e in infrasept;d chamber. However, wide lithodesnia 
ha\e been reported for Policordia Usbetae Knudsen, 
1970 (fig. 90), which has \en' different shell and pallial 
tentacular characters. The study on the incnrrent sipho- 
nal structures is of particular importance in septibnmclis, 
as the mochfied incurrent siplion is the main strnctnic 
used in prey capture (Morton, 1987). 

On the other hand, some features appear to be char- 
acteristic of Wrticordiidae, such as: elongation of lateial 
region of kidne\s; tlie muscidar stomach (see also Pur- 
chon, 1956, 1963); tlie separation between testis and 
o\ai'v. B\' the pro.ximity of the esophagus from anterior 
adductor muscle, by the lack of inciu'rent valve, and h\ 
tlie simplified buccal structures, e.g., lack of buccal caviU- 
and tongue, it is possible to suggest that Spinosipclla is a 
basal taxon inside Verticordiidae. Unfortunately, no 
member of tlie genus was analyzed in the recent com- 
parative studies on anomalodesmatans (Hai"per et al, 


The autliors are grateful to the researchers who loaned 
die material for this study: Winston Ponder and Ian Locli 
(AMS) for ts'pes of Spinosipella eiicia; Nestor E. Ardila 
(MHXMC)'and Emilio Garcia (EGG) for S. a^nci (Go- 
lonibia) and S. deshaijcsiana (Philippines); Hariy G. Lee 
for S. aones (Florida) (MHNMG) for a large lot of S. 
agnes (Golombia); and especialtv^ to Philippe Bouchet 
and Plifhppe Maestrati, MNHN, for the loan of a huge 
quantit\- of lots coming from se\ei-al places of tlie vvorld, 
niosth' from tlie Indo-Pacific. For thorongli comments 
and additional information about S. acuticosiata we 
tliank Rafael La Penia. For material oi Haliiii fisheriana 
we thank to Daniel Mansur Pinipao (PPG-BAN, 
UFRGS). For help with SEM procedures, we tliank Lara 
Guimaraes (MZSP). For Rachel Collin, Sniitlisonian In- 
stitution at Panama, we thank for the help in the te.\t and 
language. We tliank also both referees and the Editor for 
die thoughtful correction on the manuscript. This project 
is supported b\- FAPESP (Funda^ao de .\mparo a Pes- 
quisa do Estado de Sao Paulo), procs. no. 03/05860-6, 

04/02333-8, and a "Treinarnento Tecnico 3" grant, imder 
the snpenision ol Antoni;i Geci'lia Z. Aniaral and Luiz 
R.L. Simone. 


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THE NAUTILUS 122(2):79-93, 2008 

Page 79 

Late Eocene Co)uis (Neogastropoda: Conidae) 
from Florida, USA 

Jonallian K. Hendricks 

Department ot Geolog\' 
Unixersih' of Kansas 
Lawence, KS 66045-7613 USA 

Roger VV. Portell 

Division ot Invertebrate Paleontology' 
Florida Museum oF Natural Histoiy 
University of Florida 
Gainesville, FL 3261 1-7S(){) USA 


The neogastropod genus Conns is likelv tlie most diverse ma- 
rine animal genus, but has an Eocene to Pleistocene fossil 
record tliat remains poorly understood. We discuss the fossil 
record of Eocene Conns from Florida and recognize three 
species: Conns sanridens Conrad, 1833, and two new species. 
Conns palmerae and Conns allcni. We also re-describe C. san- 
ridens, identilV its likely sraonyms, and add new information 
about its geogi'aphic range. The new species C. pahncrac is 
restricted to the upper Eocene Ocala Limestone of Florida, 
while C. alh'ni new species occurs in the upper Eocene Ocala 
Limestone of Florida and the upper Eocene Moodys Branch 
Formation ol Louisiana. 

Additional Kci/iuords: Mollusca, Fossil, Gastropoda, Ocala 

Kohn and Anderson, 2008) lists names that have been 
applied to Eocene Couus trom the southeastern United 
States. Our puipose Ls not to revise the svstematics and 
describe the fossil record of all of these taxa, but rather 
to focus on those species occurring in Florida, which 
have thus far received onlv preliminai-v attention (Rich- 
ards and Palmer, 1953). This lack of attention is likely 
related to the presei'vational nature of these Floridian 
specimens, almost all of which are presented as internal 
and/or external molds rather than shell material. While 
we focus on Conns fossils trom the upper Eocene Ocala 
Limestone of Florida, vye also consider likely synonyms 
of C();;//.s sduiidens Conrad, 1833, a widespread and tem- 
porallv persistent t;L\on tliat will require additional liiture 


\Mth over 1500 described fossil and extant species, Co- 
mis Linnaeus, 1758 (cone snails) mavbe the most diverse 
marine animal genus (Rockel et al., 1995). Molecular 
sequence data have offered valuable insights into the 
relationships of extant Conns species (e.g., Diida and 
Kohn, 2005), but the deep evolutioiian- histon' ot Conus 
remains unclear: although about 1000 fossil species have 
been described (Rockel et al., 1995), manv ot tliese are 
hkelv sviionvinous (e.g., see Hendricks, in press). Many 
fossil Conus species are based onlv upon tvqje specimens 
and/or were described without reference to comparable 
material from nearb), contemporaneous fossil faunas. 
Unraveling the early evolutionary history of Conns will 
onlv be possible bv careful examination and intei^preta- 
tion of its rich and well-preserved fossil record. 

Some of the oldest substantiated Conus fossils are 
from the early Eocene (Ypresian) of England and 
France, and possibly Pakistan (Kohn, 1990). Occurrence 
records of middle Eocene Conus in North America show 
that the genus had achieved a wide geographic distribu- 
tion soon after its first appearance in the fossil record. 
For example. Table 1 (which is derived in large part from 


Formally named bv W. H. Dall (in Hall and Harris, 
1892), the Ocala limestone represented all limestones 
e.xposed in central Florida at that time. A late Eocene 
age was not determined for the Ocala limestone, how- 
ever, until Cooke (1915) correlated the unit with deposits 
in Mississippi and Alabama. Based on lithologv' and micro- 
fauna, Apphn and Applin (1944) divided the Ocala Lime- 
stone into an upper member and lower member. X'ernon 
(1951) redefined the Ocala Limestone by restricting it to 
Applin and Applin's (1944) upper member and placed 
their lower member in the Moodys Branch Formation. 
Further, Vernon (1951) subdivided the Moodvs Branch 
Formation (as pertaining to Florida) into the basal Inglis 
Member and overlying Williston Member. Puri (1953) 
renamed Vernon's (1951) Ocala Limestone (restricted) 
as the Ciystal River Formation. Later, Puri (1957) el- 
evated the Ocala Limestone to group status (Ocala 
Group) — consisting, from oldest to youngest, of the In- 
glis, Williston, and Ciystal River formations — and eUmi- 
nated the Moodys Branch Formation designation for 
Florida Eocene deposits. Based on macro- and micro- 
fossils, Toulmin (1977: 117) correlated the Inglis For- 
mation with the lower Moodys Brancli Formation in 

Pas;e 80 

THE NAUTILUS, X'ol. 122, N( 

Table 1. Names applied to Eocene Conus from tlie U.S. Coastal 
considered sxiionvins of Eocene t;ixa. Tvpe specimen abbreviations 

Plain, including names of some younger species that are here 
H, holotvpe; L, lectotvpe; and F, figured. 



Tvpe locality 

Present disposition 
according to this study 

C. sauridens Conrad, 1833 
C. claibornensis I. Lea, 1833 
C. gymtus Morton, 1834 
C. parvus H. C. Lea, 1841 
C. iiiiitilatiis Tuomey, 1852 
C. toiiilis Conrad, 185.5 
C. aheatus Conrad, 1865 

C. subsauridens Conrad, 1865 

ANSP 148.54 (L) 


ANSP 211 (H) 

ANSP 13161 (L) 


ANSP 13196 (H) 

ANSP 1.3446 (L); See MacNeil 

and Dockeiy (1984, Pi. .38, 

Fig. 26) 
AN,SP.5.3812 (H) 

C. ptdchenimiis Heilprin, 1879 AMNH-FI 10175 (H?) 

C. jcicksoncnsis Meyer. 1885 

Unknown, but specimen 
puqjorted to be holotype 
figured by Harris and 
Palmer (1947, pi. 62, fig. 17) 

PRI 264.36 (H) 

C. dcperditus var. suhdiadcma 

de Gregorio, 1890 

C. improiidus de Gregorio. 1890 Lost (see Palmer and Brann, 


C. (Conospinis) graiio/ww de Lost (see Palmer and Brann, 

Gregorio, 1890 1966) 

C. smithvillensis Harris, 1895 BEG 34656 (H) 

C. cormacki Hadiison, 1944 ANSP 16415 (H) 

C. (Leptoconus) santaiider 

Gardner, 1945 
C. (Leptoconus) liai'dliti 

Gardner, 1945 
"Conus sp. A" Palmer in 

Richards and Palmer, 1953 
"Conus sp. B" Palmer in 

Richards and Palmer, 19.53 

C. cracens Hoerle, 1976 

C. alveatus spiralis DockeiA' in 

MacNeil and Dockers', 1984 
C. (Lithoconus) sniitlnnllensis 

var. Dockery, 1980 
C. (Lithoconus) nocens Game, 

C. (Lithoconus) sntitlnillcnsis 

var. Dockery, 1980 in 

Campbell (1995) 

USNM 495181 (H) 
USNM 495182 (H) 
UF 108683 

UF 108858 (designated here as 
holotvpe of C. palmcrae, 
new species; formerly FL 
Geol. Sune\' L76.34) 

USNM 220109 (H> 

USNM 376678 (H) 

MGS 590 (F) 
UT-T.MM 962T\22 (H) 
UNt: 1.5448 (F) 

Claiborne, Alabama C. sauridens 

Claiborne, Alabama C. sauridens? 
South Carolina, localit\' unknown Nonwn duhiuni 

Claiborne, Alabama C. sauridens 

Wilmington, North Carolina Nomen dubium 

Jackson, Mississippi C. sauridens 

Vicksburg, Mississippi C. sauridens 

ProliabK- Claiborne, Alabama 
Claiborne. Alabama 

Jackson, Mississippi 

Claibonic, .Alabama 

Claibonie, Alabama 

Claibonie. Alabama 

Sniith\'ille, Texas 
Santee Cooper Canal, 

South Carolina 
Moseleys Fen"v, Texas 

Arroyo Veleno. Texas 

Gull Haiiiniock, Flmida 

Gulf PLuiimock, Florida 

Calhoun Count\, Flonda 
Smith C()iint\', Mississippi 

Near Newton, Mississippi 

Bastrop Countw Texas 

Near Cross, South Carolina 

C. sauridens 
Eosurcula pulcheniinus 

(a turrid) 
C. sauridens 

C. sauridens 

Nonu'ii duhiuni 

Noinrit duhiuni 

C. sinithi'illcnsis 
Nonu'ii duhiuni 

C. sauridens? 

C. h(li;jjlti 


C. pahnerae, new species 

C. sauiich'ns? 
C. sauridens 

C. sinithiillensis? 

C. sauridens? 

C. siiiithiillensis? 

Alabama, Mississippi, and Louisiana. He also correlated 
the Willistoii Formation with the upper Moodvs Branch 
Formation in Alabama, Mississippi, and Louisiana. For 
detailed correlations, see the Correlation of Stratigraphic 
Units of North America— Gulf Coast Region (1988). 

Based on lithology, Scott (1991) — in order to follow 
the North American Stratigraphic Code (North Ameri- 
can Commission on Stratigraphic Nomenclature, 
198.3) — reduced the Ocala Group to formational rank 
and returned to the terminology used by Applin and 
Applin (1944). Therefore, current designations used by 

the Florida Geological Sun-ev and United States Geo- 
logical Survey are: lower member oi the Ocala Lime- 
stone (formerly Inglis Formation) and upper member of 
the Ocala Limestone (formerly Willistoii and Ciystal 
River formations). See Figure 1 for a brief histon' oi 
stratigraphic divisions oi the Ocala Limestone in Florida. 
In outcrop and shallow subsurface, the Ocala Lime- 
stone occurs in northwestern peninsular Florida and a 
small area of the Florida panhandle adjacent to Georgia 
and Alabama. Litliologically, the Ocala Limestone is a 
relatively pure carbonate. Non-carbonate minerals 

J. R. Hendricks and R. W. Portell. 2{)()S 

Page 81 



Applin & Applin 

Vernon, 1951 











> O 


CC -^ 


£ -^ 

2 F 


fD OJ 

>^ o 















































— \ 


















S p 




Figure 1. Histon ot stratigraphic di\isioiis ol the Ocala 
Limestone in Florida (modified from Oyen and Portell, 2001). 

(quartz, chert, and cla\' minerals) represent less than 5% 
of the rock \ohune in the lower niemlier and less than 
10% in the upper member (Oyen, 1995). The lower 
member (formerly IngHs Formation) is primarily a clean 
packstone and grainstone that represents a higher en- 
ergw subtidal en\ironment while the upper member 
(formerh- Cn'stal Rixer Formation) has a litholog\' of 
mudd\- packstone and wackestone inteqireted to signify a 
lower energN' (below wave base), deeper subtidal deposit 
(Fenk, 1979). 

The Ocala Limestone contains ch\erse and abundant, 
shallow water marine in\ertebrate fossils consisting pri- 
marily of foraminitera, moUusks, and echinoids. Ta\a 
with calcific shells are preserved as body fossils, while 
taxa that had aragonitic shells hpicalh' occur as internal 
and external molds, although on rare occasions they can 
be found as pseudomorphs of calcite or silica. Most of 
the calcific shelled mollusks (e.g., oysters and scallops) 
and those found as pseudomoiphs ha\'e been well docu- 
mented (Harris, 1951; Richards and Palmer, 1953). Few 
of the man\- moldic Ocala Limestone mollusks, however, 
have recei\ed the critical attention that they need (Por- 
tell and \'okes, 1997). 


Most ot the specimens examined in this study are from 
die Florida Museum of Natural Histoiy (FLMNH) Di- 
\ision of In\'ertebrate Paleontology at the Universit)- of 
Florida (UF) and locaHty- detads for individual UF speci- 
men lots hsted below can be accessed online \ia the 
FLMXH Invertebrate Paleontology database at http:// Besides UF 
specimens, some type specimens from the Academy of 
Natural Sciences, Philadelphia (ANSP), the Texas Natu- 

ral Science Center at the Universit)' of Texas (specimens 
carry the acronym TMM lor Texas Memorial Museum), 
and the Texas Biu-eau of Economic Geolog\' (BEG; 
specimens now at the Texas Natural Science Center) 
were also examined. Other specimens referred to in the 
text are from the Geological Suivey of Alabama (GSA), 
the American Museum of Natural History (AMNH), the 
Mississippi Geological Survey (MGS), the Paleontologi- 
cal Research Institution (PRI), the Universit)' of North 
Carolina (UNO, and the United States National Mu- 
seum of Natural History (USNM). 

Morphospecies were recognized from museum speci- 
mens (primarily those at the FLMNH) using the criteria 
discussed by Smith (19.30), Rockel et al."(1995), and 
Hendricks (in press) and moiphological terminology and 
measurements collected follow those authors. Two terms 
are introduced here that relate to characters states of the 
subsutmal flexure, wliich is "the back^ward cuning or 
bending of the shell aperture below the suture of whorl 
contact" and "[b]eing a feature of the apertural margin it 
is strikingly manifest in the groxxlih hues [on the sutvu-al 
ramp] of well-presen-ed cones" (Smith, 1930: 2S4). 
These tx\'o terms are: symmetrically cuned subsutiu-al 
fle.xure and diagonal subsutural flexure. A symmetrically 
cun'ed subsutural flexure (also see the more generalized 
"curved type" described b\' Smith, 1930) has a luaximum 
point of curvature centered betxveen the margins of the 
sutural ramp. A diagonal subsutural flexure forms a 
straight or nearly-straight line that crosses diagonally 
(abaxioventrallv) across the sutural ramp. Also see 
Muiiiz-Soli's (1999) for illustrations of different subsu- 
tural flexure morphologies. 

Measurements were collected using digital and dial 
calipers and include: shell length (SL), niciximinn diam- 
eter (MD), aperture height (AH), height ot maxi- 
mum diameter (HMD), and spire angle (SA). From 
these measiu'ements, the following moiphometiic ratios 
(dexeloped by Rockel et al., 1995) were computed 
to characterize shell shape: relative diameter (RD; 
RD = MD / AH); position of maximum diameter (PMD; 
PMD = H.VID / AH); and relative spire height (RSH; 
RSH = [SL - AH]/SL). Recognized Floridian moipho- 
species were compared with pre\ioush' described species 
of Eocene Conns from the southeastern United States. 

Although the simple internal molds of Conus fossils 
offer little taxonomicalK- rele\ant information, room tem- 
perature vidcanizing (R.T.V.) silicone rubber casts made 
from external molds are often veiy useful for recognizing 
species. The extemal molds discussed herein were first 
gently cleaned to remove loose debris and then impreg- 
nated with poKvinyl butvral (Butx'ar-76), if needed, to 
consolidate the limestone. Once diied, clay clams were 
constmcted around the outside of the hmestone blocks 
containing the extemal molds. Then, de-aerated R.T.V. 
silicone nd;)ber was slowly poured into and above the 
molds and again de-aerated in a vacuum chamber. Later, 
the cured rubber casts were carefully lifted from the 
molds. For more information regarding this technique 

Page 82 

THE NAUTILUS, \'ol. 122, \o, 2 

see Chaney (1989) or \isit 


Our e.xamination of Eocene Conns fossils from Florida 
resulted in the recognition oi one distinctive moijiho- 
species that was first described by Conrad (1S33) as 
C. soniidens, and two new species (C. palmcmc and 
C. alleni) which are described below. 

Family Conidae Fleming, 1822 
Genus Conns Linnaeus, 17.58 

Conns sanridens Conrad, IS.3.3 
(Figures 2-10, 1.5^22; Table 2) 

Conns saundens Conrad, 1833: p. 33; Conrad, 1835: pi. 15, fig. 

7. For comprehensive synonvmy listings, see Palmer 

(1937), Harris and Palmer (1947), and Palmer and Brann 

?Conus claibornvnsis I. Lea, 1S.33: 186. 
Conns pai-vus H. C. Lea, 1841: 103, pi. 1, fig. 24. 
Conns toiiilis Conrad 1855: 260, pi. 15, fig. 5. 
Conns aheatns Conrad, 1865: 148. pi. 11, fig. 4. 
Conns snbsauridens Conrad, 1865: 148, pi. 11, fig. 9. 
Conns jacksonensis Meyer, 1885: 466. 
Conns deperditns var. snhdiadcma de Gregorio, 1890: 20. pL 1. 

figs. 56-58. 
?Conns {Leptoconns) sant(nidcr Gardner, 1945: 251, pi. 26, 

figs. 5, 9, 10, 14. 
?Co/H(s cracens Hoerle, 1976: 14, 16, pi. 3, figs. 1-3. 
Conns aheatns spiralis Docken' in MacNeil and Dockeiy, 

1984: 165, pi. .59, figs. 3, 4. ' 
?C())i!(.s' nocens Ganie, 1996: 90, pi. 19, figs. 9, 10. 

Diagnosis: Early postnuclear wliorls tuberculate, later 
teleoconcli whorls smooth; sliell often widest below 
slioulder; slioulder t\^Dicall\' ridge-like; sutural ramps 
with raised spiral tlireads; incised spiral groo\es at base 
of sliell. 

Description: .Shell medium to farge-sized (up to 116 
mm). Last whorl conical to broadK' and ventricosely coni- 
cal; outline tspicalh' convex near shovdder, nearly straight 
below. Shell often widest below shoulder. Shoulder typi- 
cally broadly carinate and forming pronoimced ridge, less 
often angulate; smootli. Spire of low to moderate height, 
spire angle (SA) txpically obtuse; outline tvpicallv straiglit 
to concave in smaller specimens, ustiallv sigmoidal in 
larger sliells. Lan'al shell midtispiral, witli at least three 
whorls. Earlv postnuclear whorls tuberculate. Subsutural 
fle.xure symmetricaify cui-ved, depth about 2.5x width. 
Teleoconch sutural ramps typically sigmoidal (rarely flat 
or convex), with raised spiral threads (tyjDically three to 
seven; number increases witli shell size) and intervening 
grooves; ornamentation liegins on earliest postnuclear 
whorls as a single incised spiral groove. Aperture opening 
of approximately uniform width from base to shoulder. 
Some large specimens bear a pronounced siphonal fas- 
ciolc. Last whorl with incised spiral grooves at base, 

sometimes extending weakK' to near center of whorl and 
occasionally to shoulder in small shells; spiral grooves 
obsolete in some large sliells. 

Shell Moiijhometrics: Thirteen shells of tspe and 
non-txpe specimens of C". saniidens were measured and 
moiphometric ratios \\'ere computed from these mea- 
surements (Table 2): relati\'e diameter (RD) ranges from 
0.57-0.74 (average = 0.67); position of maximum diam- 
eter (PMD) ranges from 0.83-0.95 (average = 0.91); and 
relative spire height (RSH) ranges from 0.09-0.21 (aver- 
age = 0.17). 

Type Specimens: Type specimens examined include: 
ANSP 14854, lectotvpe of Conns saniidens (Figures 2, 
3); ANSP 53813, tliree paralectot)pes of C. saundens 
(Figures 4—6); ANSP 53812, holot}pe of C. subsauridens 
(Figure 7); ANSP 13161, holot\pe of C. parvus (Figure 
8); 'and ANSP 13196, holotxpe' of C. tortdis (Figures 9, 
10). See Table 2 for measurements of hpe specimens. 

Type Localit\ and Occurrence: Conrad (1833) de- 
scribed the species from specimens collected at Clai- 
borne, Alabama. Palmer (1937: 10) designated the t\pe 
locality and stratum of Comts sauridcns as PRI station 
104: " 'Ferruginous sand' bed at Claiborne, on the Ala- 
bama River, Monroe County, Ala. Gosport sand" 
(Palmer, 1937: 10). The Cosport Sand is the uppermost 
formation of the Claiborne Group and is late middle 
Eocene (Bartonian) in age (Dockery, 1980). Tliis species 
has been previously reported from numerous Paleogene 
U.S. Gulf Coast strata, including the middle Eocene 
Claiborne Group, as C. tortdis in the upper Eocene 
Jackson Group, and as C. alvcatus in the lower Oli- 
gocene Vicksburg Group (the reader is directed to the 
following soinx'es for detailed chscussions of specific oc- 
currence records; Palmer, 1937; Harris and Palmer, 
1947; Palmer and Brann, 1966; and MacNeil and Dock- 
eiy, 1984). Conus sauridcns also occurs in the upper 
Eocene Ocala Limestone of Alachua, Suwannee, and 
Jackson counties, Florida. This species also questionabh' 
occm's as C. cracens in the lower Miocene Chipola For- 
mation of nortliern Florida. 

Other Material Examined: In addition to tlie tvpe 
specimen lots listed abo\e, 28 specimen lots containing 
C. sauridcns were examined (o\er 80 specimens). These 
include: UF 283, UF 290, UF 2292, UF 8647, UF 14706, 
UF 16726, UF 16661, UF 18874, UF 57713, UF 57733, 
UF 100693, UF 101995, UF 114376, UF 115875, UF 
119912. UF 119913, UF 119918, UF 119962, UF 
120021-UF 120027, UF 120032, UF 122384, and UF 

Discussion: Conus sauridcns was tlie third fossil Co- 
nus species to be described from North America, and the 
first from the Paleogene (Green described the Neogene 
species C. dclnrianns ;iihI C. nuirijUindicus in 1830; see 
Kohn, 1992). As such, this species has received much 
attention in the literature, particularly with regards to its 

J. R. Hendricks and R. W. Portell, 200S 

Page 83 

Figures 2-lS. Specniieus of Conns saiihdens Conrad, 1833 (2-10, 15-1<S), C. cract'iis Hoerle, 1976 (11, 12j, and C. lujccns 
Gai"\ie. 1996 (13, 14). See text and Table 1 for locality information. All scale bars equal 1 cm. Scale bar above Figure 5 pertains to 
Figures 2-T and 11-14. Scale bar below Figure 8 pertains only to that figure. Scale bar between Figures 9 and 10 pertains to Figures 
9-10 and 15-18. 2-3. Lectotype (ANSP 14854) of C. sauridens. shell length 33.4 mm, maximum diameter 21.1 mm. 4. Faralectotype 
(ANSP .53813-3) of C. sauridens. shell length 31.9 mm. 5. Paralectotj/pe (ANSP 53813-1) of C. sauridens, maximum diameter 22.3 
mm. 6. Paralec■tot^.pe (ANSP 53813-2) of C. sauridens, shell length 30.2 mm. 7. Holotype (ANSP 53812) of C. subsauridens Conrad, 
1865, shell length' 33.4 mm. 8. Holotype (ANSP 13161) of C. parvus Lea, 1841, shell length 6 mm. 9-10. Holot>pe (ANSP 13196) 
of C. torfilis Conrad, 18.55. shell length 90.0 mm, maximum diameter 53.0 mm. 11-12. Paratope (UF 119560-1) of C. cracens Hoerle. 
1976, shell length 32.2 mm. maximum diameter 17.8 mm. 13-14. Holotype (TM.\I-962T.\22) of C. noccns Ganie, 1996, shell length 

19.2 mm, maximum diameter 8.' 

15. R.T.V. siUcone rubber cast fn 

vtemal mold of C. sauridens (UF 120026). 16. R.T.N'. 

sihcone rubber cast from external mold of C. sauridens (UF 122384). 17-18. Highly leached, heavily pitted, and slightly silicilied 
shell of C. satiridens (UF 120027), shell lengdi 27.4 mm. maxiiriuni diameter 16. .5 mm. 

Paee 84 

THE NAUTILUS, Vol. 122, No. 2 

Table 2. Measurements (in mm) and morphometric ratios of t)'pe and non-ty|ie specimens examined. Some measurements could 
not be collected from casts made from molds; other measurements (and associated ratios) are not accurate because of shell damage 
(indicated by an asterisk, °). Morphological abbreviations: SL, shell length; MD, maximum diameter; AH, aperture height; HMD, 
height of maximum diameter; SA, spire angle (in degrees); RD, relative diameter; PMD, position of maximum diameter; and RSH, 
relative spire height. Specimen abbrexiations: H, holotvpe; L, lectot\pe; P, parat\pe; and PL, paralectotvpe. 










T\pe Specimens o{ Cunus sauridens Conrad, 1833 

ANSP 14854 (L, C. sauridens. Figures 2, 3) 









ANSP 53813-1 (PL, C. sauridens. Figure 5) 









ANSP 53813-2 (PL, C. sauridens. Figure 6) 









ANSP 53813-3 (PL, C. sauridens. Figure 4) 









ANSP 53812 (H, C. suhsauridens. Figure 7) 









ANSP 13161 (H, C. pamis. Figure 8) 









ANSP 13196 (H, C. tortihs. Figures 9, 10) 









Non-T)pe Specimens of Conus sauridens Conrad, 1833 

UF 283 









UF 290-1 









UF 8511-1 









UF 8647 



33. 1 






UF 16726 









UF 115875 









UF 120022 









Conus pahnerac new species 

UF 108858 (H, C. pahncnic, new .species. Figures 23, 24) 

UF 15886 (F) 

UF 18599 (F, Figure 25) 

UF 18711 (F) 

UF 18719 (F) 

UF 18737 (F, Figure 26) 

UF 57018 (F) 

UF 66738 (F, Figure 27) 

UF 68306 (P, Figure 28) 

UF 74473 (F, Figures 29, 30) 

UF 110360 (P) 

UF 111327 (F) 

UF 112981 (F) 

Conus alien! new species 

UF 119920 (H, Figures 31, 32) 
UF 119919 (P, Figure ,34) 
UF 119976 (P, Figure 35) 
UF 119977 (P, Figure 33) 

Other Tvpe Specimens 

ANSP 16145 (H, C. connacki. Figure 39) 

UF 119560-1 (P, C. eracens, Figiires 11, 12) 

UF 76798-1 (P, C. craceus) 

TMM-BEG 35656 (H, C. smitlioillcnsis. Figures 36-38) 

TMM-962TX22 (H, C. nocens. Figures 13,'l4) 

T.\I.\I-962TX23 (P, C norms) 






11.5° 14.6° 13.7° 77 

22.3° — — 103 

21.5° _ _ _ 

14.2° — — 101 

19.3 — — 98 

19.0° — — 98 

28.4° 53.8° — — 

11.3 — — 108 

12.4 — — 108 










0.66° 0.91 



















morphological variation ajid piobal.ilc syiioiivin.s. MllcIi 
ot the following wa.s derived from discussions in Palmer 
(1937), Harris and Palmer (1947), and Paliiu-r and Rrann 

Timothy Conrad (1S33) described C. sauridens from 
shells that he collected at Claiborne, Alabama. According 
to Palmer (1937: 461), the "Conradian collection of 
sauridens consists of 5 specimens" which were appar- 
ently glued to one card, cataloged as ANSP 14854. 
Palmer (1937: 461) goes on to state dial die "tvpe of 

Conns snl>s(niri(lcns Con. [see below] is also on the card 
with the Ciuins sauridens collection." These two passages 
suggest tliat Conrad's Claiborne collection originally con- 
sisted of six specimens. The collection now consists of 
five specimens: the lectotype (ANSP 14854; Figures 2, 3) 
of C sauridens, three paralectotypes (ANSP 53813; Fig- 
ures 4-6), and the liolot\pe of C. suhsauridens (ANSP 
53812; Figure 7); the location of any possible sixth .speci- 
men is not known. One low-spired specimen in tlie f\pe 
series (ANSP 14S54; Figure 2) closely resembles Con- 

J. R. Heiulricks and R. W. Portell. 2008 

Page 85 

rad's (1835, pi. 15, fig. 7) original figure off', Miiiridcus. 
Palmer (1937) alluded to this specimen in lier text as 
matching Conrad's figure, but did not formall)' designate 
it as the lectotN-j^e. Kohn (1992) — following Moore 
(1962) and Palmer and Brann (1966) — considered this 
specimen to be the lcctot\pc and we accept his concln- 

Palmer (1937) argued tliat the matrix lilling the aper- 
tures oi the shells in Com'ad's Conii.s .sauridens series 
suggests that the shells do not hkeh' share the same geo- 
logical provenance. The matrb; filling the lectot)pe is 
orange and appears to be sand from the Cosport For- 
mation (Palmer, 1937). The matrix filling the tin-ee para- 
lectotxpes is light-gray. The matrix filling the holotvpe of 
C. subsauridens is orange, resemliling that of the lecto- 
txpe of C. saiiiidcns (Palmer [1937: 461] described tlu' 
matrix filling the hoIot\pe of C. subsauridens as "a retl, 
silicified matrix resembling that of the Orangeburg" 
material from South (Carolina). It is important to note, 
howe\er, that these differences in matrix color ma\' be 
due to diagenetie weathering of the original matrix ma- 

The lectot\pe (ANSP 14854; Figures 2, 3) of C. sau- 
ridens differs from tlie paralectotypes (ANSP 53813; Fig- 
ures 4-6) — which, with the holotxpe of C. subsauridens 
(see below), are of the much more common moiphol- 
og\' — in the following respects: it has flat to slightly con- 
xex sutural ramps with moi"e raised spiral threads (about 
6) than is txpical, the shoulder is angulate rather than 
forming a carinate ridge, the widest point of the last 
whorl is nearh' at the shoulder lather than beneath it. 
and the spire is lower than in most other specimens of 
the species (see Table 2); further, the lectotxpe does not 
pro\ide definite e\idence oi tubercles on the earlv post- 
nuclear whorls, though these are highlv eroded on this 

Isaac Lea (1833) described Conus ehnbimiensis from 
materials sent to him b)' Judge Charles Tait of Claiborne, 
.Alabama, but lost his onlv specimen before it could be 
figured (Kohn, 1992). His description suggests that his 
specimen was a C. sauridens. a name that has priority- b\' 
three months (Kohn, 1992). Harris (1895), Palmer 
(1937), and Palmer and Brann (1966) SMionvmizecI C. 
claibornensis with C. sauridens. while de Gregorio 
(1890), Dall (1896), and Kohn (1992) regarded this taxon 
as a nomcn dubium. Gi\en that no available evidence 
suggests that more than one fossil Conus species is 
present at Claiborne, Alabama, we agree with the former 
authors that C. claibornensis is equivalent, while rjues- 
tionabh'. to C. sauridens. For a historical o\'eniew' of 
interactions between I. Lea. T. .\. Com'ad, and C. Tait, 
see Wheeler (1935). 

In 1841. H. C. Lea (I. Lea's father) described Coinis 
paiTus on the basis of a single, small, damaged shell 
lAXSP 13161; Figure 8) from the Gosport sand at Clai- 
borne, .Alabama. \A'e agree with Dall (1896), Palmer 
(1937), and Palmer and Brann (1966) that C. parvus is a 
juvenile C. sauridens. Features uniting the holotype of C. 

parvus with C. sauridens include: tuberculate early post- 
nuclear whorls, raised spiral threads on the sutural 
ramps, and incised spiral grooves on the anterior half of 
the last whorl. 

Comud (1855) described Conus loriilis Ironi one large 
specimen (ANSP 13196; Figures 9, 10) from Jackson, 
Mississippi and differentiated it from C sauridens by its 
"more prominent and con\ex spire, in the large twisted 
callus at base, & c." (p. 260). Dall (1896) and Palmer 
(1937) both considered C. toiiilis s\aionymous with C. 
sauridens and Palmer (1937) described C. toriilis as rep- 
resenting "the maximum growth of the species" (p. 459). 
We agree with these a\ithors that specimens of C. toriilis 
are large C sauridens. 

Conrad (1865) described two additional species of 
Eocene Conus: C. subsauridens and C. aheatus. Conrad 
statetl that C. subsauridens was from "the Burrstone, 
probabK', oi Alabama": we assume (see above) that the 
holotvpe (ANSP 53812) is from Clairborne. We in- 
spected the type of C. subsauridens (Figure 7) and agree 
with Dall (1896) and Palmer (1937) that it is a junior 
synonym of C. sauridens. 

The t)pe locality for Conus alveafus is \'icksburg, Mis- 
sissippi; MacNeil and Dockeiy (1984) suggested that the 
lectotvpe (ANSP 13446; MacNeil and Docken-, 1984. pi. 
38, fig. 26) and paratvpe (ANSP 13494) are probably 
from the Bvram Formation. Conrad (1865: 148) differ- 
entiated C. alceafus from C'. sauridens bv its "less el- 
evated and . . . more profoundlv carinated spire, and the 
revolving lines on the spire are less numerous than in the 
former [C. sauridens]." Dall (1896) and Palmer (1937) 
i-eeognized Conus toriilis and C. aheatus. respectively, as 
the Jacksonian (upper Eocene) and Vicksburgian 
(lower Oligocene) forms of the older, Clairbornian 
(middle Eocene), C. sauridens. MacNeil and Docken' 
(1984), however, continued to recognize C. akeatus as a 
distinct Oligocene species occurring in Mississippi and 
Mexico; thev chd not compare C. aheatus (or Dockeiy's 
subspecies C. alveatus spiralis; in MacNeil and Dockeiy, 
19S4) with C. sauridens. MacNeil and Dockeiy's (1984) 
figures of C. aleveatus (including the lectotype; pi. 38, 
fig. 26) appear consistent with C. sauridens as circum- 
scribed here. 

Meyer (1885) described — but did not figure — Conus 
jacksonensis from Jackson, Mississippi, and described the 
species as similar to Conus protracta Mever, 1885 (an 
Oligocene taxon from Vicksburg and Red Bluff, Missis- 
sippi that we accept, but do not consider fuither here; 
see MacNeil and Dockeiy, 1984 for details), but "with 
revolving lines on the .spire" (p. 466). Meyer [1886] pre- 
sented C. protracta as C. protraetns and the latter 
.spelling is the one most commonlv seen in the literature. 
Harris and Palmer (1947: pi. 62, fig. 17) figured a speci- 
men (unnumbered, but said to be from the collections of 
the Geologv Department of Johns H(jpkins Universitv) 
that they regarded as the holotype of C. jacksonensis. 
This specimen "consists of the apical whorls" and is 3.5 
mm in size (Harris and Palmer, 1947: 446). Hariis and 

Page 86 

THE NAUTILUS, Vol. 122, No. 2 

Palmer (1947) considered diis taxon a junior synonym of 
C. sauridens and we agree, particularly because of die 
presence of tuberculate early postnuclear whorls and die 
presence of raised spiral threads on the sutural ramp. 

De Gregorio (1890) described Conus dcperditns var. 
suhdiadema from Claiborne, Alabama. Palmer and 
Brann (1966) considered this subspecies equivalent to C. 
sauridens and, based upon our inspection of de Grego- 
rio's (1890) figures, we agree. We have not, however, 
viewed the holotype of C. deperditus var. suhdiadema. 
vvhich Palmer and Brann reported as PRI 26436. 

Gardner (1945: 252) described Conns santander as 
including "those species from the western Gulf that have 
formerly been included under Conns sauridens Conrad, 
described from Claiborne." Gardner's figures of the ho- 
lotype (USNM 495181; Moselevs Ferry, Burleson 
County, Te.xas) of C. santander appear consistent with C. 
sauridens and we consider this taxon a probable svii- 

Hoerle (1976) described Comis cracens (see paratype 
in UF 119560, Figures 11, 12) from the lower Miocene 
Chipola Formation of northern Florida and noted its 
strong similarity to C sauridens: "C. cracens appears to 
be a descendant of the widespread (Alabama, Vlissis- 
sippi, Texas) middle Eocene to Oligocene species, C. 
sauridens Conrad" (p. 16). She differentiated C. cracens 
from C. sauridens on the basis of several characters: 
the "nodes on the spire whorls persist for a greater num- 
ber of turns on C. cracens, also it is larger, more slender, 
with stronger and more opisthocyrt growth lines and 
more pronounced basal ornament" (p. 16). We examined 
Hoerle's paratypes at the FLMNH and could not find 
any discrete niorpliological characters separating the txvo 
taxa. Given the vast amount of geological time separating 
the species (over 12 million vears), however, we ques- 
tionably synonymize C. cracens witli C, s(niiidens. Re- 
solving the relationship bet^veen these two taxa will re- 
quire additional study. 

Finally, Game (1996) described Conns nocens on the 
basis of two small specimens from the Reklaw Foimation 
of Bastrop Count)', Texas: TMM-962TX22 (holot>iie; er- 
roneouslv published as UT-TMM 84822; Figures 13, 14) 
and TiVlM-962TX23 {pai"at)'[De, erroneously published as 
UT-TMM 84823). In addition to these two specimens, 
GaiAie (1996) reported that he examined 25 additional 
specimens of C. nocens from the Weches Formation and 
over 100 specimens from the Cook Mountain Formation. 
While we did not observe these additional, stratigraphi- 
cally younger specimens, the holotype and paratvpe of 
C. nocens appear cojisistent with Conns sauridens. They 
each have raised spiral threads on the sutiiral ramp, sym- 
metrically curved subsutural flexures, and incised spiral 
grooves on the anterior half of the last whorl. Further, 
both have last whorl sliapes similar to C. sauridens. The 
early postnuclear whorls ol the holotype are tuberculate, 
but most appear smooth on the paratype. Garvie (1996: 
90) stated that four characters separate C. nocens from 
C. scniridens: "the Hat sides, the sharp uniidgcd caiina. 

the coarse, strong, spiral basal lines, and the lack of 
strong growth lines on the ramp." We do not consider 
these characteristics — in isolation or combination — 
sufficient to discriminate C. nocens from C. sauridens (as 
circumscribed above), especially because TMM-962TX22 
and TMM-962TX23 are both shells of juveniles. Since, 
however, we have not seen the other specimens of C. 
nocens mentioned by Game (1996), we consider our 
synonymy of C. nocens with C sauridens tentative. Re- 
gardless of their taxonomic identit\', TMM-962TX22 and 
TMM-962TX23 are important specimens because of 
their likelv stratigraphic position in the Reklaw Forma- 
tion. The Reklaw Formation is thought to span the early- 
middle Eocene (or, Ypresian-Lutetian) boundaiy and has 
a relative age equivalent to nannoplankton biochrono- 
zone NP14 (Zachos antl Molineux, 2003: fig. 2), which 
has an absolute age of about 49.7 to 47.3 Ma (Berggren 
and Pearson, 2005). This age would make these two 
specimens the oldest known Conns fossils from the 
United States Coastal Plain, and only slightly younger 
than the oldest Conns in general (Ypresian of England 
and France; Kohn, 1990). This oldest regional occur- 
rence record is tempered, however, by the fact that the 
position of the original collection locality of TMM- 
962TX22 and TMM-962TX23 ("Devil's Eye, Colorado 
R."; station 11 of the Geological Sun'ey of Te.xas and 
localits' ll-T-36 of the Texas Bureau of Economic Geol- 
ogy) is uncertain (Zachos et al., 2005) and may no longer 
e.xist (Game, 1996). Further, no additional specimens of 
Conns from the Reklaw Formation have yet been found 
in the collections of the Texas Natural Science Center (A. 
Molineux, personal communication to JRH, July 11, 

Most of our knowledge of Eocene C. sauridens in 
Florida is from R.T.\'. silicone iTibber casts of external 
molds in limestone (e.g., UF 120026, Figure 15; UF 
122384, Figure 16), though UF 120027 "(Figures 17, 
18) — which is a highly leached, heavily pitted, and 
slightly silicified shell — is an exception. Conus sauridens 
is present in upper Eocene Ocala Limestone of Jackson 
(UF 18874, UF localitv-- JA002; UF 120026, UFbcalitv 
JA018; UF 119912, UF localitv JA027; and UF 119918, 
UF locahty JA031), Alacluia (UF 120027, UF localitv 
ALOOl), and Suwannee (UF 122384 and UF 120032, 
both UF localit)' SU0()3) counties. Finally, one additional 
specimen (UF 119913) that may be C. sauridens is from 
the lower Oligocene Biunpnose Limestone of fackson 
County (UF locality' JA025). Besides these records, the 
only other known record of this taxon (as recognized 
here) in Florida is Dall's (1916: 4489) account'of C. 
tortilis in the "Ocala." Conus sauridens co-occms in the 
Eocene of Florida with the new species C. palnierae and 
C. allcni; characteristics that distinguish the new species 
from C sauridens are discussed below. 

A complete review of the fossil record of Conns sau- 
ridens is beyond the sci>pe of this paper, though the 
preliminary observ'ations we have made here support 
earlier demonstrations (Palmier. 1937) that C. sauridens 

J. R. Hendricks and R. W. Portell, 200S 

Page 87 

\\'as a moq:)liologicall\- \arial)le, geographicalK wide- 
spread, and teniporalK' persistent species. While the old- 
est known (earK- Eocene) Coitus fossils were small {<35 
mm in shell length), larger species (ca. 70 mm in shell 
length) ha\e pre\iousl\- been reported from the middle 
Eocene (Kohn, 1990). During this studv, we recognized 
a ven- large (shell length, 116.1 mm) specimen (UF 
120022; Eigure 19) of C. saiiridcns from the upper 
Eocene Mood\s Branch Eormation of Grant Parish, 
Louisiana (UF locality- ZL004). This specimen may be 
the largest Conns \et known from the Eocene of the U.S. 
Coastal Plain. The large geogi-aphic range of C. smiiiclcns 
could be related to its de\elopmental mode, which was 
likeh' planktotrophic based upon its multispiral lanal 
shell (Figures 20-22), though testing this hypothesis 
widiin the context of Shuto's ( 1974) model of the rela- 
tionship between dexelopmental mode and lanal shell 
morpholog\' (;dso see Kohn and Perron, 1994) wall re- 
quire additional stud\'. While not necessaiily useful as a 
guide fossil, shells of C. saiiridcns (as circumscribed 
here) ha\e recend\" been utilized in several isotopic stud- 
ies (Kobashi et al., 2001; Kobashi and Grossman, 2003; 
Kobashi et al., 2004) of Paleogene climate and have 
pro\"en to be geologically useful in this regard. 




crae new s 

(Figures 23-30. Table 2) 


Conns sp 
fia. 14. 

B, Pah: 


and Palmer, 1953: 40, pi. 

Diagnosis: Teleoconch whorls stepped; early post- 
nuclear whorls smooth; sutural ramps tvpicallv smooth; 
last whorl smoodi. 

Description: Shell small to moderately large-sized (up 
to about 65 mm in length). Last whorl conical; outline 
slightlv concave. Shoulder shai-plv angulate, smooth. 
Spire of moderate height; outline concave to straight. 
Teleoconch whorls stepped; spire angle of earl\- whorls 
t\picall\- obtuse relative to later whorls. Lanal shell im- 
knowTi. Early postnuclear whorls smooth. Subsutural 
flexure SMiimetricalK' cuned. Teleoconch sutural ramps 
concav'e and t\picall\- smooth, though occasionalK^ 2 or 3 
weak spiral threads are present. Aperture moiphologv 
unknowTi. Last whorl t\picall\- smooth, though fine spiral 
hues ma\' co\ er the last whorl of some small specimens. 

Type Specimens: Ilolot\pe UF 108858 (Figures 23. 
24), a specimen originallv described as "Conns sp. B" by 
Palmer in Richards and Palmer (1953: 40, pi. 2, fig. 14). 
The holotype is presened as a calcite-replaced shell. All 
paratopes are moldic (consisting of just e.xtenial or exter- 
nal and internal molds) and include: UF 15886, UF 
18599 (Figure 25), UF 18711, UF 18719, UF 18737 
(Figure 26), UF 57018, UF 66738 (Figure 27), UF 68306 
(Figure 28), UF 74473 (Figures 29, 30), UF 110360, UF 
111327, and UF 112981. See Table 2 for measurements 
of diese specimens. 

Figures 19-22. Specimens of Conns sinniclciis Conrad, 
1833. 19. Largest knowm specimen (UF 120022) of C. san- 
ridens, shell length 116.1 mm, Moodys Branch Formation, UF 
locality ZL004 (Montgomery Landing 01), Grant Parish. Loui- 
siana; scale bar equals 1 cm. 20. Juvenile shell (UF 126927), 
shell length 3.4 mm, Moodys Branch Formation, UF locality 
ZL004 (Montgomeiy Landing 01), Grant Parish. Louisiana; 
scale bar equals 1 mm. 21-22. Shell (GSA.2007.005). shell 
length 60.5 mm. Mood\s Branch Formation, .Vlontgomeiy 
Landing, Grant Parish, Louisiana; Figure 21 shows the lan'al 
shell and early postnuclear whorls of diis specimen, which is 
also shown in Figure 22 (both scale bars equal 1 cm). 

Page SS 

THE NAUTILUS, \'ol. 122, No. 2 

Figures 2.3-30. Specimens of Co;/;;.? ptihncrac new species. All scale bars equal 1 cm. 2.3-24. Holotvpe (UF 108858), preserved 
shell length 22.0 nnn. preseived maximum diameter 11. .5 mm, lower member of the Ocala Limestone (formerly IngHs Formation), 
UF locaht)- LV014 (Gulf Hammock 02), Levy Count}', Florida. 25. R.T.V. silicone aibber cast of parat\pe (LIF 18599), preser\'ed 
shell length .50.1 mm, presen-'ed maximum diameter 23.4 mm, Ocala Limestone, UF localit\' AL016 (S.M. Wall Quarr\- 01), Alachua 
Countv', Florida. 26. R.T.\'. siUcone nabber cast of paratxpe (UF 18737), presened shell length 56.1 mm, presen-ed maximum 
diameter 24.1 mm, Ocala Limestone, UF locality LF0(31 (Dell Limerock Mine), Lafayette County, Florida. 27. R.T.V. silicone mbber 

cast of paratvpe, (LIF 66738), maximum diameter 14.! 

Ocala Limestone, UF locality AL028 (Newbeny 03), .Alachua Count)', 

Florida. 28. R.T.\'. silicone nibber cast of paratope (UF 68306), maximum diameter 19.3 mm, Ocala Limestone, UF locality- AL004 
(Dickerson Limerock Mines), Alachua County, Florida. 29-30. R.T.V. silicone rubber cast of paratvpe (UF 74473), preseived 
maximum diameter 19.0 mm, Ocala Limestone, UF localit^• AL()04 (Dickerson Limerock Mines), .Machua Counts', Florida; the 
topmost portion (indicated b\ arrow) of Figure 29 is magnified 3.5x in relation to Figure .30 to show details of the sutural ramps. 

Type Locality' and OccuiTence: The liolot\pe (UF 
1088.58; Figure.s 23, 24) Is from the lower member of the 
Ocala Limestone (formerh- the Inglis Formation) at UF 
locality L\'014, Gulf Hammock 02, Levy Countv, 
Florida. Richards and Palmer (1953; 5) described the 
locality (R. O. Vernon's L-93) as a "road metal pit 2.9 
miles south of the north limits of the town of Gulf Ham- 
mock just southwest ot State Road 55 in the southwest 
quarter oi Section 34, Ttjwnship 14 South, Range 16 
East." The paratvpe specimens are from Alachua Coimty 
(UF 68.306, UF 74473, UF 111.327, and UF 112981, UF 
locahh' AL004, Dickerson Limerock Mines, Ocala Lime- 
stone;' UF 18599, UF locality AL016, S.M. Wall Quariy 
01, Ocala Limestone; UF 15886, UF localit)' AL017, 
Newberry Coiporation Pit 01, Ocala Limestone; and UF 
66738, UF locality- AL028, Newbeny 03, Ocala Lime- 
stone) and Laf^xyette County (UF 18711, UF 18719, UF 

18737, UF 57018, and IF 110360, UF locality LFOOl, 
Dell Limerock Mine, Ocala Limestone). Thus, all speci- 
mens of C. palmerac are from the upper Eocene, Jaek- 
sonian Ocala Limestone of Florida. 

Etymology: This species is named in honor of 
Ka'therine V. \V. Palmer (189.5-1982), second director of 
the Paleontological Research Institution (Ithaca, NY), 
who was the first to recognize this form as a new, unde- 
scribed species and for her important contributions to 
Cenozoic paleontology (see Caster, 1983). 

Diseu.ssion: Palmer (1953) did not describe UF 
108858 as a new taxon because she did not consider this 
singk' tlaniaged shell adequate for this puipose. Newly 
collected specimens (all molds) are consistent witii the 
gross moiphology oi Palmers fossil, but offer new mor- 

R. Hendricks and R. W. Portell, 20()S 

Page 89 

Figures 31-39. Specimens ul Coiius altciii new species (31-35;, C. siitithvillcnsis Harris, JSUo (36-3S), and C. cDnnacki Har- 
bison, 1944 (39). Scale bar equals 1 cm. 31-32. Holohpe (UF" 119920) ol C. nlleni, shell lengtli .36.2 mm, preseived maximum 
diameter 20.7 mm, Moodys Branch Formation, UF locality' ZL004 ( Montgomeiv Landing), Grant Parish, Louisiana. 33. Parahpe 
(UF 119977) of C. alleni, preserved shell length 35.1 mm, Ocala Limestone, UF loealit)' LF002 (MiU Creek Quany), Lalayette County, 
Florida. 34. Faratvpe (UF 119919) of C alleni, presened shell lengtlr 33,2 mm, Ocala Limestone, UF loealit)' SU014 (Suwannee American 
Cement), Suwannee County, Florida. 35. Paratype (UF 119976) of C, alleni. preseived shell lengdi 34.5 nun, UF locality LF002 (MiU 
Creek Quarrs'), Lafayette Count}-, Florida. 36-38. Holotype (TMM-BEG 35656) of C. smithvillensis, shell lengdi 42.1 mm, pre- 
served maximum diameter 14.8 mm, Weches Formation, Colorado River at Smithville, Bastrop County, Texas. 39. Holot)pe (ANSP 
16145) of C. connacki (not a Conus; see text), preserved shell length 28.7 mm, Santee Cooper Canal, South Carolina. 

phological details that now justify description td this 

Conns pahnerae co-occurs in the Ocala Limestone 
witli two other late Eocene Conns in Florida: C. sau- 

ridcns Conrad, 1833, and C alleni new species. Conns 
pahnerae may be distinguished from botli of diese species 
by its stepped teleoconch whorls, smooth earlv postnuclear 
whorls, smooth sutural ramps, and smooth last whorl. 

Page 90 

THE NAUTILUS, Vol. 122, No. 2 

Conns alleni new species 
(Figures 31-35, Table 2) 

Diagnosis: Shoulder undulate; subsutural flexure di- 
agonal; last whorl witli raised spiral cords on anterior 

Description: Shell medium-sized (up to about 36 nnn 
in length). Last whorl conical; outline straight to slightly 
sigmoidal (convex near shoulder). Shell widest at shoul- 
der. Shoulder angulate and with large tubercles resulting 
from weak undulations. Spire ot moderate height; outline 
straight to slightly concave. Lan'al shell unknov^oi. Early 
postnuclear whorls tuberculate. Subsutural flexure di- 
agonal, depth about 1.5x width. Teleoconch sutural 
ramps concave with several raised spiral cords. Aperture 
opening about as wide at base as at shoulder. Last whorl 
with pronoimced raised spiral cords on anterior hall, 
sometimes extending weakly as threads to shoulder. 

Type Series: Holotyi^e, UF 119920 (Figures 31, 32). 
The three paratypes consist of external molds and in- 
clude UF 119977 (Figure 33), UF 119919 (Figure 34), 
and UF 119976 (Figure 35). See Table 2 for measure- 
ments of these specimens. 

TApe Locality and Occurrence: The holotspe (UF 
li9920. Figures 31, 32) was collected by J. E. Allen from 
the Jacksonian Moodys Branch Formation at Montgom- 
ery Landing (UF localitv ZL004), Grant Parish, Louisi- 
ana. The paratv-pes are all ironi the Ocala Limestone of 
Florida, including two specimens from Lafayette Count)' 
(UF 119976, Uf1i9977. UF localit)' LF002, Mill Creek 
Quarry) and one specimen from Suwannee County (UF 
119919, UF locality SU014, Suwannee American Ce- 

Etymology: Tliis species is named in honor ol James 
E. Allen (1914-1997) of Alexandria, Louisiana, who was 
an enthusiastic collector and scholar of Gulf Coast 
Eocene moUusks. 

Discussion: Conns alleni co-occurs in the Eocene ot 
Louisiana with C. saiuidens and in the Ocala Limestone 
oi Florida with C. palmerae new species and C. sau- 
lidens. Conns alleni can be readily diiterentiated from 
both species by its undulate shoulder and spiral cords on 
the anterior lialf of the last whorl (raised spiral threads on 
the last whorl niav also be present on small shells of C. 
palmerae, but if so are much weaker). 

Conns alleni shares some resemblance with a moldic 
Oligocene fossil (USNM 166720) from Decatur County, 
Georgia that Dall (1916) described as C, oanghani. Dall's 
(1916: pi. 86, fig. 1) figure of the cast shows a specimen 
(partially obscru'ed bv matrix) with an obtuse spire angle, 
undulate shoulder, and raised spiral threads on the su- 
tural ramps that are similar to the teleoconch moqihol- 
ogy of C. alleni. The presence of rows of spiral beads on 
the last whorl, the fact that the shell is widest below the 
shoulder (rather than at the shoulder, as in C. alleni), and 
the lact that the anterior end ot the shell appears com- 

pletely obscured by matrix prevents us, however, from 
considering these two forms equivalent. 

The onK' known shell material of Conns alleni is the 
holotype (from Grant Parish, Louisiana); the other three 
specimens are from Florida and all consist of external 
molds. This taxon was apparently rare, especiallv outside 
ol Florida. We recognized this new form from the moldic 
Floridian material before we — by chance — discovered 
the similar shell from Louisiana in the FLMNH collec- 
tions. We chose to designate the shell as the holotype, 
lather than one of the parahpe external molds, because 
of its greater number of characters available for obser- 

Other Records of Eocene Convs from the U.S. 
Coastal Plain 

Unidentifiable internal molds of Conns are connnon in 
the Eocene Ocala Limestone of northern Florida. We 
examined 54 such lots (over 440 specimens) from Ala- 
chua, Citnas, Jackson, Lafayette, Marion, and Suwannee 
counties. These include: UF 15884, UF 15892, UF 
15905, UF 17831, UF 17832, UF 17879, UF 17947, UF 
17950, UF 17967, UF 1S423, UF 18759, UF 18848, UF 
18864, UF 18896, UF 18955, UF 18962, UF 19140, UF 
19174, UF 19204, UF 19215, UF 20744-20746, UF 
46435, UF 68270, UF 107265, UF 119900-UF 119904, 
UF 119906-119911, UF 119914-119917, UF 119921, 
UF 120027, UF 120032-120040, UF 120047, and UF 

During the coui'se of this work, we became aware of 
several other Eocene Coastal Plain Conns species that 
are likely distinct, but are not kiiowai to occur in Florida 
and will require additional investigation; these include: 
Conns smithvillensis Harris, 1895; C. smithvillensis var. 
Docken', 1980 (also see C. smithvillensis var. Dockeiy in 
Campbell, 1995); and Conns haigljti Gardner, 1945. 

Harris (1895) described C. smitluiillensis from the 
Colorado River at Smithville, Bastrop Count\', Texas 
fWeches Formation according to TMM records). The 
holotype (TMM-BEG 35656; Figures 36-38) shares 
some characteristics with C. sauridens (including a mul- 
tispiral protoconch, tuberculate early postnuclear whorls, 
spiral threads on the sutural ramp, growth lines showing 
a deep antl symmetrically cruA'cd subsutural flexiu'e, and 
incised spiral grooves near the base of the last whorl), but 
has a \eiy different overall shell shape: the spire is much 
higher (spire angle: 53°, Table 2; in mature individuals of 
C. sanridens, the spire angle is typically over 100°), the 
conical last whorl has straight sides, and the sutural 
ramps are flat. Dockeiy (1980) figured a shell (MGS 590) 
from the slightly vounger Cook Mountain Formation of 
Mississippi that he described as a variety of C. smithvil- 
lensis. His ligmi'il shell appears to bear many of the 
discrete characteristics of C. sniitlnillensis described 
above, though has a lower spire (ca. 76°) and the last 
whorl is sigmoidal in profile. Conns smithvillensis is not 
at all simikir in form to C palmerae. While it bears some 
ot the slii'll characteristics ol C. alleni, it lacks C. alleni'^ 

J. R. Hendricks and R. \A'. Portell, 20()S 

Page 91 

distincthe raised spiral cords at tlic base of the hist w liorl. 
Campbell (1995) attributed an external mold (UNC 
1544S"> from the Santee Limestone near Cross, South 
Carolina to DockeiTS (1980) ^•ariet^• of C. smitliviUensis. 
Campbell (1995: 146) stated that this form is the "most 
common Conns in the Santee Limestone" and that it "has 
a nodose shoulder and a taller, more tabulate spire than 
the widespread Coiuis (Litlioconii.s) saiiridens." Further 
stud\" will be required to determine whether these vari- 
eties are indeed consistent with Harris's tirxon. 

Gaixlner (1945) described C. luiighti from the Laredo 
Formation of Zapata Counts', Texas. She did not fitfui-e 
her holot\pe (USNM 4951S2), but her figiu'ed (pi. 26, 
fig. 7) paratope specimen (USNM 4951S3) has a convex 
spire profile and rounded shoulder that appear chstinct 
from those characters in C. sauridcns, C. palmerae, and 
C. oUeni. Gardner's otlier figiu'ed (pi. 26, fig. 2) specimen 
of C. liaigliti lacks this distincti\"e spire form. 


The following species of Conus described from the 
Eocene of the U. S. Coastal Plain should be regarded as 
nomina clubia: Conns gijratus Morton, 1834; Conns iiui- 
tilatiis Tuome)', 1852; Conns improvidns de Gregorio, 
1890; and Conns connacki Harbison, 1944. Conus gi/ra- 
tns (holotspe, AXSP 211) is an internal mold of a shell of 
uncertain pro\'enance from South Carolina (also see 
Kohn, 1992); Campbell (1995) suggested that it could be 
an internal mold of the puiported variety of C. smithiil- 
Icnsis presented hv Docken- (1980). Conus uudtilatns 
was described (but not figm-ed) bv Tuomey from casts 
found near Wilmington, North Carolina. De Gregoiio 
(1890) described C. improvidns from Claiborne, Ala- 
bama, but his holotvpe is reportedly lost (Palmer and 
Brann, 1966) and Palmer (1937: 465) considered the 
taxon "of doubtful status as an American species." Conns 
connacki Harbison, 1944 was described from the Eocene 
Santee FormatioTi of South Carolina, but the holotspe 
(ANSP 16415, Figure 39. Table 2) is clearly not a Conus. 
Campbell (1995: 146) stated that C. connacki "is actually 
a broken volutid. " 

Several otlier Eocene Conns species are also problem- 
atic. Conus pulcheninms Heilprin, 1879 (type, AMNH- 
FI 10175) was recognized by Harris (1895) as a turrid 
(see Palmer and Brann, 1966). The t\pe specimen of 
Conus (Conospinis) granopsis de Gregorio (1890) is lost 
(Palmer and Brann, 1966) and Dall (1896) and Palmer 
(1937) suggested that the small shell figured bv de Gre- 
gorio (1890) mav be the juvenile of another species. 
Gi\"en that the type specimen is lost and that the shell is 
likel\- a juvenile specimen, C. granopsis is a name that 
should probabh' be disregarded. 

Finallw along with Conus sp. B (described here as the 
new species C. palmerae: holot\pe, UF 108858), Palmer 
in Richards and Palmer (1953) also noted a Conus sp. A. 
She said, "species A ... is a broad (21 mm.), low-spired 
(7 mm. ) shell with sharp angulation of the shoulder of the 

w'liorls; the surface was apparently smooth. The speci- 
men is a fragment, 25 mm. high" (Palmer, 1953: 40). We 
located this specimen (UF 108683), which — like UF 
108858 — is also a calcite pseudomoiph from UF locahty 
L\'()14, Gulf Hammock 02, Levv Counts', Florida (lower 
memlier of the Ocala Limestone). UF 108683 is poorlv 
preser\'ed and is too fragmentary (more than half of the 
spire is eroded away and much of the last \\'horl is miss- 
ing) to either assign to a knowii species or to describe as 
new species. 


Sable .Allen (Alexandria, Louisiana) gracioirslv donated to 
the FLMNH the large, well-documented Eocene Gulf 
Coast moUusk collection of her late husband, James E. 
Alien. Larry Rogers (Limestone Products), Wayne Bea- 
ver (Denali Limerock Mine), Joe Horton (Suwannee 
American Cement), and Leon Brooks (Hi-Cal Quany) 
kindly allowed access to collect in their Florida mines. 
Paul Callomon (Academy of Natural Sciences, Philadel- 
phia) and Ann Mohneux (Texas Memorial Museum) are 
thanked for allowing access to ty[3e specimens. Ann Mo- 
lineux also provided helpful discussions on Eocene stra- 
tigraphy in Texas. Bushra Hussaini and Sandy Ebersole 
furnished information pertaining to an American Mu- 
seum of Natural Histon' specimen and a Geological 
Sur\'e\" of Alabama specimen, respectiveK'. ^Alan Kohn 
providetl helpful discussions of and suggestions tor ter- 
minologs' related to the "subsutiu'al flexure" of cone 
sfiells. Sean Roberts (Florida Museum of Natural His- 
toiy) assisted us with some of the digital photography 
using a Sony DSC Rl camera (10.3 megapixel resolu- 
tion). We thank Alan Kohn and an anon\'mous reviewer 
for connnents that improved the quality of this manu- 
script. JRH's contributions to this work were supported 
hv the Bra\iield Award of the South West Florida Fossil 
Club and NSF EAR 0518976. RWP's contribution to this 
work was supported by the McGinty Endowanent of the 
Florida Museum of Natural History, by Barbara and 
Reed Toomey, and by the late Da\id Nicol whose SIOOO 
grant allowed RWP to permanentlv transfer the Florida 
Geological Suney Eocene mollirsk collection from die 
Paleontological Research Institution (where it was under 
study by Katherine V.W. Palmer) to the Florida Museum 
of Natural Histoiy. This is University of Florida Contri- 
bution to Paleobiolooy 602. 


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THE NAUTILUS 122(2):94-9S, 20()S 

Pasie 94 

Synonymization olNeohi/aliinax Simroth, 1896, and Omalonijx 

d'Orbigny, 1837, with a redescription of Omalonijx bmsiliensis 

Simroth, 1896) (Gastropoda: Succineidae^ 

Janine O. Arnida 

Laboratono de Malacologia 

Museu de Ciencias e Tecnologia 

Pontificia Universidade Catolica do Rio 

Grande do Sul 

A\enida Ipiranga 6681, 

90619-900 Porto Alegre, BRAZIL 


Jose W. Thome 

jose\vthoine(s'p< J. 


Neolu/alimax was established as a monotypic genus for A', bm- 
siliensis. Cliaracters of the reproductive system prompted au- 
thors to subsequently treat the taxon as a subgenus of Omal- 
onijx. Examination of the holot)'pe and discussion of these char- 
acters allowed Neohyaliniax to be synonyniized witli Omalonijx. 
Onialonijx bmsiliensis remains a \alid species based soleh' on 
the holotyqie. 

Additional Kci/ieords: Mollusca, Pulmonata, Brazil, Rio Grande 
do Sul 


Tiie genus Onwloiu/x (.lOrbigm, LS.37, comprises siicci- 
neids slugs of West Indies, Central and South American 
and Juan Fernandez, with a reduced. Hat and fingernail- 
hke shell. Omaloiu/x has commonly been classified with 
two subgenera (e.g., Patterson, 1971; Tillier, 19S1), 
namely Omalom/x sensu stricto and 'Neohijaliinax Sim- 
roth, 1896. Zilch (1959-1960) and Thiele (1992) indi- 
cated that the separation between the subgenera Oma- 
lonijx and Neolu/alimax is based on the mantle covering 
either only the margin or the entire shell and the spire 
being small or scarcely indicated. The iiionot)'pic subge- 
nus Neohi/aUmax, tyjje-species Neohijalimax 
Simroth, 1896, was described Irom a nnicjue specimen 
collected irom Rio Grande do Sul, Brazil. In revising the 
family Succineidae Beck, 18.37, Patterson (1971) dis- 
cussed tlie similarities of Omaloni/x fclina (Guppy, 1872) 
[= O. mathemni (Potiez and Vlichaud, 1835), according 
to Tillier, 1981] and N. hrasiliensis, and brought atten- 
tion in particular to details of the original description of 
the latter species (namely the number of seminal vesicles 
and cephalic tentacles) that suggested recognition of 
Neohijalimax at the subgeneric level. It should i)e noted. 

however, that Patterson (1971) did not examine the type 
of A', hrasiliensis, nor made a direct comparison with O. 
unguis (d'Orliignv, 1837), the t^pe species of Omak^nt/x. 
In a revision ol the South American and Juan Fernandez 
succineid slugs, Tillier (1981) similaiK did not examine 
the ty-pe of N. hrasiliensis, but accepted Patterson (1971) 
assessment of taxonomic status based on the original de- 
scription provided by Simroth (1896). In support of rec- 
ognition at the subgeneric rank, TiUier (1981: 144) noted 
"an altogether different way ol life (terrestrial), a plesio- 
moi"phic radula (marginal teeth still longitndinalK elon- 
gated) and slug-like characters more apomoiphic than in 
Omalonijx sensu stricfo\ 

Simroth (1896) mentioned that the t\pe specimen of 
A', hrasiliensis was deposited with the "Berlin Museum". 
Indeed there is a specimen in Museum fiir Naturkunde 
der Humboldt Universitat zu Berlin (ZMB 45.913) la- 
beled Neohijalimax hrasiliensis, but it bears no indication 
of type status. That the specimen corresponds well wth 
the description given liy Siinr(.ith (including most details 
of the internal anatomy) and that no other specimen of 
this species has been located, the ZMB specimen is here 
considered to be the holotyjie. 

With the recent I'ediscoveiy of Simroth's t\pe speci- 
men, we take the opportunit)^ to redescribe N. hrasilien- 
sis and reassess its taxonomic status. 


The following lots were examined: GUADELOUPE, 
Pointe-a-Pitre, Omalonijx matheroni , 4 spec, 
27.IX.I972, Pointier leg. (MNHN); O. matheroni. 11 
spec, S. Tillier det. 1979. (AINHN); FRENCH C;UI- 
ANA, along the road Cavenne-Kourou, 17 km SE 
Kourou bridge, O. matheroni, 5 spec, Tillier leg. 
(MNHN); PARAGUAY, O, unguis (lectotype designatexl 

J. O. AiTuda ami j. W. Thome, 200S 

Page 95 

bvTillier, 1981), 1 shell. (MNHN); Asiineion, Asuncion, 
6. iiitiiiii^!. 1 spec, canje F.H. Sehade. (MACN19968, it 
is labeled as Hoin. unguis d'Orb.); ARC;ENTINA, For- 
mosa, Mllat'ane (26"14.17' S .59"07.()S' W, Arnno Bel- 
laco, Estancia La Marcela), O. iiiignis, 2 spec, 
30.X.20()4, Cristian Ituaite leg. (MLP 1 187S); BRAZIL, 
Rio Grande do Sul, Xcohijalimax brasilicnsis Simrodi, 1 
spec (ZMB 45.913); Cachoeirinha, O. convexii.s. 34 
spec, 29.IV.2006, A. Paladini and J. O. Armda leg., J. O. 
Armda det. (MCP 8839); Pelotas (Estrada Pelotas-Rio 
Grande, 31°45'53.1" S 52°22'48.2" W), O. convcxiis. 48 
spec, 1S.III.2006, A. Paladini and J. O. Arrnda leg., J. O. 
Armda det. (MCP 8836). 

Based on examination of the holotspe oi' Nciiluidliiiuix 
brasilicnsis, we redeseribed the reproductiw system 
(Figure 1). The specimen had been previouslv dissected, 
\\ith a sagittal cut from its head to the mantle edge, 
which dien continues towards the left side of animal's 
tegument. The reproducti\e system was separated in t\vo 
parts: one including the \agina (pre\iously dissected and 
\\itli the papillae on its internal surface exposed), the 

Figure 1. Reconstruction of the reproductive svstem of 
Omalomjx brasilicnsis (Simroth, 1896) (Z.VIB 45.913). Abbre- 
\iations: ag, albumen gland; b, epiphallial bulb; dd, deferent 
duct: ep, epiphaUus; fc, fertilization complex: fo, free oviduct; 
gg, ganietoHtic gland: o, ovariotestis; ov, oNiilispermduct; p, 
evertophallus; po, pallial oviduct; pr, prostate; ps, evertophal- 
lus sheath: rm, retractor muscle: sv, seminal vesicles; v, vagina. 

gametolitic gland duct and its gland antl the tree oviduct; 
the other including the pallial oviduct, prostate, albumen 
gland, fertilization complex, ovuli.spermduct, ovariotestis, 
deferent duct, epiphallus and e\ertophallus (= penis). 

Institutional ablu'eviations: MACN, Museo Argentino 
de Ciencias Naturales "Bernardino Rivadavia" e Instituto 
Nacional de Investigacion de las Ciencias Naturales 
(Buenos Aires, Argentina), MCP, Museu de Ciencias e 
Tecnologia da Pontificia Universidade Catolica do Rio 
Grande do Sul (Porto Alegre, Brazil); MLP, Museo de La 
Plata (La Plata, Argentina), MNHN, Museum national 
dllistnire naturelle (Paris, France) and ZMB, Museum 
liir Naturkunde der Humboldt Universitiit zu Berlin 


Order Hetemrethra Pilsbiy, 1900 
Superiamily Succineoidea Beck, 1837 
Family Succineidae Beck, 1837 

Genus Oiniihmiix d'Orbigny, 18.37 

Type Species: Siiccinea (Oinaloni/x) uiiis^iiis d'Orbigny, 
1837: 229 (year according Sherborn and Woodward, 
1901), by original designation. 

Diagnosis: Shell veiy flat, fingernail-shaped, obtuse 
and small spire, body whorl and aperture veiy large and 
elongate-oval, animal body twice the shell lengdi; e\'er- 
tophallus exceptionally long. 

Oinaloin/x brasilicnsis (Simroth. 1896) 

Neolnidliiniix hrasiliensis Simroth, 1896: 39—4.5; Morretes, 

1949: 130; Simone, 2006; 237. 
Oinalonyx {Neohijaliinax} brasilicnsis. — Zilcli, 1959-1960; 202; 

Patterson, 1971: 187: Tillier: 1981. 
Hoinalonyx {Neohijalyinax) brasilicnsis. — Thiele, 1992. 
Omalomjx brasiliemis. — Salgado and Coellio, 2003: 153. 

Diagnosis: A species oi Onwloiu/x with a thick-walled 
vagina with robust diamond shaped papillae on its inter- 
nal sui'face. 

Holohpe: ZMB 45.913 

T\pe Locality: Brazil, Rio Grande do Sul. 

Redeseription: The shell is thin, oval and higliK' flat- 
tened (it is fragmented, wliich precludes its measure- 

The jaw and radula are missing (see Discussion be- 
low). Within the reproductive system, the epiphallus is 
approximatelv one fourth of tlie length of the exertophal- 
lus. The former possesses a thick wall, a smooth outer 
face and a bulb, and its internal surface has transverse 
folds. The opening of the epiphallus into the evertophal- 
lus is a simple opening, without any papilla. 

The evertophallus has a tliick, wide wall, and the in- 
ternal surface is covered with aligned chamond shaped 
papillae. At its proximal extremity, the papillae are 

Page 96 

TIIK NAUTILUS, Vol. 122, No. 2 

smaller and di.scoid. At its distal extremibi' there are small 
longitudinal folds. The eveitophallus sheath, thick and 
muscular, covers practically the entire distal half of the 
eveitophallus. It graduallv becomes a thin and transpar- 
ent membrane, \vhich covers the other halt of the ever- 
tophallus and the epiphallus. The evertophallus retractor 
muscle inserts at the jiniction of the deferent duct with 
the epiphallus. 

The prostate is long and the section of the deferent 
duct that is embedded in the prostate has small orifices 
throughout its length within the gland. 

The vagina is approximately half of the length of the 
evertophallus. It has a thick wall and aligned, diamond 
shaped papillai' on its internal surface. These papillae are 
similar to those on the evertophallus, but more robust. 
Tlie gametolitic gland is spherical, with a diameter ap- 
proximatelv half the length of its duct. The gametolitic 
gland duct is slim and about two thirds the length of the 

The free oxdcluct is long, thick, and convoluted with 
rippled longitudinal folds on its internal surface. Tlie 
pallial oviduct, approximateh' half the length of the lower 
oviduct, is vei-v' inflated and convoluted. 

The albumen gland is small, bean-like and about one 
third the length of the prostate. The ovniispennduct is 
slightlv conxoluted and more inflated in its distal portion. 
The o\'ariotestis is spherical and large. 

The pallial cavity and the neiA/ous and digestive sys- 
tems are damaged. 


Sinn-oth ( lS9fi: 40, pi. \\ Fig. .3) characterized the shell as 
'a small flat plate of 9 mm in length and 6 nun in breadth. 
Beneath the light-vellowdsh periostracum the lime was 
deposited in concentric rings, with an excentric micleus 
to the left of the posterior end. The nearer the nucleus, 
the thicker is the shelK" matter. The periplien' is solid, 
surrounded, near the posterior right-hand margin, by a 
brown conchiolin line. The nucleus is somewhat thick- 
ened; the shell throughout is perfectly flat, and therefore 
does not enclose any of the soft parts". The shell, accord- 
ing to Sinu'oth (op. cit.), would place the species close to 
the genus Hijalbnax H. & A. Adams, 1855. However, that 
author recognized that the shell In Hi/alimax it is slightly 
arched (Fischer)'. Sirnroth (1896: 41) described the jaw 
as 'brown and [it] has a shaip edge, with a slight median 
process (oxi/gnaflj) and a lighter-coloiu-ed pakital plate 
(clafimot^nafh), the two posterior corners of which ai'e 
somewhat prominent. This plate is finely sculptured, the 
thread-like markings converging towards the median 

Simroth (1896: 41) described the ratlular teeth as fol- 
lows: 'the median tooth is tricuspid, the middle cusp 
Ijeing vei-y large. In the lateral teeth the median cusp is 
still larger, and directed ol)liquely towards the middle 
line of the radula; the inner cusp is reduced in size, whilst 
the outer one is duplicated. In tlie marginal teeth the 

inner cusp increases in size, whilst the exterior outer 
cusp subdivides, giving rise to three outer cusps'. 

In the description oiO.felina, Guppy (1872) indicated 
that the shell was usuallv covered bv the mantle, but 
when retracted, it occasionally exposed the central region 
of the shell. Gibbons (1879), however, repeatedly ob- 
sei^ved living specimens of Omalomjx and never noticed 
more than a limited portion of the shell border covered 
by the mantle. In observations of live specimens oi Oma- 
lomjx from the states of Rio Grande do Sul and Minas 
Gerais, Brazil, we found that the mantle covered tlie 
shell in different degrees, and although the shell was well 
covered in some, in none presented its shell completely 
covered by the mantle. Based on Guppy (1872) and on 
our obseivation of live specimens oiOmalonijx, the char- 
acter "mantle covering partially or totally the shell", in- 
dicated by Zilch (1959-1960) and Thiele (1992) for the 
separation of the subgeiu'e Omalomjx and Neohijalimax, 
are imjustifiable. The other character, "small spire or 
e.xtremely small spire" is not a systematic character. 

Patterson (1971), using Simroth's description of iVeo- 
lii/alimax hrasiUcnsis as reference, indicated that it has 
similarities with Omalomix fcliiia concerning the general 
bodv's shape, the mantle pigmentation and the anatomy 
of the reprochictive system. Because the reproductive 
system of Ncohi/alimax hrasilicnsis is similar to that in 
Omalomjx s.s and different from that in Hijalimax, which 
resembles Siiccinca Draparnaud, 1801, Patterson consid- 
ered Neohi/alimax as a subgenus oi Omalomjx. However, 
Patterson also noted that Ncolii/aUmax brasiliensis has 
one seminal vesicle, whereas there are two in O. felina, 
and that N. brasiliensis has only one pair of tentacles, 
while O. felina has two. We verified the presence of two 
vesicles and one fecundation pouch in N. Ijrasilieusis, 
which contradicts its original description by Simroth 
(1896) and conciu's with the obsenations of Patterson 
(1971) for Omalomjx sensu stricto. 

Simroth (1896) obsen'ed only one pair of tentacles in 
N. brasiliensis, the onnnatophores. Patterson (1971), 
liowever, stated that Omalomjx sensu stricto has two 
pairs of tentacles. Through an external ventral view exam 
of the anterior region of N. brasiliensis, we could obsei've 
that the lower tentacles are present, verv close to the 
animal's mouth opening. 

Simroth (1896) did not mention the presence of an 
epiphallus in N. brasiliensis. However, we verified the 
presence of tlris structiu'e, which is slightly narrower than 
the evertophallus and is also present in the otlier species 
of Omalomjx s.s. We also olisei-ved small longitudinal 
folds at the distal extremit)' of the evertophallus, which 
are present in all Omaloni/x species with kniown anatomv . 

Bai'ker (2001) noted tlie placement of the genital ap- 
erture in Neohijalimax immediately adjacent to the right 
tentacle. We confirmed that this does not differ from its 
position in other species of Omalomjx sensu stricto. 

In the state of Rio Grande do Sul, t\pe localitv of O. 
brasiliensis, no specimens with the characteristics ol this 
species were found, despite extensive investigation and 
collecting. In the mimicijialities of Gachoeirinha, Pelotas 

O. Arrada and T. W. The 


Page 97 

and other locaHties in Rio Grande do SnI, onI\ C)iiuiloiu/x 
convcxiis (HeMieniann, 1S6S) \\as found, suggesting its 
o\'erlapping geographical area with O. hmsilioisis. The 
reproducti\e systems of these species are distinct. In O. 
convcxns. the ovariotestis is hemispherical or discoidal; 
the lower o\iduct is generallv halt the length ot the upper 
oviduct; half of the proximal region of the e\ertophallus 
is generallv a little wider than the rest, and its internal 
surface has longitudinal folds in the wider proximal re- 
gion and cordiform papillae in the distal region; the in- 
tern;il surface ot the \agina has smooth longitudinal 
folds. In O. bmsilicnsis, the o\'ariotestis is spherical and 
large; die lower o\iduct is approximateK' b,\ice the length 
of the upper oviduct; the internal surface ot the ever- 
tophallus has diamond shaped papillae antl the vagina 
has similar but more robust papillae. 

The reproductixe s\stem of O brasiliensis resembles 
diat in the studied specimens oi O. matheroni (Potiez 
and Michaud, iS35), which agrees with the meticulous 
description of Tillier (1981), with the exception of the 
inteniiil surface of the \'agina. According to our obsena- 
tions and to Tillier, the vagina in O. luatlicroni has ir- 
regular longitudinal folds, inconsistent in number or 
shape, whereas O. bra.siUeiisis has papillae instead of 
folds, similar to the ones in its evertophallus. Despite the 
reproductive s\steni similarities of these species, they do 
not have overlapping ranges. Tillier (1981) considered 
the probable range of O. matheroni to be the entire 
Amazon basin, the eastern coastal regions of South 
America from Caracas (Venezuela) to Rio de Janeiro 
(Brazil) and part of the Lesser Antilles. Neohijalimax bra- 
siliensis has been reported only once, from Rio Grande 
do Sul (Brazil). 

When compared to the O. unguis (d'Orbigny, 18.37) 
1)^6 species, O. brasiliensis presents specifics different 
characteristics such as a smooth external surface of the 
epiphaUus. the internal surface of the evertophallus and 
\agina has diamond shaped papillae, more robust in the 
latter, the length of the gametolitic gland corresponds to 
appro.ximatelv a half the length of its duct and the lower 
oviduct, which is little convolute, is generally two times 
the lengdi of the upper oviduct. In O. unguis, according 
to .\rruda et al. (2006), the external surface of the 
epiphaUus has a looped fold, the internal surface of the 
evertophallus has discoidal papillae, the internal surface 
of the vagina has inflated elliptic papillae, the length of 
the gametolitic gland corresponds to appnjximatelv a 
quarter the length of its duct and tlie lower oviduct, 
which is convolute, is approximatelv three times the 
lengdi of the upper oviduct. 


Considering the foregoing discussion regarding the num- 
ber of seminal vesicles, the number of tentacles and the 
position of the genital aperture, as well as the observa- 
tions of the reproductive system, we here consider Neo- 

liildliiiiax Siiinoth, 1896 as a jiuiior sviionvm of Ouialo- 
ni/x d'Orbignv, 1837. Ouialoni/x bmsiliensi.i (Simroth, 
1896) remains a valid species, based on the only known 
specimen, which we consider to be the holoty^^e. 


We thank the reviewers. Dr. Gaiv Barker and Dr. Robert 
Cowie, for their great contribution with veiy good criti- 
cisms and improving the English; Alejandro Tablado 
(MACN), Cristian Ituarte (MLP), Lucia Maria Z. Rich- 
initti (MCP), Virginie Heros (MNHN) and Matthias 
Glaubrecht (ZMB) for loaning material; Guacira Gil for 
comments and suggestions; Luiz Augusto F. Araiijo for 
the translation from Portuguese to English; to Conselho 
Nacional de Deseiivolvimento Cientifico e Tecnologico 
for financial support (CNPq 131716/2005-0 for JOA, 
500032/2003-.3 for JAVT). 


Armda, J. O., S. R. Gomes, R. Ramirez, and J. W. Thome 2006. 
VIorfoanatomia de duas especies do genero Omalonijx 
(MoUusca, Pulmonata, Succineidae) com novo registro 
para Minas Gerais, Brasil. Biociencias 14 (1): 61-70. 

Barker, G. M. 2001. Gastropods on land: phylogeny, diversitv' 
and adaptive moqshologv. In: Barker, G. M. (ed.) The 
biology ot terrestrial molluscs. CABI Piililishing, New- 
York, pp. 1-146. 

Beck, H. 18.37. Index MoUuscorum praesentis aevi Musei Prin- 
cipis Augustissimi Christiani Frederici. Gopenliagen, 124 

Gibbons, ]. S. 1879. Comparison of Omtiloni/x i(i7gi«,s d'Orb., 
with O, felinci Guppv. Jo\n"nal of Gonchologv' 2: 98-101. 

Guppy, R. J. L. 1872. Third series of additions to the catalogue 
of the land and freshwater MoUusca of Trinidad, with a 
revised list of all species. Proceedings ot the Scientific 
Association of Trinidad 2: 17-25. 

Morretes. F. L. 1949. Ensaio de Gatalogo dos Moluscos do 
Brasil Arquivos do Museu Paranaense 7(1): 1-216. 

d'Orbigny, A.D. 1837 [18.3.5-1847]. Voyage dans 1' Amerique 
Meridionale: MoUusques. Paris: Pitois-Levraut 5(3): 

Patterson, C. M. 1971. Taxonomic studies ot the land snail 
family Succineidae. Malacological Review 4: 131-202. 

Pilsbrv', H. A. 1900. On the zoological position of Partula and 
Achatinella. Proceedings of tlie Academy of Natural Sci- 
ences ot Philadelphia 3: 561-567, pi. 17. 

Salgado, N. C. and A. G. S. Goelho. 2003. Moluscos terrestres 
do Brasil (Gastropodes operculados ou nao, exclusive 
\'eronicellidae, Milacidae e Limacidae). Revista de Bi- 
ologia Tropical (International Journal ot Tropical Biologv 
and Gonseivation) 51 (suppl. 3: Malacologi'a Latinoameri- 
cana): 149-189. 

Sherbom, C. D. and Woodward, B. B. 1901, Notes on the dates 
of publication of the Natural Historv- poitions ot some 
French X'oyages — Part I. '.Amerique meridionale'; 'Indes 
orientales'; 'Pole Sud'; ('Astrolabe' and Zelee'); 'La Bon- 
ite'; 'La Coquille'; and "L'Uranie et Physicienne'. Annals 
and Magazine of Nat\n-al Histoiv' (series 7) 7 (40): .388- 

Paee 98 

THE NAUTILUS, V()I. 122, No. 2 

Simone, L. R. 2006. Land and treshwater inolluscs ol Brazil. 

Fapesp, Sao Paulo, 390 pp. 
Simroth. H. 1896. On Neoluialiinax hrasiliensis, n. gen., n. sp. 

(allied to Hi/aliimix), from Brazil. Proceedings of the Mal- 

acological Societ\' of London 2: 39-45. 
Thiele, J. 1992. Handbook of Systematic Malacolog)'. Part 2 

(Gastropoda: Opistliobranchia and Pulmonata). In: Bieler, 

R and P. Mikkelsen (translatoi's). Smitlisnnianlnstitntion 

Libraries. The National Science Fomidation, \Vashington 

627-1189 pp. 
Tillier, S. 1981. South Anieincan and Juan Fernandez succineid 

slugs (Pulmonata). Journal of Molluscan Studies 47: 12.5- 

Zilch, A. 1959-60. Euth>iienra. In: Wenz, W. (ed.). Handbuch 

der Palaozoologie, Gebriider Borntraeger, Berlin, vol. 2, 

834 pp. 

THE NAUTILUS 122(2):99-106, 2(K).S 

Page 99 

Rehabilitcition oi Er^^alatax iiiaiiciisi (Schepman, 1892^ 
(Gastropoda: Miiricidae), senior synonym oi Ergalatax ohscurci 
Honart, 1996, and description of ErgaJatax junionae, new name 
for Moriild iiuuioisi Dall, 1923 

Roland Houart 

Institut ro\'al des Sciences naturelles de Beli;ii|ue 

Rue X'autier, 29 

B-1000 Bioixelles, BELGIUM 



Eraalatax jiinionac nomen iioxuin is proposed as replacement 
name for Morula maiiensi Dall. 1923. from the Gulf of Oman 
and the Persian Gulf The latter name is a junior secondary 
homonMU oiPentadactijlus mai-tensi Schepman, 1892, from tlie 
Red Sea. of which £. obscura Houart, 1996, is here considered 
as a junior s^•non\^^^. Ergalatax junionae nomen novxim is com- 
pared with Ergalatax martensi (Schepman) and E. margariti- 
cola (Broderip. 1S33). The geographical distribution of E. 
junionae and E. maiiensi is updated and the introduced Medi- 
terranean species is identified as E. junionae. 

Additional Ki'ijuords: .MoUusca, Gastropoda, .Muricidae. Gulf 
of Oman, Persian Gulf, Red Sea, Mediterranean Sea, hom- 
on\Tn\'. s\Tion\Tn\'. new nan ; i ■ 


Some vears ago (Houart. 1996: 13), I described Ergalatax 
obscura. a muricid from the Red Sea. Shortly after that 
paper was published, R. G. Moolenbeek (in Htt.) told me 
about a name introduced bv Schepman (1892), Pcnta- 
dactijliis maiiensi. which he also described from the Red 
Sea. Ha\ing then examined a s\iit\pe of that species 
from ZMA (Figure 14), I concluded that P. maiiensi 
Schepman, 1892, was a junior svnonvm of Ergalatax 
maroariticola (Broderip, 1833). 

However, after recenth requesting additional infor- 
mation about the other svnt\p)es of P. maiiensi in 
NMNL, I receixed digital images that illustrate the re- 
maining three specimens (Figures 18-23), among them 
the 17 mm long specimen that probabK' ser\'ed for the 
description (Figures 18-19). Those s\iit\p)es are un- 
doubtedl\' conspecific with E. ohscurn, while the ZMA 
s\Tit\pe represents E. margarificola. 

For manv vears (Houart, 2001). 1 had identified an 
introduced eastern Mediterranean species as Ergalatax 
obscura Houart, 1996 [= E. maiiensi (Schepman, 1892)]. 

However, this concerns yet a third species, which occru's 
primarily in the Gulf of Oman and in the Persian Gulf. 
The species was first wrongly illustrated as Puiyura 
(Ricinula) siderea Reeve (a buccinid) bv von Martens 
(1874: 95, pi. 5, fig. 49). Later, Dall (1923: 304) proxided 
a new name for that species, naming it Morula maiiensi. 
Unfortimately, both species, Pentadacti/lus maiiensi 
Schepman, 1892, and Monda maiiensi Dall, 1923, are 
no\\' included in Ergalatax, and Dall's name becomes 
ij^so facto a jvmior seconclan" homonym and needs to be 

I personally mixed botli species because of their close 
relationship and the presence of some related forms 
(compare Figures 30 and 35). First (in lift.), I provision- 
ally identified the Mediterranean specimen as E. inar- 
tensi (Dall. 1923), but aftenvards I erroneously consid- 
ered it as conspecific xxith E. obscura [= E. maiiensi 
(Schepman, 1892)] from the Red Sea and the Gull of 
Aden (Houart, 1996: 12). 

Abbre\iations tised herein are: AMS: Australian Mu- 
seum, Sydney, Australia; MNHN: Vluseum national 
d'Histoire naturelle, Paris, France; NMNL: National 
Museum of Natin-al HistoiT Naturalis, Leiden, The 
Netherlands; RH: collection of the author; ZMA: Zoolo- 
gisch .Vluseum, University of .Amsterdam, The Nether- 
lands; ZMB: Museum fiir Naturkunde der Humboldt 
Unix-ersitat zu Berlin, zoologisches Museum, Germany. 
Terminolog\'' for shell spiral cords (after Merle. 1999 ;uid 
2001) gi\'en in Figure 1. 


Subfamily Murieoidea Rafinesque, 1815 

Family .Muricidae Rafinesque. 1815 

Subfamily Ergalataxinae Kuroda, Habe and 0\'am;i, 1971 

Genus Ergalatax Iredale, 1931 

Type Species: Ergalatax reeurrens Iredale, 1931, -\us- 
traha (Figure 17) [?= E. contraefa (Reeve, 1846)] by 
original designation. 

Page 100 

THE NAUTILUS, Vol. 122, No. 2 



Subsutural cord 


Infrasutiiral primary cord (primary cord on shoulder) 

adis : 

adapical infrasutural secondary cord (shoulder) 

abis : 

abapical infrasutural secondary cord (shoulder) 


Primary cord 

s : 

secondary cord 

t : 

tertiary cord 

PI ; 

Shoulder cord 

P2-P6 : 

Primary cords of the convex part of the teleoeonch whorl 

sl-s5 : 

secondary cords of the convex part of the teleoeonch whorl 

example; si = secondary cord beUvcen PI and P2; s2 = secondary cord between P2 and P3, 



adapertural primary cord on the siphonal canal 


median primary cord on the siphonal canal 



Infrasutural denticle 

Dl toD5: 

Abapical denticles 

Figure 1. Terminology used to describe the spiral cords (al- 
ter Meile, 1999 and 2001) (sporadic sculpture is showai in pa- 

Ergalaliix iiuiticnsi (Schepiiian, 1892) 
(Figures :3, 5, 6, 11, 1:3, lS-.3(), 46) 

Pcntachicti/his {Morula) maiicnsi .Schepnian, 1892: 104. 
Cronia iiiai-tensi Dall. — Sharabati, 1984: pi. 19, fig. 9, 9a, 9b; 

Singer and Mienis, 1991b: 58, fig. 19; Coulombel, 1994: 

73, text fig. (not Morula maiicnsi Dall, 192.3); Verbinnen 

and Dirkx, 2000: 69, fig. 9 (not Monila maiimsi lOall. 

Dnipella nigosa. — Singer and Mienis, 1991a: 18, fig. 6 (not 

Murex rugosus Born, 1778). 
Ergalatax obscura Houart, 1996: 13, figs 1, 3-6; Houart, 2001: 

108 (in part), figs 17, 31, 106 and 449 only; Heiman and 

Mienis, 2003: 22-23 (text fig.). 
Not Morula martensi (Schepman, 1892) -Tan, 1995: 160, figs 

52, 192 g, h (= Ergalatax jujiioiiae nom. nov.). 
Not Ergalatax obscura. — Houart, 2001 (in part): lOS (in part), 

figs 31 and 450-451 only; Delongueville and Scaillet, 2007: 

57, tig. 31 (= Ergaliitcix junioiuie nom. nov.) 

Description: Shell medium sized for the genus, up to 
25.5 mm in length at maturity. Length/width ratio 1.39- 
1.92. Heavy, stout. Spire liigh with 3+ protoconch whorls 
(tip partially broken), and up to 7 broad, strongly shoul- 
dered teleoeonch whorls. Suture adpressed. Protoconch 
conical, acute, whorls smooth. Axial sculpture of teleo- 
eonch whorls consisting of high, rounded, nodose ribs: 10 
or 11 from first to penultimate whorl, 6 to 8 on last whorl, 
rarely 5 or 9. Spiral sculpture of high, strong, primaiy, 
secondaiy and tertiary cords. Last teleoeonch whorl with 
adis, IP, abis, PI, immediately followed by small P2, s2, 
P3, (t), s3, P4,(t), s4, P5, (s5); P6, ADP, MP. PI and P2 
small, P3, P4 and P5 similar in strength, P6 veiy small or 

Sculpture forming high, nodose knobs at intersection 
of spiral cords and axial ribs. Aperture relatively small, 
ovate. Columellar lip with 2 or 3 weak kmobs abapieally, 
rim adherent. Anal notch broad, moderately deep. Outer 
lip weakly crenulate, with 7 strong elongate denticles 
within [ID, D2 (D1-D2 fused), DS'split, 04, D5 split]. 
Siphonal canal short, broad, i)roadly open. Milky white, 

Figures 2-5. Ergalatax species. Spiral .sculpture and apcrlnral denticles. 2, 4. Ergalatax juiiiniiac noiiicii nox iiiii (lectotvpe ZM15). 
3, 5. £. auirtcnsi (Schepman, 1892) (Red Sea, RH). 

R. Houait. 20C).S 

Page 101 


Figures 6-13. Ergalatax species. 6. Radula oi Ergalatax martensi (Schepman, 1893), Gulf of Aden. Scale bar = 150 jxm. 7. Radula 
oiE.junionae nomen no\xini. Gulf of Iskenderun, Turkey. Scale bar = 120 \x.m. 8. Radula of £. margaiiticola (Broderip, 1833), Palau. 
Scale bar = 120 \i.m. 9-10. Radula of £. contractu (Reeve, 1846). 9. Aden, juvenile specimen. Scale bar = 60 |jim. 10. Korea, large 
adult specimen (lateral denticles worn). Scale bar = 150 (jLm. 11. Operculum of £. martensi Scale bar = 1 mm. 12. Operculum of 
E. junionae. Scale bar = 1 mm. 13. Protoconch of £. niartensi. Scale bar = 0.5 mm. (Figures 6-11, photos A. Waren). 



THE NAUTILUS, Vol. 122. No. 2 


\. ^ 17 

Figures 14-17. Ergalatax species. 14. En^ahitax iiuii'ij^drilifiihi (liroclerip, 18.3.3). LectoUpe of Pciildcliuiiiliis {Monila) martcnsi 
.Schepiiian. 1893. "Red Sea, coll. Forskal", ZVIA Moll. 24.8 mm. 1.5-16. Erffilafax junionae noiu. nov. Lectot)pe oi Morula 
iiuii-tcnsi Dall. 1923. ZMB 21596, 21.4 mm. 17. £rg«/rtte.v rcciirrens Iredale, 1931. Hulohpe AMS C57761, 25.5 mm (photo courtesy 
E. H. Vokes). 

cream\' white, or tan, iisiialK with some light to dark 
bro\\a'i colored .spiral cords on shoulder and on more 
prominent nodes. Aperture cream or pale vellow within. 

Operculum ergalat;L\ine (Fig. 11), D-shapecl with lat- 
eral nucleus in lower right. 

Radula (Fig. 6) with a racliidian bearing a long, narrow 
central cusp, a small, narrow, triangular lateral denticle 
and a broad, long lateral cusp on each side. Sickle-shaped 
lateral teeth broad at base and narrow at their end. 

Type Material Examined: .3 syiityi">es NMNL, Red 
Sea: RMNH.MOL.57165, here selected as lectot^pe and 
paralectot)pes, 1 symtype ZMA Moll.,5, Red Sea, 
here selected as paralectotype; Ergalatax obsciira 
Houart, 1996: Perim, Strait of Bab el Mandeb, holo- 
t\pe and 4 paratypes MNHN, 1 paratspe RII; Djibouti, 
Obock, Gulf of Aden, 9 paratypes MNHN; Yemen, 
Aden, 4 paratopes MNHN, 1 paratxpe RH. 

Othe Material E.xainined: Djibouti: Obock, Gull of 
Aden, 2 RH; Gulf of Aden: 2, RH Rod Sea: (no other 
data), 1 RH; Massawa, Taulud Is., 1 RH; Egy^jt, Sinai, 
mider stones at low tide, 10 RH; Eg)pt, Sinai, Shanu El 
Sheik, 1 RH; Egypt, Sinai, Shark's Bay (niarsa umm 
mureihha), under stones, low tide, 2 RH. Gulf of Aqaba: 
Israel, Eilat, 0.5-1.0 m, 3 RH; Gulf of Aqaba, Israel, 
Eilat, imder stones, 1-2 m, 4 RH. 

Distribution: From the Gulf of Aden to Eilat, Gulf of 
Aqaba, 0-2 m, on and under stones (Fig. 46). 

Remarks: For a comparison witli Eriiahihix junioiidc 
see that species below. 

Ergalatax junionae noiuen novmu 
(Figures 2, 4, 7, 12, 15-16, 31-40, 46) 

Monila sidcrea Ree\e. — von Vlartens, 1874: 95, pi. 5, fig. 49 

(not Riciniila siderca Reeve, 1846). 
Monda inai-teiisi Dall, 1923: .304. new name for Monila siderea 

von Martens, 1874, not Reeve, 1846. 
Crania konkanensis^ — Bosch and Bosch, 1982: 95, text fig; 

Sinythe, 1982: 60, pi. 1, fig. i; Bosch and Bosch, 1989: 60, 

text fig. (not Riciniila konkanensis MeKill, 1893). 
Crania cf. konkanensis. — Giunchi and Tisselli, 1995: 8, text 

Ergalaiax maiicns. — Buzzurro, Engl ami Tiimtiirk, 1995: (no 

pag,), text fig,; Engl, 1995: 46,' fig. 10, 
Crania cf konkanensis^—Bosch et al,"l995: 121, fig. 480. 
Manila martensi (Schepman, 1892).— Tan, 1995: 160, figs 52, 

192 g, h (not Pcntadactijliis (Monila) inaiiensi Schepman, 

Ergalatax obscura. —Houart, 2001: 108 (in part), figs. 450^51 

only; Delongneville and Scaillet, 2007: 57, fig. 31 (not 

Ergalatax abscura Houart, 1996). 

Description: Shell medium sizetl for the genus, up to 
29 mm in length at maturit)'. LengtlVwidth ratio 1.91- 
2.03. Slender, lanceolate, heavy, nodose. Shoulder 
weakly sloping, conca\e. 

White or ci'eamy white with dark browii or blackish 
colored primary spiral cords and occasionally s2 or s3. 
Aperture glossy white. 

Spire veiy high with 3.5 protoconch N^lioi'ls ;uk1 teleo- 
conch up to 7 weakly convex, strongly shouldered, no- 
dose whorls. Suture adpressed. Protoconch small, coni- 
cal, acute; terminal lip raised, of sinusigeral tyj^e. 

A.xial sculpture of teleoconch whorls consisting of liigh, 
strong, broad, nodose ribs and erratically placed strong 
varices. Last teleoconch whorl with S-1 1 ribs, occasion- 
ally with one or two erratically placed, broad x'arices. 
Spiral sctdpture of high, strong, nodose and squamose 

R. Houart, 2008 

Pase 103 

Figures lS-30. Eroalatax martemi (Schepiiuiii. isy2,). lS-23. Lectohpe and paralectoUpcs liMMlMOL. 5T165 iplioto J. 
Goud). 18-19. 17.4 mm; 20-21. 15.T mm: 22-23. 17.3 mm. 24-25. Holohpe of £. ofescHra Houart, 1996. Perim, Strait of Bab el 
Mandeb. MXHN 0159, 24.2 mm (photo MNHX). 26. Eilat, Israel, RH, 20.8 mm; 27. Obock, Gulf of Aden, RH, 21.6 mm; 28. Eilat, 
Israel, RH, 25.1 mm; 29. Red Sea (no other data), RH, 21.4 mm; 30. Aden, Gulf of Aden, RH. 21 mm. 

Page 104 

THE NAUTILUS, Vol. 122, No. 2 

Figures 31-45. Ergalatux .species. 31-40. Ergalatax jwtionac iioinen ikimiiu, 31-33. Doha, VValiia, Qatar, RH. 31-32. 26.5 iiiiii; 
3.3-34. 22.9 mm; .3.5. Yalikent, Lskenderuii, Turkey, RH, 17.2 min; 36-37. Leliancui, Bay til' Juunieli. 19.6 mm; 38-40. Iskendenin, 
Bay of Lskendenin, Turkey, RH. .38. 25.6 mm; .39. 22.2 nun; 40. 22,9 uim. 41-45. Erotilaiax margariticola (Broderip, 18.33). 41. 
Raroia , Tumaotu Archipelago, RH, 28.4 mm; 42. Kai Is., Mollucas, RH, 20.5 mm. 43. Sdiilli (if New Caledonia, RH, 23.7 mm; 44. 
Park Beach, coast of Singapore, RH, .30,4 mm; 45. Kwajalein Atoll, RH, 25,9 mm 

R. Houart. 2008 

Page 105 

priiiuin-, secondan- and tertian' cords. Shoulder of last 
teleoconch whorl with broad SP, adis, IP. abis, PI, P2, s2, 
[t). P3, s3, P4,(s4), P5, i.s.5), P6. .ADP. MP: PI and P2 
narrow, P3, P5 and PR broad, similar in strength, P4 

Aperture large, narrow, o\ate; columellar lip smooth, 
entireh' adherent, with low parietal tooth at adapical ex- 
tremih'; anal notch deep, broatl; outer lip weakly erect, 
with lb, D2-D5. ID largest, broad; D2-D4 decreasing in 
strength abapicallv, D5 split; denticles elongate within 
apertiu'e. Siphonal canal short, broad, dorsally recurved, 
broadK' open. 

Operculum (Fig. 12) dark browii, D-shaped, witli lat- 
er;il nucleus in lower right; attached surface with about 8 
growth lines and broad, callused rim, about 30-40 % of 
opercular width. 

Radula (Fig. 7) with a rachidian bearing a long, slender 
central cusp, a small, short lateral denticle and a broad, 
long lateral cusp on each side. Lateral teeth sickle- 
shaped, with broad base and narrow end. 

T\'pe Material Examined: Morula luaiicnsi Dall, 
1923, lectot\"pe (selected by Houart, 1996) and 6 syn- 
t^pes ZMB 21596. 

Other Material Examined: Persian CTulf: Kuwait, 
Kuwait Cit^-. 4 RH: Qatar. Doha, under rocks, 30 RH; 
Sharjali, 25°20' N, 55°21' E, on rocks, 4 RH; .A.bu Dhabi, 
0.5-1.0 m, 3 RH. Gulf of Oman: A\ Hamra, near Qurm, 
10 km N\V of Muscat, 16 RH; Al Bustan, under rocks, 1 
RH. Lebanon: Beirut, harbour entrance, breakwater, 
max. 15 m. 2 RH; Bav of Jounieh (N), 10-25 m, 2 RH; 
Batroun, 1-4 m, under stones, 2 RH. Turkey: Gulf of 
Iskenderun, Iskenderun, under rocks, harbor, 1 m, S RH; 
Kale, beach, 6 RH; Yumurtalik, rock pools, 6 RH; Ya- 
likent. shallow water, on rocks, I RH; Bav of AntaKa, 6 
km off Kemer, 36°39' N. 30°33' E, on rocks with mussel 
banks, 0.5-1.0 m, 7 RH; Fethive-Oludeniz, on rock at 2 
m. 1 RH. 

Distribution: Gulf of Oman, Persian Gulf and eastern 
Mediterranean Sea, intertidal to 4 m, on and under rocks 
and stones (Figure 46). 

Etvmology: This species is named in honor of Mrs. 
Marie-Louise Buvle-Jimion (1916-2003), former librar- 
ian of the Belgian Malacological Societ)-. She was one of 
tlie mainstays of the Societ)^ together with her husband, 
Jean Buvle. 

Remarks: Ships docking at oil terminals in the Gulf of 
Iskenderun (eastern Turkey) could have introduced the 
species into the eastern Mediterranean Sea. As noted by 
Delonguexille and Scaillet (2007), the transport via ship 
hulls or baUast water may be suspected. 

The shell morphologv' of some specimens oi E. jiinio- 
nae (Fig. 35) is nearlv the same to £. mai-tensi (Fig. 30), 
however E. martensi differs in having a more strongly 
shouldered, broader shell, and a yellowish aperture in- 
stead of white, with 7 denticles within instead of 6 in £. 
junionae. Moreover, the spiral cords differ in number 
and strengdi, as described above and illustrated in Fig- 

Er^alalax manendi (Schepman. 1893) 
O Er^alatax funiunac nomen novum 

Fig. 46. 15istribution of Eri^/ihitax inartciisis and E. junionae. 

ures 2-5. Twenty-five specimens of E. inaiienai and a 
few more of E. junionae were examined to confirm the 
stability of these chfferences. 

Ergalatax martensi usually has 6—8 axial ribs on the last 
teleoconch whorl, rarely 5 or 9, compared to E. junionae 
which bears 8-11 ribs and varices on the last whorl. 

Ergalatax niargariticola (Figures 8, 14, 41—45), a very 
common Indo-West Pacific species is also related to E. 
junionae, however it is generally stouter and broader, 
with a wider shoulder, a more squamose spiral sculpture, 
more uniformly colored shell and different aperture 
color, being bluish-white, occasionalK' with a tinge of 
pink or mauve on the columellar lip r.v. completelv white 
in E. junionae. The shell morphology and color of E. 
niargariticola are highlv variable, however it is always 
easily distinguishable from E. junionae by one or more 
differences cited above. 

The three species are related to the ergalataxine Er- 
galatax contracta (Reeve, 1845) (Figures 9-10), a prob- 
able senior synonym of Ergalatax recurrens Iredale, 1931 
(Fig. 17), the type species of Ergalatax. 

Buzzurro, Engl and Tiimtiirk (1995) were the first to 
mention the presence of Ergalatax junionae [as Erga- 
latax niaiiensi (Dall, 1923)] in the eastern Mediterra- 
nean Sea. 

Tan (1995:147) in his Ph D. thesis also selected a lec- 
totype for Morula martensi Dall. 1923. However, he des- 
ignated a specimen of Ergalatax margariticola (Tan, in 
litt.) from Kingsmill Is (USNM 52472). Nevertheless, 
this designation being published in a thesis that does not 
satisfy Articles 8.1.2 and 8.1.3 of the International Code 
of Zoological Nomenclature, it is not available as such. 


I am verv grateful to Jeroen Goud. National Vluseum of 
Natural Histoiy Naturalis, Leiden for the digital images 

Page 106 

THE NAUTILUS, Vol. 122, No. 2 

of the type material ofPentaclacti/his niai-tcnsi and for his 
comments, to David Reid, Natural Histon' Museum, 
London, for providing me useful photocopies and for 
comments, to Robert G. Moolenbeek, Zoologisch Mu- 
seum, University of Amsterdam for the loan of the lec- 
totvqoe of P. inai-tensi, to the late Prof R. Kilias, Museum 
fiir Naturkrmde der Humboldt Universitiit zu Berlin, (or 
the loan of the specimen illustrated by Martens (1874), 
lectotype of Manila maiiensi Dall, to Anders Waren, 
(Natural Histon- Museum, Stockholm) for radula prepa- 
ration and SEM work of radula and operculum, to E. H. 
Yokes (Prof Emeritus, Tulane University, Louisiana) for 
the photograph of E. recurrens Iredale, 1931, to Greg 
Herbert (Universitv- of South Florida, Tampa, USA) (or 
his uselul conunents and suggestions, and to [ohn Wolfi, 
Lancaster, Philadelphia, for checking the English te.xt. 


Bosch, D. and Bosch, E. 1982. Seashells ot Oman. Longiiuin 

Croup, England: 206 pp. 
Bosch, D. and Bosch, E. 1989. Seashells of Soutlicm Arabia. 

Motivate, Dubai: 95 pp. 
Bosch, D. T., Dance, S. P., Moolenbeek, R. G. and Oliver, 

P. G. 1995. Seashells of Eastern Arabia. Ed. P. Dance, 

Motivate Publishing: pp. 1-296. 
Buzzurro, C., Engl W. and TUnitiirk, I. 1995. Bivalven und 

Gastropoden der Europaischen Meere (4): Ergalatax mar- 

tensi (Dall, 1923) (Muricidae), Ein neuer Lesseps'schere 

Einwanderer von der Tiirkischen Siidkliste. Club Con- 

chylia Informationen 27(1): 17-LS. 
Coulombel, A. 1994. Coquillages de Djibouti. Edisud, La 

Calade, Ai.\-en-Provence: 143 pp. 
Dall, W. H. 1923. Notes on Dnipa and Manila. Proceedings of 

the Academy of Natural Sciences of Philadelphia 75: 303- 

Delongueville, C. and ScaiUet, R. 2007. Les especes invasives 

de moUusques en Meditei'ranee. Novape.x 8(2): 47-70. 
Engl, W.1995. Specie prevalentemente Lessepsiane attestate 

lungo le coste Turche. Bolletino Malacologico 31(1-4): 


Giunchi, L. and Tisselli, M. 1995. Crania cf. kankanensis 
(Mehill, 1893), new Indo-Pacific host in the Mediterra- 
nean Sea. La Conchiglia 27(275): 8-9. 

Heiman, E. L. and Mienis, H. K. 2003. Shells of East Sinai, an 
illustrated list. Muricidae (2). Triton: 22-23. 

Ilouart, R. 1995. The Ergalataxinae (Gastropoda, Muricidae) 
from tire New Caledonia region with some comments on 
the subfamily and the description of thirteen new species 
from the Indo-West Pacific. Bulletin du Museum national 
d'Histoire naturelle, Paris, 4e ser., 16A(2-4):197-245. 

Houart, R. 1996. On the identity' of Mon//fl martensi Dall, 1923 
and description of a new species of Eroalatax from the 
Red Sea (Gastropoda: Muricidae: Ergalataxinae). The 
Nautilus 110(1): 12-16. 

Houart, R. 2001. A review of the Recent Mediterranean and 
Northeastern Atlantic species of Muricidae. EvoKer: pp. 

Martens, E. von 1874. Ueber \'orderasiatische Conchylien 
nach den Sammlungen des Prof. Hausknecht. Cassel, T. 
Fischer: 127 pp, 9 pi. 

Merle, D, 1999. La radiation des Muricidae (Gastropoda: Neo- 
gastropoda) an Paleogene: approche ph}logenetique et 
evolutive, These du Museum national d'Histoire naturelle, 
Paris: 499 pp. 

Merle, D. 2001. The spinil cords and the internal denticles of 
the outer lip of the Muricidae; terminolog\' and method- 
ological comments. Novapex 2 (3): 69-91. 

Schepman, M. .M., 1892. Note VI. Two supposed new species 
ot Pcntailacti/his. Notes from the Levden Museum, Vol. 
XV: 103-104. 

Sharabati, D. 1984. Red Sea Shells, KPL London: 128 pp. 

Singer, B, S. and H. K, Mienis. 1991a. Shells of the Red Sea. 
The famih- Thaidididae (sic) (1). La Conchiglia 27(260): 

Singer, B. S. and H. K. Mienis. 1991b. Shells of the Red Sea. 
" Tlie family Thaididae (II). La ConchigUa 27(261): 54-60. 

Smythe, K. 1982. Seashells of the Arabian Gulf George Allen 
& Unwin, London: 122 pp. 

Tan, K-S. 1995. Taxonomy of Tliais and Manila (MoUusca: 
Gastropoda: Muricidae) in Singapore and \icinit)'. Ph.D. 
thesis. National Phiiversitv of Singapore. 

Verbinnen, G. and Dirkx, M. 2000. Red Sea Mollusca, Part 6. 
Gloria Maris 38(4-5): 64-76. 

THE NAUTILUS 122(2):107-l 14, 2()()S 

Page 107 

Tw^o new deep-sea muricids (Gastropoda) from Argentina 

Guido Pastorino 

Museo Ari^entino de Ciencias Naturales 
A\-. Ana;erG;illardo 470, 3° piso lab. 57 
C1405bjR Buenos .\ires, ARGENTINA 

Fabrizio Scarabino 

Direccion Nacional de Recursos Acuatico.s 

and Museo Nacional de Ilistoria Natural y 


c. c. 399'- C. P. li.OOO 

Montevideo, URUGUAY 



Two new" species of muricids belonging in the genus TropJton 
are described from the upper slope off the Atlantic coast of 
.\rgentina. Bodi species ha\e a small size tor the genus. The 
radulae show similarities with those of Antarctic species of die 
same genus. Troplion columbarioides new species has a smooth 
shell widi spines pointed adapically and was collected at 37-38° 
S, in 209-382 m. Trophon fasciolarioides new species has 
prominent spiral cords and was collected at Bunvood Bank in 
286-292 m depth and off Baln'a Blanca in ca. 1000 m depth. 

Additional Keywords: MoUusca, Muricidae, TropJioiu South- 
western Adantic, Gastropods, Ta.\onomv, Patagonia 


The genus Troplion include.s a group of species of un- 
doubtedl\" austral origin. The older species can be traced 
as far back as the Oligocene, fi'oni Patagonian deposits 
(Griffin and Pastorino, 200.5). Pastorino (2005) re- 
described all known living species of Trophon from 
southern Soutli America, and also some new species be- 
longing in the genus. \\'lule die subtamilial affinities of 
diis genus are discussed bv several authors (e.g. Kool, 
1993, 1993a; \'ermeij and Carlson, 2000, among others), 
die genus is firmK- established for all fusiform and/or 
lamellate gastropods — usually with spiral ornamenta- 
tion — from die southwestern Atlantic. Two of the species 
characterize the shallow waters along the Patagonian 
coast, i.e., Troplion geversianus and T. plicatiis. Both are 
SMiipatric in the southern part of Argentina; however 
onlv r. geversianus could be collected intertidally as far 
North as Buenos Aires province. AH the other species are 
niostlv subtidal. Trophon species, as far as it is known, 
are all predators, feeding on the mussel banks and bar- 
nacles that are fairlv common along the coast of the 
soutliem part of South America. 

Living in moderately deep waters are Trophon acan- 
thodes \\'atson, 1882, and the recently rediscovered T. 
clenclu (Carcelles. 1953), a rare lamellose and spinv spe- 
cies originaUv assigned to the genus Miirex (Pastorino, 
2005). Bodi of them undoubtedly belong in Trophon. In 

addition, Houart (1991) and Pastoiino (1999) described 
Trophon miicrone and T. veronicae from deep waters off 
South America. The two species were included in Tro- 
phon despite some minor but distinctive differences in 
radulae, protoconchs and penises. Recentlv Houart 
(2003) and Houart and Sellanes (2006) described new 
species from deep waters off Chile. 

In this paper two new deep water species from the 
southwestern Atlantic are described and compared with 
the other related ones living around the region. 


Specimens of T. colninhtirioidcs new species studied 
herein were collected by one of us (FS) on board the 
Umguayan IW Aldebaran. The other specimens are 
from the 2002 cniise to Antarctica of the German IW 
POLARSTERN. Additional material was collected by Uiii- 
guayan fishing boats. Dissections were performed on 
these ethanol-preseived specimens to study radulae and 
male reproductive system when it was available. Radulae 
were cleaned v\itli commercial bleach and ultrasound, 
and obsened using a Philips XL 30 scanning electron 
microscope (SEM) at the Museo Argentino de Ciencias 
Naturales (MACN). Critical point dning of the penises 
(when available) was prepared at the MACN. Radular 
terminology follows Kool (1993: fig. fiB). Shell photo- 
graphs were taken using a digital camera. A[\ images 
were chgitallv^ processed. The material is housed at the 
MACN and the Museo Nacional de Ilistoria Natural y 
Antropologi'a, Montevideo (MNHNM). 


Class Gastropoda Cuvier, 1797 
Order Neogastropoda Wenz, 1938 
Family Muricidae Rafinesque, 1815 
Subfamily Trophoninae Cossniann, 1903 
Genus Trophon Montlort, 1810 

Type Species: Mun 

Trophon gevcrsianns ( 


'IS Gmelin, 
liv orifrinal 

1791, = 

Page 108 

THE NAUTILUS, Vol. 122, No. 

Trophon columharioides new species 
(Figures 1-11. 15-18) 

Diagnosis: Shell veiy small, thin, iusilorm, chalky; 
axial ornanientation of 30-35 regular axial lamellae grow- 
ing along the entire whorl surface, attached to the shell, 
producing open, long, regularly spaced spines along pe- 
riphen', pointing up\\'ard.s. Siphonal canal vei-y long. 

Description: Shell small (up to 14 mm), slender, thin 
profile, chalk)', white, somewhat bright; protocoiich of 
two whorls, slightly globose, symmetrical; teleoconch of 
five oblifpe, shouldered whorls; spire height less than Vi 
of total shell height. Spire angle about 40°; suture im- 
pressed; subsutural shelf well defined, oblique; aperture 
semicircular, interior chalky white; anterior siphonal ca- 
nal veiy long (longer than aperture height), narrow, 
straight, open; outer lip shaip, rounded, inner lip ad- 
pressed. Axial ornamentation of 30-35 regular axial 
lamellae growing along the entire whorl surface, but at- 
tached to shell and producing open, long, regularly 
spaced spines along peripheiy (almost in the middle of 
the whorl), pointing adapically; last three whorls \vith ten 
lamellae each . A second obsolete series of spines ap- 
pears to rise at the peripheiy of older specimens (Figure 
11). Spiral ornamentation lacking. Growth lines present 
throughout shell. 

Operculum subtriangular, nucleus terminal. External 
surface covered by concentric, irregular, growi:h lines. 
Inner surface attachment area reaching upper side or 
center, with horseshoe-sliape scars (Figures 17-lS). 

Rachidian teeth of radula with thin, small central cusp; 
lateral cusps wider and larger than central cusp; deiiticle 
between central and lateral cusp lising from base. Base 
of rachidian tooth sliglitly cnn'etl. Lateral teeth with 
single, long cusps; attachment area thick (Figures 15- 


Type Material: Holotype MACN-In 373S0 (Figures 
1-3, 7, 9-10) ;md t\vo paratyi^es, MACN-In 373S1 (Fig- 
ures 4-6, 8) and MNHNM'15540 (Figure 11). 

Tyjje Locality: RA' Aldebaran cniise 200,3/01, sta- 
tion 37, 37^43' S, 55^00' W, 209 m, October 26 2003, 
5.3°C of bottom temperature, Piccard dredge; (holotype 
and one paratype); bet\veen 37°05' S, 54°12' W in 255 m 
and 37°02' S,'54°02' W in 382 m (one paratype). 

Etymology: The general shell morphologv' reminds 
some species of the genus Columbarium (Caenogas- 
tropoda: Turbinellidae), to which it has no close relation- 

Distribution: Known onlv from three specimens from 
the type localit)' and vicinity. 

Remarks: At first glance the general moiphology of 
the shell shows some similarities with juveniles of Tro- 
phon acanthodes Watson, 1SS2. However the typical 
spiral cords of the latter appear early in ontogeny (see 
Figures 12-13) and are completely absent in the new 
species. In addition b(jth species have clear radidar dif- 
ferences (see Pastorino, 2005:69). In addition, T. plicatus 
and T. cicnclu are comparable species. The latter has 
unmistakable early developed spiral ornamentation; the 
former has complete lamellae and a shorter siphonal ca- 
nal, besides radular differences. 

Despite the geographic distance separating them, Tro- 
plion scolopax and T. septus described by Watson (1882) 
are comparable species. They live around Kerguelen Is., 
in the southernmost Indian Ocean. As a main difference 
Trophon cohiinbarioides new species has a higher spire 
and only one series of large and open spines pointing 
upwards, instead of the three series shown by T. scolopax 
or the triangular upturned ones of T. septus. 

Trophon echinatus (Kiener, 1840), an extremely vari- 
able species (according to Bouchet and Waren, 1985: 
141), from deep waters off Northeastern Atlantic and 
Meditei'ranean shows a remarkable shell siniilarih' with 
T. cohiinbarioides new species Despite this, we do not 
support phvlogenetic affinities between these and we 
made the comparison just for showing the existence of 
specific contrasting differences, particularK' at radular 
level. As far as we can see in the material available there 
are no signs of spiral cords in the new species which are 
common in the deeper specimens of the northern one. 
However, some porcelkmeous (not chalky) smooth speci- 
mens are in fact comparable. All moiphs of T. ecliinatus 
have a shorter protoconch. Some grown specimens of T. 
cohimbarioides shows the apparently presence of a sec- 
ond rows of spines while T. echinatus has only one. In 
addition, the radulae (illustrated bv Bouchet and Waren, 
1985 figs. 333, 335, 336) shows a pair of almost obsolete 
intermediate denticles between the lateral cusps while in 
T. cohimbarioides new species they are slightly smaller 
than the central and lateral cusps. The base of the rachid- 
ian is sinuous in T. ecliinatus and somewhat straight in 
tlie new species. The intermediate denticles of the 
rachidian teeth rise from the internal side of the lateral 
cusps in a wa\' that is t\pical of the southwestern Atlantic 
Troplion species. The attached portion of the marginal 
teeth are also different. 

Houart (2001) considered T. echinatus as belonging in 
to the genus Pagodida Monterosato, 1884 despite the 
chfferences that Bouchet and Waren (1985) pointed out 
with the protoconch of the type species oi Pagochda: the 
Pleistocene species Murex vaginatus Cristofori and Jan, 

Figures 1-11. Trii])lum cohiinhfiritnilcs new species . 1-3. Holotyjje, MACN-In 37380, IW Aldebakan cruise 2003/01, station 37, 
37°43' S, 55°00' W in 209 m. 4-6. Paratype, MACN-In 37381. Same locality as hoiotyise. 7. Apical view of the holotype. S. Apical 
view of the paratype. 9-10. Protoconch of the holot)pe, scale bar = 1 mm. 11. Paratype, MNMNM 15540, between 37°05' S, 54°12' 
VV in 2.55 m and .37°02' S, 54°02' W in 382 ni. 12-14. Tropluni ,i,;nilli<«lrs Watson', 1882, MACN-In 25165-2, 37°35' S, 54°55' W. 
102 III. Sc'aie liar = 1 cm lor all lignres except 9-10. 

G. Pastorino and F. Scarabino, 2008 

Pas;e 109 

Page 110 

THE NAUTILUS, Vol. 122. No. 

Figures 15-18. 'frojilioii ctiliiiiilxiiioUlc.s new specifs. 15. Kaihila Iroiital \ie\\', .scale bar = 1(1 |xiii. 16. Kadiila lattaal xicw, scale 
bar= 20 |xiii. 17. Operciiliiin ol the holohpe and, 18. Parahpe in tigs. 4-6. Scale bars= 1 mm. 

Unlortunately both dissected specimens ot the new 
species here described were females so nothing can be 
said about the moq3llolog^' of the penises. 

Truphon fasciolaiioicles new species 
(Figures 19-3]) 

Diagnosis: sliell \'eiA small, thiji, transiiicid or chalk\'; 
very weakly developed axial ornamentation oi regular, 
veiy low, varices attached to the shell. Spiral oi'namen- 
tation of 2, 3 and 8 or 9 spiral rounded cords in the 
second, third and last whorl respectively; cords of similar 
width; intersection of spiral cords with axial varices con- 
veying a slightly cancellate appearance to shell surface. 

Description: shell small (up to 12 mm), slender, tliiu 
prolile, translucid white or chalky; pi'otoconch ol 2 
whorls, symmetrical; teleoconch of 4'/2 tabular whorls; 
spire height less than '/-> of tfital shell height. Spire angle 
about 45°; suture impressed, subsutural shelf very 

oblique; aperture suboval; anterior siphonal canal long 
but never longer than aperture height, slightly curved 
adaxiallv, open; outer lip shaip, rounded, inner lip ad- 
pressed. Axial ornamentation ol poorl)' developed va- 
rices, regular, veiy low, attached to the shell, growing 
along the entire whorl surface except the siphonal canal. 
Spiral ornamentation ol 2, 3 and 8 or 9 spiral rounded 
cords in the second, third and last whorl respectixeK; 
cords of similar width; intersection ot spiral cords with 
axial varices conveying a slightly cancellate appearance to 
shell surface; growth lines present dn'oughout shell, be- 
coming scaly at intersection with spiral cords. 

Operculum suboval, nucleus terminal. External sur- 
face covered by concentric, irregular, extremely thin 
growth lines. Inni'r surhice attachment area reaching up- 
per side or centej', witli horseshoe-sliaped scars (Figure 

Rachidian teeth ol ladula with lliin central cusp, 
hitrlier tlian laterals; denticle bt.'tween centi;il ;ind lateral 

G. Pastorino and F. Scarabincx 2008 

Page 111 

Figures 19-26. Troplio}i ju.iLUihirioides^ new species. 19-21. llolutvpe MACN-IN 373S2, Banco Bunvoud, Pularstern St. 150, 
beh\-een .54=.3().22' S. .56°0S'5S' \\' in 286 m and 54°29.64' S 56°08.09' W in 292 m. 22. Parahpe MACX-IN .3738.3. same localih- 
as holohpe. 23-24. Protoconch of the paratype of figure 22, arrow head transition to teleoconch, scale bar = 1 )iiiii. 25-26. Panit\pe 
MACX-In 37384. off Baliia Blanca ca. 1000 m depth. Scale bar = 5 mm for all figures except 23 and 24. 

cusp tliin and long, rising from the base. Base of rachid- 
ian tooth cur\'ed. Marginal area with single cusp. Lateral 
teetli with single, long cusps; attachment area thick (Fig- 
ures 27-29). 

The penis shows an unusual moi"phologv' among the 
Patagonian species of the genus: it is wide, slightly 
cun-ed and lateralh* flattened with a small slit at the tip 
(Figure .30). 

T\-pe Material: Holot\pe MACN-IN 37.382 and two 
paratopes MACN-IN 37383 and MACN-In 37384. 

Type Locality: Banco Buiwood, Polarstern station 
150, 6 Apr. 2002, ACT net, between 54°30.22' S, 
56°08.5S' W in 286 m and 54°29.64' S, 56°08.09' W in 
292 m. 


some spec' 


shell moi"phology reminds 

les of the genus FascioJaria (Caenogastropoda: 

Distribution: known from tvpe localit\' and appro.xi- 
matelv off Bahia Blanca in ca. 1,000 m depth (MACN-ln 

Page 112 

THE NAUTILUS, Vol. 122, No. 2 

Figures 27-31. Troplion fasciolarioUles new speeie.s. 27. Kadiila ot a paratvpe, MACN-In 37384, liontal view, scale bar = 40 |JLm. 
28. Lateral view of the same radula as 27, scale bar = 50 [jliii. 29. Radula of the parabi^pe MACN-In 37383, scale bar = 20 ixm. 30. 
Critical point dn' o( the penis, scale bar = 500 jxin. 31. Two \iews oi die opcrculnm of the parat\pe of figures 25-26, scale bar = 
1 mm. 

Remark.s: The iiioqDliolog)' of the shell and the radula 
match some oi the Patagonian and Antarctic species of 
the genus Trophon. From the first group, T. ohlini 
Strebel, 1904, a rare species from the Magellanic area, 
has a similar profile. However, its distinctive protoconch 
points out a clear difference. In addition, the moiphology 
of the penis is far from the ty[:)ical Patagonian represen- 
tatives of the genus. Trophon cmihjae Pastorino, 2002, 7". 
cleclinans Watson, 1882, and T. ciispkkirioicles Powell, 
1951, are comparable Antarctic species (Pastorino, 
2002a). The first two species differ from the new species 
in having well developed axial sculptm-e only. Also T. 
cuspiclarioides has 5V2 whorls including the protoconch 
and five blunt spiral cords in the body whorl, while the 
new .species has a larger spire and 6V2 whorls, and 8 or 9 

rounded and well defined cords in the last whorl. The 
a\ial ornamentation in the new species is very irregular 
but higher in number than the 25 axials present in T. 



In previous papers two groups of species belonging in the 
genus Trophon — i.e., from South America and Antarc- 
tica — were pointed out. The main differences between 
these two groups are several radular and anatomical fea- 
tures (Pastorino, 2002b, 2005). The shell morpholog\- of 
the two new species described herein agrees with that of 
Trophiiii living off the South American coast. However, 

G. Pastorino and F. Scaiabino, 200S 

Pasre 113 

Buenos Aires ' 

Bahia Blanca 




Malvinas Is. 

Staten Is. 

Figure 32. Map showing the t\pe locaUties of Troplion 
cohimbaiioidcs new species • (filled circle) and Truphon fa.s- 
ciolarioides new species Q (blank square). 

the radula has some features found in the Antarctic rep- 
resentati\'es of tlie genus — albeit the taxonomic status of 
tlie .\ntarctic species still needs confirmation. Like in the 
Antarctic group of species, the central cusp of the rachid- 
ian teeth is shorter and thinner than the laterals, the 
internal denticle rise from the base of the rachidian in- 
stead of the upper third of the internal side of the lateral 
cusp, and the lateral teeth have a wide attachment area. 
The geographic ranges of species of these tvv-o groups 
do not o\"erlap. Therefore, the moiphological features 
seem to be clearh- separated geographicafly too. How- 
ever, in the species described herein, this seems not to be 
die case. W'liile diere is no geographic overlapping of 
species, the morphological features that characterize the 
Antarctic group do appear in these Uvo new Patagonian 
species. Contrarily, diis does not occur in the case of 

Patagonian species, the moii^hological features of wliich 
are restricted to South America. Moreoxer. thev are bet- 
ter de\eloped in species from the Vhigellanic among 
those described from southern South America (see Pas- 
torino, 2005). 

Recently, Houart {2()()3) introduced three new species 
lie assigned to Troplion from off Chile: T. ceciliac; T. 
condei and T. vaugoctJjemi. The morphology of the 
radula, only known for the last hvo species, together with 
that of T. mucrone Houart, 1991, T. veronicae Pastorino, 
1999 and both new species described here, is that of the 
Antarctic group, despite the geographic distance sepa- 
rating them. All mentioned species were collected from 
more than 300 m deptli and in most cases they reach 
more than 1,000 m (see Table 1). Most of the Patagonian 
species of Troplion were collected alive from the inter- 
tidal zone to about 300 m depth. Exceptions are some 
specimens of T. acanthodes, T. clenchi. and T. hahmnon- 
dei that come from deeper areas. Howe\'er, there is no 
Antarctic species — or with their characteristic radular 
and anatomical features — that li\e at shallower depths or 
closer to the continent. Both new species described here 
were collected from about 300 m depth, which is the 
shallowest for a Trophon with Antarctic features at South 
American latitudes. 

Generic assignment in the whole Trophoninae group 
is actually under revision. The concept of the genus Tro- 
phon sensu stricto is easy to apply to the shelf species. 
However, when deeper species are studied, the presence 
of several features in common among the shallower spe- 
cies appear somewhat contradictoiy. Nevertheless, we 
think that biogeographic boundaries are hard to suipass 
for species without free lanae as both new species ap- 
pears to be according to the shape and whorl number of 
the protoconch. Contrasting historical biogeographic 
processes must be considered when suggesting affinities 
between species and dierefore comparison with species 
lixdng far away from the Southwestern Atlantic area can 
be considered an empty exercise. Some exempli are cases 
like T. coJnmbarioides new species — T. ccliinatus which 
have \en' few characters that clearK' spht both species. In 

Table 1. Recently described Soutii American sj:ecies ofTrophDii. 

Depth (m) 

Tvpe localitv 

T. coiuhi Houart, 


Ancud, Chile 


T. ceciliac Houart. 

434-100(1, 1300 

Antofagasta. Chile 


T. vangoetheini 

About 350 

Itata, North of 

Houart, 2003 

Concepci'on, Chile 

T. mucrone Houart. 


Off Rio de 


Janeiro. Brazil 

T. veronicae Pastorino, 




T. coluinhinioides 


37°43'S, 55°00'W 

new species 

T. fasciolarioides 


Buiwood bank 

new species 

Paee 114 

THE NAUTILUS, \ ol. 122, \o. 2 

the future molecular characters could add to the under- 
standing of the evolution ot moiphological features in 
this group. 


We are grateful to L. Paesch and R. Bird (DINARA) 
for facilities in the collecting of Trophon specimens, as 
well as to y. L. Viggiano and H. Racz-Lorenz (Monte- 
video) for the material provided. We thank Miguel Grif- 
fin (UNLPam) who helped to impro\'e the English ver- 
sion of the manuscript. R. Houart (Belgium) re\iewed 
this paper; his comments and discussions undoubtedly 
contributed to enhance the original \'ersion despite dis- 
agreement on some points. This contribution was sup- 
ported in part by the project PICT No. 03-14419 from 
the National Agency for Scientific and Technical Promo- 
tion, Argentina. We acknowledge funding by the Consejo 
Nacional de Investigaciones Cientificas y Tecnicas 
(CONICET) of Argentina, \\'hich sponsors the research 
of G.P. 


Bouchet, P. and A. Waren. 1985. Rexasion of the northeast 
Atlantic bathval and abyssal Neogastropoda e.xcluding Tiir- 
rklae (Mollusca, Gastropoda). Bolletino Malacologico 
supplemento 2: 123-296. 

Carcelles, A. 1953. Nuevas especies de gastropodos niarinos de 
las Repiiblicas Oriental del Uruguay y Argentina. Comu- 
nicaciones Zoologicas del Museo de Historia Natural de 
Montexldeo, 4(70): I-LS. 

Griffin, M. and G. Pastorino. 2005. The genus Troj)lion Mont- 
fort, 1810 (Gastropoda: Muricidae) in the Tertian- of Pa- 
tagonia. Journal of PaleontoIog\' 79: 296-311. 

Houart, R. 1991. The southeastern Brazilian Muricidae col- 
lected by R V Marion-Dufresne in 1987, widi die descrip- 
tion of three new species. The Nautilus 105: 26-37. 

Houart, R. 2001. Ingensia gen. no\'. and efe\en new species of 

Muricidae (Gastropoda) from New Cafedonia, Vanuatu, 
and W'allis and Futuna Islands. In: P. Bouchet and B. A. 
Marshall (eds.) Tropical Deep-Sea Benthos, volume 22. 
Memoires du Museum national d'Histoire naturelle 185; 

Houart, R. 2003. Description of three new species of Trophon 
s. 1. Montfort, 18f0 (Gastropoda: Muricidae) from Chile. 
Novape.x 4; 101-110. 

Houart, R. and J. Sellanes. 2006. New data on recently de- 
scribed Chilean trophonines (Gastropoda: Muricidae), de- 
scription of a new species and notes of their occurrence at 
a cold seep site. Zootaxa 1222; 53-68. 

Kool, S. P. 1993. Phvlogenetic analysis of the Rapaninae (Neo- 
gastropoda: Muricidae): Malacologia 35: 155-260. 

Kool, S. P. 1993a. The systematic position of the genus NticeUa 
(Prosobranchia: Muricidae: Ocenebrinae): The Nautilus 
107: 43-57. 

Monterosato, A. di, 1884. Nomenclatura generica e specifica di 
alcune conchiglie Mediterranee. Palermo, 152 pp. 

Pastorino, G. 1999. A ne\v species of gastropod of the genus 
Troplion Montfort, 1810 (.Mollusca; Gastropoda; Muri- 
cidae) from sufiantarctic waters. The Veliger 42; 169-174. 

Pastorino, G. 2002a. Two new Trophoninae (Gastropoda; Mu- 
licidae) from Antarctic waters. Malacologia 44: 353-361. 

Pastorino, G. 2002b. S\stematics and phvlogen\- of the genus 
Troplion Montfort, 1810 (Gastropoda; Muricidae) from 
Patagonia and Antarctica; moiphological patterns: BoUet- 
tino Malacologico 38; 127-134. 

Pastorino, G. 2005. A re\ision of tfie genus Trojihon Montfort, 
1810 (Mollusca; Muricidae) from southern South 
America: The Nautilus, 119: 55-82. 

Powell, A. W. B. 1951. Antarctic and subantarctic mollusca; 
Pelec\poda and Gastropoda. Discoven' Reports, 26; 47- 
196. ' 

Vermeij- G. and S. Carlson. 2000. The muricid gastropod sub- 
famih' Rapaninae: pli\logen\' and ecologiciil liiston'. Pa- 
leobiolog)' 26: 19-46. 

Watson. R. B. 1882. Mollusca of 11. M. S. Challenger" Expedi- 
tion. Part 13. The Journal of The Lmnean Societ\'. Zool- 
otry, 16; 358-392. 

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Vohiinc 122, Nuntbcr 3 

Sepfcmbcr 26, 200H 

ISSN 0028-1344 

LouElla R. Saul Cretaceous trichotropid g;astrop(i(ls from the Pacific slope of Noitli 
Richard L. Squires America: Possible pathways to cal)ptraei(l moqjhology 115 

Samuel S. Espino Feeding behavior, phylogenv, and toxinologx' ni Conin fniTus Reeve, 

.-Vlau J. Kohn 1843 (Gastropoda: Neogastropoda: Conidae) 143 

James A. \'illanue>a 
Frank M. Herakle III 
Patrice Showers ComeH 
Gisehi P. Concepcion 
Baldomero M. ()U\era 

Ridiard Ducrr Two new species oi Mit reiki (Gastropoda: Neogastropoda: 

Cokimbellidae) from the lower Miocene Chipola Formation of 
nortliwestern Florida 151 

Fresia ViUalobos-Rojas Catalogue of the tvpe material ol mdllusks deposited at the Zoologv' 

.\na. G. Guzman-Mora Museum. Universih' of Costa Rica 155 

Yohinda E. Camacho-Garcia 

Takenori Sasaki Dilemma jtiponieum new species (BivaKaa: Anomalodesmata: 

Jose H. Leal Poromyidae): A new record oi the genus trom the 

Northwest Pacific 166 

^'ictor Scarabino On the genus Heteroschismoides Ludbrook, 1960 (Scaphopoda: 

Carlos Henrique Soares Caetano Gadilida: Entalinidae), with descriptions ot two new species 171 

Norma Emilia Gonzalez-Vallejo Parasitism of Monogamii'i minihitlhi (Olsson and McGintw 1958) 

(Gastropoda: Eulimidae) on the red sea-urchin Echbwmetm lueiinter 
(Linnaeus, 1758) (Echinodermata: Ecliinometridae) on the Caillibean 
coast of Mexico 1 78 

Erratum 182 

Notices 183 

5^iBLWH0! Library 



Massachusetts 025^3 

THE NAUTILUS 122(3): 115-142, 20()S 

Page 115 

Cretaceous trichotropid gastropods from the Pacific slope of 
North America: Possible pathways to calyptraeid moi^phology 

LouElla R. Saul 

Invertebrate P;ileontolog\' Section 

Natural Histor\' Museum of 

Los Angeles Count^• 

900 Exposition BouIe\ard, 

Los Angeles. CA 90007 USA 

Richard L. Squires 

Department of Geological Sciences 
California State University, 
Northridge, CA 91330-8266 USA 


Late Cretaceous gastropods belonging to genus Lysis Gabb, 
1864, from die Pacific slope of North America, bridge die mor- 
phologic gap bet\veen turbinitorm trichotropids and limpet-like 
cal\ptraeids. De\elopment oi the depressed and broadened 
inner lip/columella of Lysis resulted in a larger aperture diat 
allowed more space for die foot to grasp a hard substrate. 
Pacific slope species of Lysis are represented by five species 
that collectively span an interval from late Coniaeian to late 
Maastrichtian. They stem from two lineages of the trichotro- 
pine genus Ariadnaria Habe, 1961. The first Hneage, which 
includes Ariadnaria ainikta new species of late Albian to Cen- 
omanian age, Ariadnaria stibara new species of Cenomanian 
age, and Ariadnaria obstricta (White, 1889) of late Coniaeian? 
and Santonian age, gave rise to die Lysis diiplicosta group of 
neritiform to hahotiform, coarse-ribbed Lysis, including Lysis 
mickeyi new species (earliest Lysis in the world), Lysis diipli- 
costa Gabb, 1864, Lysis jalanmca new species, and Lysis lo- 
maensis new species. The second lineage o{ Ariadnaria consists 
of the Turonian Ariadnaria aldersoni new species, which gave 
rise to die Lysis siiciensis (Whiteaves, 1879) group. Morpho- 
logicallw this group, which show crepiduliform and fine ribbed 
shells, appears Ukel\' to be a stem group from which Cenozoic 
Crepidula-hke genera evolved. Garzasia new genus, which 
ranges from late Campanian or early Maastrichtian age to the 
mid Maastrichtian, evolved from the Lysis duplicosta group 
and includes Garzasia intermedia (Cooper, 1894) and Garzasia 
diabla new species. The very broad, depressed spiraling inner 
hp of Garzasia is suggestive of Calyptraea Lamarck, 1799. We 
propose placement of Lysis and Garzasia in Lysinae new sub- 
famil\' of the Trichotropidae. In addition to their occurrence 
along the Pacific Slope of North .A.merica, Lysis or Lysis-like 
gastropods are known from middle Santonian to lower Cam- 
panian strata in Soudi Africa, upper Campanian in the Congo, 
and Maastrichtian strata in Mozambique and Japan. 

Additional Keywords: Trichotropidae, Lysinae, Calyptraeidae, 
evolution, paleogeographical occurrence 


This stud\- deals with the fossil record of the e.xtinct ge- 
nus Lysis Gabb, 1864, a small group of enigmatic gas- 

tropods which has received little or no study regarding its 
ancestry, point of origin in terms of time and geographic 
locale, and evolutionary histoi"y. Specimens have been 
found in shallow-marine Cretaceous deposits from 
southern Vancouver Island and neighboring Gulf Is- 
lands, British Columbia, Canada to northern Baja Cali- 
fornia, Mexico (Figure 1) and, ahhough Lijsis-like gas- 
tropods have been reported at a few locales in die world, 
its familial placement has been uncertain. This study 
brings neu' information about all of these items. 

Material for this study included t)pe specimens, addi- 
tional collections from their type localities, and speci- 
mens of Late Cretaceous age (Coniaeian to Maastrich- 
tian) from previously unreported-upon localities. Figure 
1 provides an index to areas which Yielded specimens 
used in this study. 

During the study we discovered undescribed species 
of the trichotropine genus Ariadnaria Habe, 1961, which 
appear to have given rise to two groups of Li/sis. a ner- 
itiform coarse-ribbed group and a crepiduliform fine- 
ribbed group. As will be discussed under "Evolutionary 
Implications," we propose that the neritiform group 
evolved into Garzasia new genus, which appears to be a 
precursor to caKptraeiform genera. 

A total of three genera (one of them new) and 11 
species (seven of them new) make up this study. The taxa 
are: four species of Ariadnaria (three of them neyv), five 
species of Lysis (three of them new), and two species of 
Garzasia (one of them new). The ranges in time of all 
these species, as well as two recognizable groups of Lysis, 
are plotted on Figure 2. The Lysis duplicosta group con- 
sists of four species, spanning a total inteival of late Co- 
niaeian to late Maastrichtian. This group was also the 
most geographically widespread, with specimens col- 
lected from British Columbia to Baja Cahfoniia. The 
Lysis suciensis group is known only from the Campanian 
to possibly early Maastrichtian, with specimens knowai 
from British Columbia to Baja California Sur. Lysis per- 
sisted for a total of approximately 18 million years in the 
study area. 

The specimens studied here are mainlv from fine- 

Page 116 

THE NAUTILUS, Vol. 122, No. 3 


--vV "^^ ' 


1 -V:uicoLi\'er and adjacait 

13 Agua CaUeute Canyon. 

islands, Britisli Colimibia 
.', - Siicia Island, San Juan Co., 

Santa Barlaara Co., CA 
14 - Warm Springs Mtn., 

1. !• L^ 


3 - Soutli Cow & Bear creeks. 

Los Aigdes Co., CaUf 
5 - Live Oak Canyon, 


^ ' 1 JH) 

Shasta Co., CaHfomia 


. 1 


Kem Co., CaUfomia 

4 ~ Ono area, Sliasta Co., Calif. 

2« 2 

6 - Simi FBUs, Los Angeles 

■ I .W 

5 - Cliico Creek, Butte Co., 


and Ventiu'a counties. 



6 - Pentz and Dry Creek, Butte 

17 - Santa Mcaiica Mtns, | 

Co., CaUfomia 

Los Angeles and 

7 - Texas Flat & Gnuiite Bay, 
Placer Co., California 

Vaitiu-a comities, CaUf. 
IS - Saitta Ana Mtns., 

~. r -/ ^^ — ' ~~~-«.^ 

1 I , ^ 


S - Pigeon Point, San Mateo Co., 

9 - Garcas Creek, Stimislaiis Co., 

Oi-ange Co., California 
1 9 - near Carlsbad, San 
Diego Co.. CaUfomia 



- Point Loma, San 

/ 1 

10 -soutli of Garzas Creek, 

Diego Co., CaUfomia 

' f-"^ ' 1 

Mared Co., CaUfomia 

21 - East of La Mision, | 

1 Q 1 ^^*—**..,.,^^ ' 1 

1 1 -Cooper Canyon, Fresno 

Baja California, Mexico 1 

Co., CaUfomia 

22 - Ciuion San Fernando, | 

y , 1 -~--_L^ 

12 -Jalama area, Santa Barbara 

Baja California, Mexico 

Co., CaUfomia 

23 - Pttita Abreojos, Baja | 

c. _ 

California Siu; Mexico 

g 5» 5» 


I 7 


sJ^^Vio /^ \ 

?W' ^ \ 

^ ^ -§ 







12j»^#l4i^^ r \} 



' k^^f \ :J 




'0 15^ 

* 16( 

1 ^2 
» 16 


J 14. 


20*_ [ > ( 




• ^3« 






. 20» 





1^ "S 

1 p 

r"----r4 u 

1 .s 

^ I % \ 



^ ^^' 



» 22 


/\ iX^ 

'^i ) y 


1 f\fj^^^ 

50 100 250 SOOknrC' ''^^Z/^'/'N 

III 1 =f 2i^-(''\ 



Figure 1. Index map ol collecting localities. 

L. R. Saul 

R. L. S 



Page 117 









75 — I 

80 — 

85 — 

90 — 


100 — 





125 — 























58 7 


61 7 



70,6 — 















Figure 2. Age ranges of species oi Ariadnaria. Li/sis, Gai~asia, Cahjptraea, and Crepichda discussed in text. Lines connecting the 
species indicate inferred descent based on nioiphologic simiarit)--. Lijsis duplicosta gave rise to two species one of which, L. lomaeiisis, 
produced proto-calvptraefomi Gai-asia and the other, L. jalamaca, is crepiduhforni. An additional crepiduliform line was founded 
bv L. suciensis. No knowai intermediate forms connect these Cretaceous calyptraeiform and crepiduliform gastropods to Tertiaiy 
species. The earhest knowna Pacific Slope cahptraeid, 'C diegoana lacks the strong ribbing oi Garzasia, but crepiduliform Spiro- 
cnjpta pileum differs largely from L. suciensis in its small size. Time scale after Gradstein et al. (2004). 

Page 118 

THE NAUTILUS, Vol. 122, No. 3 

grained argillaceous sandstone or siltstone that constitute 
shelfal deposits that accumulated at depths near or just 
below wave base. 

Specimens are low in number and almost always in- 
complete. Protoconchs are rarely preserved, the lai-val 
shell is in part missing, its shape partially represented by 
its inner cast in all available specimens. Apertures are 
usuallv missing their anteriormost area. Adliering matri.x, 
commonly consisting ot well-cemented sandstone usuallv 
plugs the aperture, thereby necessitating caretul clean- 

As will he discussed under "Evolutionaiy Implica- 
tions," the shape oi Lysis, with the exception of L. mick- 
eiji new species, approaches that of Crepidida, resulting 
from flattening of the aperture, development of a broad 
shelf and lateral coiling of the spire. The crepiduliform 
shape, as well as the CahjptraeaAike shape oi Gai-asia, 
pose problems for terminology used for describing cer- 
tain figures of specimens, as well as for describing certain 
shell dimensions. For some specimens, a full view ot the 
aperture could onlv be depicted by ha\ing the the shell 
tipped out of the plane of its axis. The tioie height of 
these shells, therefore, is not shown; hence, the "vertical 
dimension" of the view is given instead, and peipendicu- 
lar to it is the "horizontal dimension." In most cases, the 
abapertural view is in the plane of the shell axis; hence, 
the terms "height" and "diameter" applv. 

Abbreviations used in the text are: ANSP: Academy of 
Natural Sciences of Philadelphia; CAS: CaUfornia Acad- 
emy of Sciences, San Francisco; CGS: Cieological Sunev 
of Canada, Ottawa; CIT: CaUtornia Institute of Technol- 
og)', Pasadena (collections now housed at LACMIP); 
CSMB: California State Mining Bureau (specimen at 
CAS); IGM: Instituto de Geologia, Universidad Nacional 
Autonoma de Me.xico; LACMIP: Natural History- Mu- 
seum of Los Angeles County; SDNHM: San Diego 
Natural Historv Museum; UCLA: University' of Califor- 
nia, Los Angeles (collections now at LACMIP); UCMP: 
University of California, Berkeley, Museum of Paleon- 
tology; USGS: United States Geological Suney, Menlo 
Park (collections now housed at UCMP); USNM: Na- 
tional Museum ot Natural Histon', Smithsonian Institu- 


sis are known elsewhere in tlie world. They are: Lysis 
capensis Rennie, 1930, from the middle Santonian to 
lower Campanian of South Africa (Kiel and Bandel, 
2003); Lysis congolensis (Brebion, 19.56) from upper 
Campanian of the Congo, Africa; Lysis africana (Cox, 
1925) from the Maastrichtian (undifferentiated) of Mo- 
zambique; and Lysis izuiniensis Kase, 1990, from the 
earliest Maastrichtian of [apan. 

Rennie (1935) reported Lysis caffra Rennie, 19.35, 
from the Upper Cretaceous near the eastern border of 
the Eastern Cape Pro\ince (Pondoland), southeast Af- 
rica, but its swollen naticiform shape with a very large, 
inflated body whorl and broad, non-depressed inner Hp/ 
columellar region, that appears to have a thin callus, are 
features not found in Lysis. 

Gai~asia ranges from late Campanian or early Maas- 
trichtian to the mid Maastrichtian and is endemic to the 
Pacific slope ot North America. 


Lysis has been found attached to a few specimens of 
large \'olutid gastropods. One specimen o{ Longoconcha 
eumcka Saul and Squires, 2008, from the Point Loma 
Formation near Carlsbad, California has two specimens 
of crepiduliform Lijsis jalamaca new species on its shell, 
near the outer lip margin (Figure 34), as well as an at- 
tachment scar on the abapertural side of the shell. Speci- 
mens of Lysis siicicnsis from the Chatsworth Formation 
near Chatsworth, CaUfornia, although not found in situ 
as are the younger Carlsbad specimens, have aperture 
shapes that could fit on the exterior of a specimen of 
Vohitoderma Gabb, 1877. Indication of such an associa- 
tion is absent prior to middle Campanian. Whether the 
specimens of Lysis species were using dead shells as a 
substrate or had de\'eloped an association with living Vo- 
hitodvrma is undetermined. 

Comparison to probable family members indicates 
that Lysis was a sedentan' faculative ciliaiy feeder. Tri- 
chotropines, as well as calyptraeids, are ciliary feeders 
that ]i\e a sedentan- life on hard substrates and are pro- 
trandrous hermaphrodites among whom brooding their 
young is common (Graham, 1954; Yonge, 1962). These 
mode-of-life characteristics ha\e historicallv been used 
to classify trichotropines as being close to calyptraeids. 

Ariadnaria ranges from late Albian to Recent with its 
earliest appearance in Pacific slope deposits of North 
America, where it is found in strata ranging from late 
Albian to Santonian age. From Japan, Kase (1990) fig- 
ured, as Trichotropis?, a possible Ariadnaria species of 
earliest Maastrichtian age. 

Lysis ranges from the late Coniacian to late Maastri- 
chitan, and that is also its range for the Pacific slope of 
North America. The genus apparently originated in Cali- 
f(5rnia, with five species known from the Pacific slope of 
North America. Four additional probable species of Ly- 


Superfanuh CaKptraeoidea Lamarck, 1809 

Discussion: The taxonomv of calvptraeoids, as with 
most gastropods, was tradition;illv based on shell mor- 
phologv and later modified In' increasing anatomical 
knowledge. The inclusion of patclliform Capulidae 
Fleming, 1822, tnrbiniform Trichotropidae, and coiled 
liiripet-shaped CaKptraeidae in the superfamilv CaKp- 
traeoidea (e.g., Thiele, 1929; Wenz, 1940), has provided 

L. R. Saul and R. L. Squires. 2008 

Pas^e 119 

calvptraeoids a considerable moiphological range and a 
somewhat complicated classification histoiy (Bandel and 
Riedel, 1994). In addition to the above three families, 
Bandel and Riedel (1994), includetl Hipponicidae 
Troschel, 1861, in Ciilvptraeoidea, but Collin (2003: 632) 
rejected hipponieids from a close relationship with ca- 
Kptraeids. Ponder and Waren (1988) and Ponder (1998), 
equated iamih- Capulidae \vith family Trichotropidae 
Gra\', 1850, on anatomical grounds, reducing the in- 
cluded families to two. Capulids, as exemplified by 
Capulus Montfort, 1810, have limpet-shaped shells; tri- 
chotropids, as exemplihed h\ TiicJjotropis Broderip and 
SoNN-erbw 1829, usuallv ha\e coiled shells; and cal\p- 
traeids, as exemplified by Cahjptraca Lamarck, 1799, 
Crepidula Lamarck, 1799, and Cmcibulum Schumacher, 
1817, have limpet-shaped shells with an internal shelf of 
\ariable shape. 

In her anaKsis of calyptraeids, Collin (2003a, 2003b) 
utilized shell characters, anatomical characters, and mo- 
lecular characters. Collin (2003a) argued that although 
the so-called echinospira laiva of Capulus and ofTricho- 
trppis do not appear to be "true" echinospira, the thick- 
ened and elaborate lanal shell oi these two groups is 
clearly different from the simple lai-val shell of e.xtant 
cal\ptraeids, and she (Collin, 2003a, 2003b, 2005) has 
continued to recognize the tamilies Capulidae Fleming, 
1822, CaKptraeidae, and Trichotropidae. Collin (2003a) 
mentioned that despite the detailed studies done on ca- 
lyptraeids, tlieir taxonomy remains contentious and un- 
certain. Because specimens studied here show no resem- 
blance to capulids but do, in pait, resemble trichotro- 
pids, and caKptraeids such as Cahjptraca and Crepidula, 
we foUow Collin in recognizing families Calyptraeidae 
and Trichotropidae. 

FamiK" Trichotropidae Crav, 1850 
Subiamilv Trichotropinae Gray, 1850 

Description: Small to medium sized (usualK* 15 to 25 
mm, but up to 40 mm in height), coiled, high-conic to 
broad, low-conic, or nearlv cap-shaped; spiral sculpture 
usually better developed than coUabral sculpture and 
represented bv distinct cords and/or keels; umbilicus 
broadl\- open to slit-like or completelv closed; aperture 
\ariable in shape, ranging from irregularly triangular and 
broadly oval to elongate-oval; some forms with more or 
less attenutated canal; operculum small, horny; radula 
taenioglossate; periostracum forming combs, bristles, 
spines usualK- in places of intersection of spiral and col- 
lab ral sculpture (from Egorov and Ale.xeyev, 1998). 

Discussion: Trichotropids provide the geologically 
oldest representatives of the three families Trichotropi- 
dae. Capuhdae, and Calyptraeidae. We did not follow 
Ponder and Waren (1988), Ponder (1998), and Bouchet 
and Rocroi (2005) in placing Trichotropidae in Capul- 
idae because trichotropids, such as the high spired Aii- 
adnaria spp. discussed herein, differ distinctly from cap- 
shaped capulids. Collin (200.3b) referred to trichotropids 
plus capulids as die closest outgroup to calyptraeids, sug- 

gesting that she recognized tliese tlnee as separate fami- 
hes. At present the geologic record finds trichotropids 
occurring earlier than capidids and also earlier than ca- 
lyptraeids, and we suggest that based on species de- 
scribed herein, trichotropid-like gastropods gave rise to 
calyptraeids and that trichotropid specimens reported 
upon herein demonstrate a progressive development to- 
ward either a crepiduliform or a calyptraeiform shell. 

Atresius Gabb, 1869, of Early Cretaceous (Valanginien 
to Hauterixdan) age is the earliest trichotropine recog- 
nized by Wenz (1940), but its sole named species A. 
liratufi Gabb, 1869, is a prominent constituent of chemo- 
synthetic paleoconununities in northern California and 
probably not a trichotropid. Lirpsa Stephenson, 1952, of 
Cenomanian age from the Woodbine Formation of Texas 
may be the earliest previously known trichotropine. The 
earliest reported cap-shaped capulid is Capulus venis 
(Bohm, 1885) of Late Cretaceous (early Campanian) age 
from Aachen, Germany. 

Genus Ariadnaria Habe, 1961 

[= Ariadna Fischer, 1864; nan Autk)\nn, 1826]. 

Type Species: Trichotropis borcaUs Broderip and 
Sowerby, 1829, by monot)py; boreal Arctic circumpolar 
(Egorov and Ale.xeyev, 1998) and cool temperate seas: 
northern North Atlantic south to Nortlumibria and all 
Scottish coasts (Fretter and Graham, 1962) and Massa- 
chusetts Bay (Emerson and Jacobson, 1976); the Bering 
Sea and north Pacific south to British Columbia (LACM 
collection, Forrester Island). 

Description: Shell turbiniform (oval-conic) with 
raised spire; spiral sculpture well developed and consist- 
ing of raised cords separated b\' interspaces of variable 
width; collabral sculpture consisting of raised growth 
lines; aperture wide; inner lip slightlv conca\e; canal 
short and straight; umbilicus slit-like; operculum thin; 
periostracum forming long, closely spaced bristles on spi- 
ral ribs. 

Discussion: No prior records of Ariadnaria as a fossil 
were found by us. Ariadnaria differs from Trichotropis 
Broderip and Sowerby, 1829, by having an umbilicus. 
Turbinopsis Conrad, 1860, a turbiniform trichotropid of 
late Campanian or early Maastrichtian age from Missis- 
sippi, differs from Ariadnaria by having a wider umbili- 
cus (profound according to Conrad, 1860), a last whorl 
that is more inflated and is tabulate, and having a very 
oblique fold near the basal margin of the columella. 

Ariadnaria ainikta new species (Figures 3-4) 

Diagnosis: Medium-size Ariachiaiia with sturdv shell, 
rounded whorls, strong sculpture with nine spiral cords 
widely spaced; collabral sculpture thickly foliate; umbili- 

Page 120 

THE NAUTILUS, Vol. 122, No. 3 





9 ^^^^6^" 10 

Figures 3-10. ArkiiUiana species. ,Speciinens coated witli aiiiuiomuiii chloiKle. 3-4. ArkuUiiiria diiiikta new species, holot}pe 
CAS 61794.00, CAS loc. 61794, height 20 mm, diameter 17 mm. 3. Apertural view. 4. Abapertural x-iew. 5-6. Ariadnaiia stibara new 
species, holots'pe LACMIP 13371, LACMIP loc. 23464, height 19 mm, diametei' 13.5 mm. 5. Apertural wew. 6. Abapertural view. 
7-8. Ariadnaiia aldersoni. holotype LACMIP 13372, LACMIP loc. 26370, height 12 mm, diameter 8.5 mm. 7. Apertural view. 8. 
Abapertural \'iew. 9-10. Ariadiiaria obsfricfa (White, 1889), h\pot)pe LACMIP 13373, LACMIP loc. 28717, height 21 mm, diameter 
15 mm. 9. Apertural \ie\v. 10. Abapertural \iew. 

cus elliptical: inner lip broad, expanded anterior and pos- 
terior to nnihilicus. 

Description: Shell mechum (approximatelv 22 mm 
height), sturdy, turbiniform, spire moderately high, ap- 
proximately 36% of total shell height; apical angle 110°; 
most ot protoconch missing, remanent low and appar- 
ently smooth; teleoconch wlioils three, whorls well 
rounded and enlarging ven' lapiillv; sutin"e appressed but 
appearing channeled; umbilicus open, deep, and ellipti- 
cal; growth line prosocline; spiral sculpture consisting of 
regulail)' spaced and ec|uallv narrow coi'tls; loru^ cords on 
penultimate whorl; nine cords on last whorl, becoming 
stronger and more raised near umbilicus; spiial cords on 
last whorl widely spaced and occasionally \vith spiral 
thread in interspaces, especially anterior of peripheiy; 
collabral sculpture consisting of thickly foliate ridges co- 
incident with growth checks, especially near outer lip; 
aperture D-shaped; inner lip broad, expanded anteriorly 
and posteriorly of umbilicus; abapertural edge of inner 
lip delineated by shaip ridge; l)asal lip broadened. 

Holotype: CASC; 61794.00, hciglil 20 mm 
17 mm, spire lieiglit 7.5 mm. 


Type Locality: CASG 

61794 [ = CASG loc. 

Distribution: Basal Bald Hills Member of the Budden 
CauNon Formation, (area 4) Ono area, Shasta Co., Cali- 

Geologic Age: Late Albian. 

Discussion: C)nl\' tlii' holotvpe is known. It evidently 
had a ve\y foliate, thick shell. What remains is riddled 
with endobiont borings. Remnants of the protoconch are 
present, there is no clear e\idence of an anterior sinus to 
the aperture, and the shell does not appear to have been 

The new species most resembles the trichotropid 
Tiirbinopsis lulgarcli Conrad, 1860 (Conrad, 1860: 289, 
pi. 46, fig. 29; Sohl, 1960: 91, pi. 10, figs. 17, 18) from the 
upper Campanian/lower Maastrichtian Ripley Formation 
of Tippah Co., Mississippi, except A. ainikta has a 
smaller umbilicus and narrower cords with much wider 
interspaces. AtlditionalK', the innei' lip of A. ainikta is 
more expanded both iiiitcrior :m(l posterior to the um- 

Ariadiiaria ainikta and .\, \lihara new species are simi- 
kir in that there is no ritlge separating the imibilicus from 
the irmer lip. Ariadiiaria ainikta dillers from A. stibara 
li\ ha\ing ;i less ('longate shell, \\idei' penultinuitt' whorl, 
stionger spiral cords with much wider interspaces, a 
shorter umbilicus that is oval rather than slit-like, foliate 

L. R. Saul and R. L. Squires, 2008 

Paire 121 

collabnil sculpture, and no parietal swellint:; on the inner 
lip. Ariadnoria ainikta differs from A. aldcrsoni new spe- 
cies hv being; larger, haxinsr nnich stronger siiiral cords 
with nuich wider interspaces, a well rountled last whorl 
I, not angulate), shorter nnibilicus that is o\al rather than 
slit-like, and no ridge separating the umbilicus from the 
inner lip. Ariadnaiia ainikta differs from A. ohstricfa 
(\\'hite, 1SS9) hv having a lower spire, wider penultimate 
whorl, round last whorl (not angulate) more spiral cords, 
toliate collabral sculpture, shorter umbilicus that is oval 
ratlier than slit-like, and no ridge separating the nnibili- 
cus from the inner lip. The strong spiial libbing ol A. 
ainikta resembles that of A. obstiicta. 

EtATiiology': Ainiktos, Greek, meaning: baifling, ob- 
scure, or enigmatic. 

Ariadnaiia stibara new species 
(Figures 5-6) 

Diagnosis: A medium size sturtK Aiiadnariu with 
rounded whorls, mediiuii strong scidptru'e with many 
spiral cords moderateh- closely spaced; collabral sculp- 
ture \en" fine and lattice-like on spire whorls; umbilicus 
chink-hke; inner lip with low parietal swelling. 

Description: Shell medium small (height appro.xi- 
mateK' 20 mm), stin^dy, turbiniform, somewhat elongate, 
spire high and approximately 50% of total shell height; 
apical angle approximateh' 67°; protoconch not pi'esent; 
teleoconch whorls toiu', whorls well rounded and enlarg- 
ing rapidlv; last whorl tapering anteriorly; suture ap- 
pressed, appearing channeled, and rapidly descending 
near apeiture; umbilicus narrow, chink-like and present 
onh' adjacent to niechal and posterior paits of inner lip; 
growth line prosocline, with several irregularly spaced 
growth checks near outer lip; spiral sculpture consisting 
of numerous fine snbequal roimded cords; approximately 
ten closely spaced cords on penultimate whorl with in- 
terspaces of nearly equal width; approximateK 18 cords 
on last whorl with interspaces slightK' wider than cords; 
cords strongest, most wideK' spaced, and occasionally 
widi spiral thread in interspaces on medial and anterior 
portions of last whorl; collabral sculpture consisting of 
tliin, raised growth lines, forming nearly microscopic lat- 
tice-like pattern on spire whorls; aperture D-shaped, 
moderateh' large, oblique, narrowed at posterior end by 
low parietal swelling; inner lip Ijroad. somewhat exca- 
vated (concave) mecUallv and flattened anteriorly; aba- 
pertural edge of inner lip delineated b\ low but distinct 
ridge; basal lip broadened. 

Holotype: Holotxpe LACMIP 1.3371, height 19 mm 
(incomplete), diameter 13.5 mm, .spire height 9 mm. 

T\pe Locality: LACMIP loc. 23464 is also type local- 
it\ of Tiinilites dilleri Vluiphv and Rodda, I960. 

Distribution: Bald Hills Member (unit W of Matsu- 
moto. 1960) of the Budden Canvon Formation, (area 4) 
Ono area. Shasta Co., California. 

Geologic Age: Middle Cenomanian. 

Discussion: OiiK the holotype is known, and it lacks 
the protoconch antl tiie anterior end of the teleoconch. 
W^iether or not an anterior siphonal notch was present is 
unknowai, but the shape of the last whorl suggests that at 
least a small one was present. The species is assigned to 
Ariadnaiia based on shell shape, but it differs from typi- 
cal Aradnaiia in its veiy sturdy shell, the fineness of its 
sculpture, and the presence of a parietal swelling at the 
posterior end of the aperture. 

Ariadnaiia stibara diflcrs Ironi ,\. ainikta \)\ having a 
more elongate shell, narrower peiinltimate whorl, much 
weaker spiral cords with nnich narrower interspaces, 
longer and narrower umbilicus, parietal swelling on inner 
lip, and absence of foHate collabral sculpture. Ariadnaiia 
stibara differs from A. aldersoni by larger size, less elon- 
gate shell, more rounded whorls that are not lax in their 
coiling, coarser spiral ribs, lattice-like vei-y fine collabral 
sculpture on spire whorls, paiietal swelling on inner lip, 
less delineated abapertiual edge of inner lip, and no fas- 
ciole-like flange bounding the umbilicus. Ariadnaiia 
stibara differs from A. obsirida h\ having rounded 
whorls, many more spiral ribs that are much weaker and 
much more closely spaced, and a less well demarked 
abapertural edge of the inner hp. 

Etymology: Named for its sturtlv shell, stibaros, 
Greek, meaning strong or sturdy. 

Ariadnaiia aldcrsoni new species 
(Figures 7-S) 

Diagnosis: Small .Aria(ln;Lri;i with elongate shell and 
last whorl medially subangulate, coiling stretched a.xially, 
whorls bearing many veiy fine and closely spaced ribs, 
umbilicus chink-like, spiral sculpture veiy fine, abaper- 
tural edge of inner lip well delineated and raised, lunbi- 
licus bordered by fasicole-like llange. 

Description: Shell small (approxnriatelv 13.5 mm 
height, estimated); elongately turbiniform, with medially 
subangulate whorls; upper spire missing; teleoconch 
whorls enlarging rapidly; suture appaivntlv appressed on 
earlier whorls, becoming channeled on penultimate 
whorl, rapidly descending; umbilicus chink-like, 
bounded bv strong rountled fasciole-like ridge; growth 
line prosocline, well marked with numerous growth welts 
on last whorl; spiral sculptm^e consisting of fine, well 
spaced ribs of unequal strength crossing irregular growth 
welts; interspaces between ribs widei- than ribs; aperture 
D-shaped. narrowed posterioiK but not angled, with 
short, broad anterior sinus; outer lip thin, simple; inner 
lip moderately narrow, abapertiu-al edge raised and well 

Holotype: LACMIP 13372, height 12 mm (incom- 
plete), diameter 8.5 nun. 

Tjpe Locality: LAC;M1P 26.370. 

Distribution: Panoclie Formation, (area 11) Alcalde 
Hills. Fresno Co., Cahfornia. 

Page 1 


THE NAUTILUS, Vol. 122, No. 3 

Geologic Age: Late Turonian. 

Discussion: Oiil)' the holo1y[3t' is known. It is incom- 
plete, consisting only of the last two whorls, and its small 
size may indicate that it is not mature. The elongate 
shape is a distinctive characteristic oi this species. Ari- 
adnaria aldersoni apparently had a relatively high spire, 
and the coiling is lax and reminiscent oi Liijisa Stephen- 
son, 1952. It somewhat resembles Lhysa teres Stephen- 
son, 1952, but the new species has a narrow, chink-like 

Aiiadnaria aldersoni is most similar to A. stihara and 
differs by being smaller, having a more elongate shell, 
angulated whorls that are lax in their coiling, much 
weaker spiral ribs, abapertural edge of inner lip better 
delineated, fasciole-Uke ridge bounding the umbilicus, 
absence of lattice-like very fine collabi'al sculpture on 
spire whorls, and absence of parietal swelling on inner 
lip. Aiiadnaria aldersoni differs from A. ainikta bv being 
smaller, having much weaker spiral cords with much nar- 
rower interspaces, angulate last whorl, longer umbilicus 
that is sht-like rather than oval, and having a ridge sepa- 
rating the umbilicus from the inner lip. Ariadnaria al- 
dersoni differs from A. ohstricta by being smaller and 
having fewer and much weaker spiral cords with much 
narrower interspaces. 

In some respects A. aldersoni is similar tt) Lijsis sncien- 
sis. Both have fine ribbing, a reladvel)' high spire, rather 
lax coihng, and a somewhat slower increase of whorl 
diameter. Whereas the suture of A. ohstrieta and Lijsis 
niickeiji is very close to or at the perimeter of the previ- 
ous whorl, in A. aldersoni and Li/sis sueiensis the suture 
is nsualK abapical U> the pre\it)us wliorl's perimeter. 

Etymology: The species is named for John M . Alder- 
son who collected the holotvpe from Cooper Canvon. 

Ariadnaria ohstricta (\\''hite, 1889) 

(Figures 9-10) 

Stomatia ohstricta Wliite, 1889: 18-19, pi. 4, figs. lO-H. 

Diagnosis: Medium size Ariadnaria with high spire 
and angulate last whorl, sculpture of a few widely spaced 
strong cords, umbilicus chink-like or covered. 

Description: Shell medium size (appro.ximately 21.4 
nnn height), elongate turbiniforni; whorl profile rounded 
with slight angulation at third strong cord on last whorl; 
spire high and appro.ximately 50% of total shell height; 
apical angle approximately 67°; protoconch missing; 
teleoconch whorls four, enlarging rapidly and last whorl 
tapering anteriorlv; suture appressed, anterior to suban- 
gulate peripheiy; umbilicus chinklike or covered by inner 
lip expansion; bounded abaperturally by strong ridge; 
growth line prosocHne; sculpture of three strong, widely 
spaced cords on spire, five or six on bocK whorl; inter- 
spaces commonly with mid thread; aperture large and 
ovate with abapertural edge raised and sharply de- 
marked; outer lip apparently simple; inner lip somewhat 
expanded and standing high along umbilical chink; basal 
lip barely drawn out into slight spout-like sinus. 

Holotype: USNM 20124. 

Type Locality: Little Cow Creek l^ut additional speci- 
mens have not been found there. The species is abun- 
dant at some localities along South Cow Creek, Shasta 
Co., California. 

Hvpotyi>e: LACMIP 13.'37.3, height 20 mm, diameter 
1.5 nnn' from LACMIP loc. 28717." 

Distribution: Redding Formation, Bear Creek Sand- 
stone Member, especially (area 3) along South Cow 
Creek and Bear Creek, Shasta Co.; Chico Formation, 
Musty Buck Member, (area 5) Chico Creek, Butte Co., 

Geologic Age: Late Coniacian? to Santonian. 

Discussion: The above description is based on 16 
specimens; all but one are from LACMIP loc. 28717. 
Most specimens are poorly preserved, and specimens 

with the shell surface presened are difficult to find. The 
primaiy cords are strong and almost ilange-like. 

White's species is here assigned to Ariadnaria based 
on shell shape, sculpture, umbiUcus, and presence of a 
small spout-like sinus in the aperture. This would be the 
earliest unquesticmed occurrence of this genus that pre- 
\ iousK' was knowm only from the Recent (Wenz, 1940). 

In shape and probably sculpture (preservation makes 
comparison difficult) A. ohstricta resembles illustrations 
oi'Trichotropis? sp. in Kase (1990: .568, figs. 2.26, 2.27). 
Kase's specimen was from the Izumi Group of Japan of 
early Maastrichtian age. 

Ariadnaria ohstricta is veiy similar to Li/sis niickei/i 
new species, but .A. ohstricta has more I'egiilar spiral ribs, 
a shghl\- higher spire, the strong spiral delimiting an um- 
bilical chink, and a free standing imier lip. Ariadnaria 
ohstricta differs from Ariadnaria ainikta hv haxing a 
higher spire, narrower penultimate whorl, angulate last 
whorl, fewer spiral cords, shorter umbilicus that is slit- 
like rather than oval, ridge separating the umbilicus from 
the inner lip, and an absence of foHate collabral sculp- 
ture. Ariadnaria ohstricta differs from A. aldersoni by 
being larger, having fewer spiral cords that are much 
stronger and much more widely spaced, aiid lacking a 
fasciole-like ridge bounding the chink-Uke umbilicus. 
Ariadnaria ohstricta differs from A. stihara hv havino; 
angulate whorls, fewer spiral ribs that are much stronger 
and mucli more widely spaced, and a more demarked 
abapertural edge of the inner lip. 

Snbtannlv Lysinae new subfamily 

Description: Small to moderatelv large (15 to 80 nnn 
in height), low turbiniform to almost haliotiform, barely 
siphonate shells with spiral ribbing. Final whorl some- 
what to greatly enlarged; spire ven' short; apertm'e large, 
nearly circular to elongate oval; columella and inner lip 
flattened, expanded, and depressed to form a shelf within 
the apeiture; some witli shelf tliat spirals into an "umbi- 

Discussion: i'lic sublannK L\sinac nichides Li/sis 

L. R. Saul and R. L. Squires, 2008 

Page 123 

Gabb, Gai-zasia ne\\' genus, and probably Spirogalerus 
Finlay and Mar\\ick, 1937. These gastropods are inter- 
mediate in form between trichotropines and calyp- 
traeids. If their characteristics were better knowai, some 
other species such as those discussed under Global Dis- 
tribution oi Cretaceous Lvsiform Gastropods, probably 
could be included here, some as Li/sis or Gai-asia others 
in as \'et undescribed genera. 

Genus Lijsis Gabb, 1864 
Tropidothais Cox, 1925: 213-214. 

T\pe Species: Lyais duplicosta Gabb, 1864, by mono- 
t^px- (Stewart, 1927: 345); Campanian of Pacific slope of 
North .America. 

Description: Turbinate to crepiduliform gastropods 
with a rapidh- e.xpanchng whoi'l diameter haxdng the col- 
umella/inner lip flattened, expanded, and submerged to 
form a narrow to broad shelf or deck. Shell sculptured by 
spiral cords or smooth. Nonumbilicate. Aperture with 
\"ei"\" slight anterior siphonal notch. 

Discussion: Li/sis differs from Tricliotmpis and Ari- 
adnaria in haxing die inner lip completely appressed to 
tlie columella. Typical Lysis (i.e., tlie group of L. dupli- 
costa) has a carinated whorl in the juvenile stage and a 
few moderately strong to strong spiral cords. The stron- 
ger spirals are t^picall^■ scalv. Included in this group is L. 
duplicosta and the following new species: Lysis mickeyi, 
L.jalamaca, and L. lomaensis. The group oi Lysis sucien- 
sis has a more rounded whorl profile and more subdued, 
finer spiral sculpture; included in it is L. suciensis. 

Group of Lijsis duplicosta 

The genus Lysis was proposed by Gabb (1864) for a 
lo\\"-spired, tiu-biniform gastropod with a depressed inner 
lip. He had onK' immature specimens of a single species 
(i.e., the type species) and did not recognize their simi- 
laritv' to genus Crepidida Lamarck, 1799. He gave no 
indication of the familial affinities of Lysis, other than 
stating the general form is like genus Stouiatia Helbling, 

During tlie last 127 years. Lysis has been placed in at 
least 11 famihes scattered among "archaeogastropods" to 
the neogastropods. A review of this placement histon" is 
gi\-en here. Species that were eventuallv placed in Lysis 
were originalh' placed in Stomatia of the Stomatellidae 
Gray, 1840, bv Whiteaves (1879, 1903) and White 
(1889). Stoliczka (1867-1868: 157-158) suggested that 
Lysis should be placed near Separatista Gray, 1847, in 
the Trichotropidae Gray, 1850, in the event that Lysis 
does not belong in either die Naticidae Guilding, 1834, 
or the \'elutinidae Gray, 1840. Fischer (1885) placed 
Lysis in die Naticidae near Eunaticina Fischer, 1885. 
Tr\"on (1884: 112) did not hesitate to refer Lysis to the 
Muricidae Rafinesque, 1815 (as Murexia) [= Purpuradae 
Clnldren, 1823], but on page 208 Tryon suggested a re- 
lationsliip to Velutina Fleming, 1821, of the Lamellari- 
idae d'Orbignv. 1841. Gossmann (1903) wrote that Lysis 
could not be a muricid but must be placed near Fossarus 

Philippi, 1841, presumably in the Fossaridae Adams, 
1860, where Stewart, 1927,' Rennie (1930), Wenz, 1940, 
and Anderson (1958) also put it. Cossmann (1925) con- 
sidered Lysis to be a subgenus oi Micreschara Cossmann 
(1891) in the family Vanikoroidae Gray, 1840. DaU in 
Eastman (1913) and Packard (1922) placed Lysis in the 
"Thaisiidae" (=Thaididae) Suter, 1909. Saul ("l959) and 
Saul and Alderson (1981) placed Lysis in Galyptraeidae, 
and, in 1990, Saul included it in superfaniily Calyptrae- 
oidea. Kase (1990) discussed previous taxononiic treat- 
ments of Lysis and suggested that, based on its inner hp 
shelf, the genus should be placed within the Cahp- 
traeidae Lamarck, 1799. Bandel and Riedel (1994) and 
Kiel and Bandel (2003) supported this placement. Cox 
(1925) named and described genus Tropidothais Cox, 
1925, which is a junior synonym of Lysis. He based 
Tropidothais on T. africana Cox, 1925, from the Maas- 
trichtian of Mozambique [formerly Portuguese East Af- 
rica] and tentatively placed his genus in the Thaididae 
Jousseaume, 1888. Upon reahzing its similarit}' to Lysis. 
Cox (in Rennie, 1935) synonomized the two genera and 
placed Lysis in the StomateUidae. In this present paper, 
we place Lysis in the family Trichotropidae, subfamily 
Lysinae because Lijsis appears to have evolved from tri- 
chotropids by expanding the columella/inner lip area 
(width and length) to form an interior shelf suggestive of 
the calyptraeid Crepidula. 

Lysis mickey i new species. 
(Figures 11-16) 

Diagnosis: A relativeh- high spired Lysis with eight or 
nine strong cords on last whorl; shelf moderately broad, 
somewhat depressed, and shallowly concave. 

Description: Shell medium small (height approxi- 
mately 20 mm), turbiniforni; whorl profile overall 
rounded with medial angulation on last whorl; spire mod- 
erately low and appro.ximately 30% of total shell height; 
apical angle approximately 90°; protoconch 1.5 whorls, 
low and smooth; teleoconch whorls 3.5, moderately ex- 
panding and last whorl tapering anteriorly; suture abut- 
ting and becoming laxly channeled on later whorls; sculp- 
ture of strongly raised cords, either moderately closely 
spaced or \\idely spaced; penultimate whorl with two to 
six and last whorl with eight or nine strong spirals, with 
variable number (two to five) of finer spirals in inter- 
spaces; both cords and interspaces crossed by fine coUa- 
bral ribs, producing beaded appearance; angulation mod- 
erately strong on last whorl, coincident with strongest 
spiral cord; spiral cord anterior to angulation neai'ly same 
strength, thereby producing bicarinate appearance to 
medial part of last whorl; aperture large, oblique, barely 
notched anteriorly; outer lip simple; shelf moderately 
broad, somewhat depressed, and shallowly concave. 

Holotype: LACMIP 13374, height 13 mm, diameter 
9.5 mill, spire height 5 mm. 

Paratypes: LACMIP 13375 from LACMIP loc. 23617 
and 13376 and 13377 from LACMIP loc. 10757. 

Paee 124 

THE NAUTILUS, Vol. 122. No. 3 





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Figures 11-28. Ly.sw species, 11-16. Lijsis mickciji new species, 11-12. Paratvpe LACMIP 13375, LACMIP loc. 23617, height 
21 mm, diameter 16 mm. 11. Apertural \'iew. 12. Right-lateral view, 13. Paratype' LACMIP 13376, LACMIP loc. 10757. left-lateral 
view, height 9 mm, diameter II mm. 14. Holotvpe LACMIP 13374, LACMIP loc. 10757, abapeitiiral \iew. height 13 mm, diameter 
9.5 mm. 15-16. Paratope LACMIP 1.3377, LACMIP loc. 10757, height 5 mm, diameter 9 mm. 15. Left-lateral view. 16. Apical view, 
17-28. Lijsis diiplicosta Gabb, 1864. 17. Plasto-lectoty|3e oi Stomatia sticiensis caiinifera Whiteaves, 1879, CCS 5772, height 10 mm, 
diameter 19.5 mm. 18-19. Hypoty|3e LACMIP 13378, LACMIP loc. 24128. IS. Apertural view, vertical dimension 27 mm, 

horizontal dimension 21 i , 19. Abapertural view, height 22 mm, diameter 26,5 mm. 20-21. Hvpohpe L.'\CMIP 13379, LACMIP 

loc. 24340. 20. Apertural view, veitical dimension 28 mm, horizontal dimension 25 nnii. 21. Lateral \iew. vertical dimension 13 mm, 
horizontal diameter 29.5 mm. 22-23. Hvpoty|3e LACMIP 13380, LACMIP loc. 24340, height 19 mm, diameter 18 mm. 22. 
Apertural view. 23, Abapertural view. 24. Hypotvpe LACMIP 13381, LAC^MIP loc. 2434(J, abapertural view, veitical dimension 28 
mm, diameter 37.5 mm. 25. IIvp(.)tvpe LACMIP 13382, LACMIP loc. 24.349, abapertural view, height 17 mm, diameter 22 mm. 26. 
Hvpc)t)pe L.'VCMIP 13383, LACMIP loc. 10095, crushed specimen, abapertural view, height 24 mm, diameter 22.5 mm. 27. 
Hypotype LACMIP 13384, LACMIP loc. 26951, abapertural view, height 21 mm, diameter 25 mm. 28. Hypot)pe LACMIP 13385, 
LACMIP loc. 24340, abapertural view, height 7.5 mm, diameter 12 mm. 

L. R. Saul and R. L. Squires, 2008 

Page 125 

Type LocaliK: LACMIP loc. 10757. 

Distribution: Redding Formation, Bear Creek Sand- 
stone Member of Haggart (area 3) on Bear Creek, Shasta 
Co.; Chico Formation, top of Ponderosa Way Member 
and Must\- Buck Member (200 m to 650 m above the 
base oi the section) (area 5) on Cliico Creek, Butte Co.; 
basal Tuna Can\'on Formation (area 17) at liead of Ga- 
rapito Creek, Santa Monica \h)untains. Los Angeles Co., 

Geologic Age: Earl\ Coniacian to Santoisian. ^-= 

Discu.ssion: The above description is based on 80 
specimens; most of tliese are from LACMIP Iocs. 10846 
and 23617. Most specimens are internal molds. Many 
show endobiont boreholes, especialK' on the spire 
\\'horls. Two specimens show the protoconch. Tlie oldest 
specimen is from LACMIP loc. 26967 in the Santa 
Monica Mountains. 

Ltisis michct/i resembles Lijsis ■ntcicnsis (Whiteaves, 
1879") in height of spire but is closer to L. duplicosia in 
sculpture. Lijsis mickciji differs from L. cluplicosta in 
ha\ang a higher spire, less expanding last whorl, much 
less expanded shelf, more o\al aperture, and thimier spi- 
ral cords. Lysis mickeiji greatly resembles Trichotropis 
obstiicta (White, 1889), but on L. mickciji the spiral ribs 
are less regular, and its spire is slightly lower. In addition, 
L. mickciji lacks an umbilical chink and a free-stancUng 
inner lip. 

Li/sis mickciji is the earliest knov\ai Lysis from any- 
\\'here in the world. 

EtMnolog\^: Named for Mickev of Mickev's House on 
Cliico Creek near the locahty, LACMIP 23617, from 
which tlie species was first recognized. 

Lysis diiplicosta Gabb, 1864 
(Figures 17-28) 

Lysis cluplicosta Gabb, 1864: 1.38, pi 21, fig. 98a-98c; Tiyon, 

188.3: 112. pi. 44, ficr. 2.5-26; Cossmann, 1903: 70; .Stewart, 

1927: .34.5-346, pL21. figs. 7, 7a; Anderson, 1958: 169. 
Stomatia suciensis \-c\het\' cariiiifeni Wliiteaves, 1879: 128-129, 

pi 16, fig. 5. 
Lysis oppansus White, 1889: 17. pi 4, figs, l-t-15; Anderson, 

1958; 169. 
Li/sis suciensis \ar. carinifera (\\'liiteaves). — ^^^^itea\■es, 1903: 

367, pf. 45, fig. 4. 
Micreschara (Li/sis) cluplicosta (Gabb). — Cossmann, 1925: 173, 

pi 9. figs. 6. 21. 
Lysis duplicostafa Gabb. — Wenz, 1940: 880. fig. 2587 (reprint 

of Stewart, 1927): Elder and Saul. 1993: pf 2. figs. 14-15. 
Lysis carinifera f\\'liiteaves). — Anderson, 1958: 170. 
? Lysis duplicosta carinifera (Whiteaves). — Dailev and Pope- 

noe. 1966: 6. 
Xot Lysis duplicosta Gabb. — Saul and Alderson, 1981: 35-36, 

pi 3, figs. -3-4 [= Lysis suciensis (Wliiteaves) /(f/p Saul, 


Diagnosis: \'ariabl\' sculptured Lysis, with man)' fine 
cordlets or with six to eight strong cords, including 
prominent (often flange-like) carina on periphery; col- 
umella and inner hp depressed and expanded to form 

crescentic shelf, occup)'ing at least one third of aperture 
in larger specimens. 

Description: Shell mecUum size (height up to approxi- 
matel\- 26 nun), neritiform to crepiduliform; spire mod- 
erately low, approximately 20% of total shell height; pro- 
toconch 1.5 whorls, low and smooth; teleoconch approxi- 
matelv two whorls, overall rounded, enlarging very 
rapidly, and medially carinate; sculpture consisting of 
spiral ribs, generally six to eight prominent ones, but 
highh' v;niable in number, spacing, and strength; periph- 
eiy alwa\s dernarked b\" \en' strong (occasionally flange- 
like) carina, located anteriorfy of medial position on spire 
whorl and located mediallv on last whorl; remainder of 
whorls covered bv spiral sculptiu'e, ranging from nimier- 
ous closely spaced fine cordlets to several uidelv spaced 
moderatefy strong (can alternate iii strength) cords, with 
interspaces smooth or bearing many cordlets or threads; 
cords just anterior and, to a lesser degree, just posterior 
of medial carina on last whorl commonlv approaching 
strength of medial carina, thereby imparting either a bi- 
carinate or tricarinate appearance to whorl profile; aper- 
ture circular with a scarcely discernable anterior canal 
notch; abapertural edge of apertiu'e shaipK* demarked bv 
raised edge; inner lip and columella fliittened, and ex- 
panded to form shelf; shell moderately wide (occupwig 
at least one third of apertui'e in larger specimens), sub- 
merged within the aperture, wrapping far past suture and 
attached to inside of outer lip; medial part of outer lip 

Lectotjpe: Oi' Lysis duplicosta UCMP 11975, height 
10 mm, diameter 19.5 mm. Gabb (1864) did not indicate 
a holotyjje. Merriam (1895) recognized UCMP 11975 as 
the figured specimen. Stewart's (1927) statement that 
this is the tvpe specimen is taken as designation of lec- 


Paralectot>pe: Of Lysis duplicosta ANSP 4242. 

Syntypes: Stomatia suciensis varietv carinifera 
Whiteaves. 1879. CGS 5772, a-d (Bolton, 1965). 
Whiteaves (1903; pi. 16, fig. 5) figured one of the five 
sviity-pes CGS 5772. 

Holotvpe: Of Lysis oppansus White, 1889, USNM 

Hypotypes: Oi Stomatia suciensis carinifera, CCS 
5939 (Whiteaves, 1903); Of Lysis duplicosta LACMIP 
13378-13385; USNM 468585, 468586. 

Type Locality: Of Lysis duplicosta, Texas Flat, near 
Rock Corral, from a mine shaft at a depth of 12 m [40 
ft.], near the Phicer-Sacramento Co. line. Placer Co., 
northern California; Of Stonuitia suciensis carinifera, Su- 
cia Island, San Juan Co., Washington. Oi' Lysis oppansus, 
Pentz Ranch, Butte Co., northern California. 

Distribution: Cedar District Formation, Nanaimo Ba- 
sin, (area 1) Vancouver Island area, southern Britisli Co- 
lumbia and (area 2) Sucia Island, San Juan Co., Wash- 
ington; Chico Formation, Ten Mile Merjiber on (area 5) 

Page 126 

THE NAUTILUS, \'ol. 122, No. 3 

Chico Creek and Must}- Buck Member along (area 6) 
Dr\' Creek, near Pentz, Butte Co., California; Chico For- 
mation, (area 7) Granite Bav and Te.xas Flat, Placer Co., 
California; Pigeon Point Formation, southern sequence, 
(area 8) north of Pigeon Point, San Mateo Co., Califor- 
nia; Jalama Formation, (area 12) western Santa Ynez 
Mountains, Santa Barbara Co., California; Ladd Forma- 
tion, uppermost Holz Shale Member and Williams For- 
mation, Schulz Member, (area 18) Santa Ana Mountains, 
Orange Co., CaBfomia. 

Geologic Age: Campanian. 

Discussion: The above description is based on 101 
specimens; most of these are ironi the Pentz area (LAC- 
MIP loc. 24340). At any locality, most of the specimens 
are internal molds. Some show endobiont boreholes. 
One specimen, LACMIP 13385 (Figure 28), shows the 
shape of the protoconch. 

The holot^'pe of L(/.s';.s' dnplicosta is a juvenile and had 
not yet developed the wider deck of an adult. The largest 
specimens of this species thus far found are from the 
Must\' Buck Member of the Chico Formation at LAC- 
MIP loc. 24340 near Pentz (area 6). The specimens are 
from a matri.\-supported pebble conglomerate richly fos- 
siliferous in places. The fauna, which includes scraps of 
cvpraeids, suggests warm, shallow water. Li/sis diipUcosta 
is veiy rare in the coeval Ten Mile Member on Chico 
Creek (area 5) which probably represents deeper water 
than at LACMIP 24340, and the specimen from LAC- 
MIP loc. 23639 on Chico Creek may have been trans- 
ported dowmslope. 

The strength of the cords, especially of the strongest 
one, varies greatly between individuals. Gabb's specific 
name refers to a doubled appearance of each major cord, 
but Gabb's (1864) shaqD, deep channel along the strone 
ribs results from the wearin 
on the ribs. 

Dailey and Popenoe (1966: fig. 3) listed Lysis dnpli- 
costa carinifera from the Jalama Formation at LACMIP 
loc. 24128. This somewhat distorted specimen (Figures 
18, 19) has some stronger ribs as in L. dnplicosta. but 
may represent a strong ribbed variant of L. jalamaca. If 
L. dnplicosta. it is the geologicallv youngest specimen of 
this species. 

Rennie (1930: 1935) described two species of Li/sis 
from the Umzamba Formation of the Eastern Cape 
Province (formerly Pondoland), South Africa, one of 
which Lysis capensis Rennie, 1930, is veiy similar to L. 

or breaking off of the scales 

dnplicosta. According to Klinger and Kennedy (1980), 
the lower Umzamba Formation at its type locality is lat- 
est Santonian or earliest Campanian in age and, there- 
fore, similar in age to the Chico Formation near Pentz, 
Butte Co., California, where L. dnplicosta is common. 
Rennie apparentK' had onlv two specimens of L. capensis 
which he said had six stout, sharp, spiral ribs. His holo- 
type is small, similar in size to the holotype of L. dnpli- 
costa, and both apparently have a relatively narrower 
deck than is found in large specimens of L. dnplicosta. 
Kase's (1990) report of L. dnplicosta in the Chats- 
worth Formation is based on a misidentification of L. 
snciensis in Saul and Alderson (1981). 

Lysis jalamaca new species 

(Figures 29-36) 

Lysis dnplicosta Gabb. — Dailev and Popenoe, 1966: 6. 

Not Li/sis dnplicosta Gabb, 1864. 

Diagnosis: A Lysis with sculpture of fine spiral 
cordlets of alternating strength and within the aperture 
the shelf wrapping considerably past suture. 

Description: Shell medium size (height up to 26 mm), 
crepiduliform, with rapidl\- expanding whorls; spire low. 
and approximately 30% of total shell height; protoconch 
1.5 whorls, low and smooth; teleoconch approximately 
two whorls, well rounded; whorls flattened adjacent to 
suture; periphery near mid-whorl height; suture abutting 
just below periphen'; growth line prosocline, occasionally 
forming coUaliral wrinkles; sculpture consisting of nu- 
merous closely spaced fine spiral cordlets alternating in 
strength and somewhat scaly; cordlets strongest at and 
near peripheiy where two or three can be stronger be- 
coming cords and the scales bead-like, with approxi- 
mately three cordlets in interspaces; aperture ovoid, its 
abapertural margin shaiplv demarked and steeply de- 
scending; inner lip and columella flattened and expanded 
to form shelf; shelf narro\v anteriorK becoming moder- 
ately wide mediallv, submerged within aperture, wrap- 
ping past suture and attached to inside of outer lip. 

Holotype: LACMIP 13386, height approximateh- 35 
mm, diameter 34 mm. 

Paratvpes: LACMIP 13387 (LACMIP loc. 24137) 
-13388 (LACMIP loc. 24122) and SDNHM 114595, 
114596 (SDNHM loc. 3405). 

Type Locality: 


LACMIP loc. 24137, Jalama Forma- 

Figures 29-39. Lysis species. Specimens coated witli ariniionluni chluridc, 29-36. Lysis icihiiiuiai new species. 29-30. Holotype 
LACMIP 13386, LACMIP loc. 24137. 29. Apertural view, vertical dimension 34 mm, liorizontal dimension 30 mm. 30. Abapertural 
view, vertical dimension 27 mm, diameter 34 mm. 31-32. Paratype LACMIP 13387, LACMIP loc. 24137. 31. Left-lateral view, 
height 18 mm, horizontal dimension 19 mm. 32. Lateral view, vertical dimension 11.5 mm, diameter 28 mm. 33. Parahpe LACMIP 
13388, LACMIP loc. 24122, abapertural view, height 11 mm, diameter 13 mm. 34. Paratypes SDNHM 114595 and 114596, SDNHM 
loc. 3405 on outer lip of volute gastropod Lon^oconcho ciiincka Saul and Squires, 2008 (SDNHM 70974), SDNHM loc. 3405, height 
133 mm, diameter 33 mm. .3.5-.36. Posteiiormost paratvpe SDNHM 114596 on volute shown in previous figure. .3.5. Abapertural 
view, height 13 mm, diameter 20 mm. 36. Oblique apical view, vertical dnnension 17 mm, diameter 20 mm. 37-39. Ij/.sis loiiiaciisis 
new species, holotvpe SDNHM 67150, SDNHM loc. 3403, height 22 mm, diameter 23 nmi. 37. Apertural view. 38. Abapt'rtural 
view. 39. ApicaMateral view, vertical dimension 8 mm, diameter 23 mm. 

L. R. Saul and R. L. Squires, 200S 

Page 127 

















^ ' 





^ 34 

Distribution: |alaiiia I'^onnation, (area 12) western 
Santa '^'nez Mountains, Santa Barbara Co.; Debris Dam 
Sandstone, (area 13) Agua Caliente Canyon, San Rafael 
Mountains, Santa Barbara Co., Califomia: Chatsworth 
Formation, upper part (area 16) at Lang Ranch, Ventura 
Co., California; Point Lonia Formation, (area 19) near 
Carlsbad, San Diego Co., California; Rosario Formation, 
(area 22) Caiion San Fernando, northwestern Baja Cali- 
fornia, Vlexico. 

Geologic Age: Late Campanian and early Maastrich- 

Discussion: The above description is based on 13 
specimens. The best presen'ation occurs in the Point 
Loma Formaton near Carlsbad. 

Li/sis jalamaca is similar to L. sucivusls but has 
coarser, more scaly costae than L. suciensis, and available 
specimens of L. jalamaca are smaller and less elongate 
than are large specimens of L. suciensis. Lysis jalamaca 
is also similar to L. dupUcosta in haxdng variable sculp- 
ture and in having the strongest cords on the peripheiy, 
but L. jalamaca is less angnlate at the peripheiy and has 
weaker cords there. In addition, L. jalamaca differs from 
L. dupUcosta by being more elongate, having finer sculp- 
ture, and having the shelf broader posteriorly. The speci- 
men (LACMIP 13378, Figures 18, 19) from the Jalama 
Formation has ribbing similar to L. dupUcosta, but its 
shelf appeal's to wrap farther (jnto the interior of die 
outer whorl than is usual for L. dupUcosta. 

Etymology: The specific epithet, a name in apposi- 
tion, reflects the new species occurrence in the Jalama 
Formation, Santa Barbara Co., Califomia. 

Lysis loDiacusis new species 
(Figures 37-39) 

Diagnosis: Li/sis with low spire and last whorl bearing 
numerous flanged carinae separated by wide interspaces 
bearing se\eral spiral threads. 

Description: Shell medium size (up to 17 mm height 
and 30 iiiiii diameter, same specimen), neritifonn (last 
whorl rapidly expanding); spire very low, approximately 
10 to 15% of shell height; protoconch 1.5 whorls, low and 
smooth; teleoconch 2 to 2.5 whorls, veiy carinated; su- 
ture abutting just anterior to peripheiy; sculpture con- 
sisting of up to 11 strong cords; cord at periphen- and 
next three cords posterior of peripheiy veiy thin, flanged, 
and protniding (with cord at peripheiy protruding most); 
posteriormost part of last whorl (in vicinity of suture) 
with approximately three low scaly cordlets; up to three 
beaded to lowly spinose cordlets anterior to medial ca- 
rina at periphery; interspaces between all cords wide and 
bearing up to seven spiral threads (occasional thread can 
locally develop into small flanged cord); area anterior to 
medial carina can be covered with only fine cordlets; 
aperture circular with margin sharply demarked by 
raised edge; shelf moderately wide, wrapping past suture 
and attached to inside of outer lip; posterior lialf of outer 
lip digitate. 

Page 128 

THE NAUTILUS, \"ol. 122, No. 3 

Holotype: SDNHM 67150, height 22 mm, diameter 
23 mm. 

Paratope: SDNHM 67152 from SDNHM loc. 4071. 

Type Locality: SDNHM loc. 3403. 

Distribution: Basal San Fraiici.s(juito Formation, 
(area 14) Warm Springs Mountain, Los Angeles Co.; 
Point Loma Formation, (area 19) San Diego Co., Cali- 
fornia; Rosario Formation, (area 21) five miles east of La 
Mision, northwestern Baja California, Mexico. 

Geologic Age: 


Latest Campanian to late Maastrich- 

Discussion: Five specimens \\ere studied. Presetva- 
tion is generally excellent for three specimens from the 
Point Loma Formation near Carlsbad, although the pro- 
toconch of the holotype is imperfectly presei-ved. The 
incomplete, crushed specimen from Warm Springs 
Mountain is from lieds of latest Maastrichtian age at the 
base of the San Francisquito Formation (LACMIP loc. 

14310). Altliough tlic abapertural side is not availaiile, 
the flatness of tlie apical whorls suggests that it is prob- 
abh' L. lomaensis. 

The new species is most similar to Lysis diiplicosta but 
available specimens are smaller than a large L. diipli- 
costa. Lysis lomaensis also has a lower spire, more cari- 
nate last whorl (especially posterior to the medial carina), 
and spiral threads on interspaces between carinae. 

Abaperturally, L. lomaensis is similar in shape and 
sculpture to Garzasia intermedia, Iwt L. lomaensis has a 
lower spire and in the apertural \'ie\\' the shelf margin is 
less arcuate, not sigmoid, and the shelf does not spiral 
into the umbilicus. 

Group of Lysis suciensis 

The group ol L. sneiensis differs b'oni tliat of L. diipli- 
costa in ha^ing a more elongate aperture and finer spiral 

Lysis sneiensis (Whiteaves, 1879) 
(Figures 40-50) 



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U^' 50 

Figures 40-50. Ltjsis siiciaciisis (Whiteaves, 1879). Specimens coated \vitli ammonium chforidc. 40-41. Hvpotvpe LACMIP 
133S9, LACMIP foe. 26020. 40. Apertural view, vertical dimension 44 mm, horizontal dimension 40 mm. 41. Abapertural view, 
height 25 mm, diameter 43 mm. 42-43. HyiDotype LACMIP 13390, LACMIP loc. 1071 1. 42. Abapertural view, height IS mm, 
diameter 26.5 mm. 43. Lateral view, vertical dimension S mm, diameter 26.5 nmi. 44. Ilvpotyije LACMIP 13391, LACMIP loc. 
10095. lateral view, vertical dimension 5 mm, diameter 12.5 irnn. 4.5. Ilvpotvpe LACMIP 13392, Canon San Fernando, 32 km 
southeast of El Rosario, Baju (.'alilornia. Mexico, abapertural view, heiglit IS mm, diameter 21 mm, 46-50. Ihpotvpe LACMIP 
10495, LACMIP loc. 26020. 46-47. Vertical dimension 70 mm. horizontal dimension 38 nun. 46. .Apcrtmal \icvv. 47. Abapertural 
view. 48. Right-lateral view, vertical dimension 70 mm, horizontal dimension 25 mm. 49. Slightly oblique right-lateral \iew. vertical 

dimension 70 nnn, horizontal dimension 27 nun. .50. Lateral view, vertical dimens 

25 nnu, horizontal dimension 36 mm. 

L. R. Saul and R. L. Squires, 2008 

Page 129 

Stomatia siiciensis AMiitea\es. 1SV9: 12S-129, pi. 16, Fig. 4. 
Li/sis sticicnsis (^\"hitea^•es). — \\"hitea\e,s, 1903: 367, pi. 45, tia;. 

3. Stecheson, 2004: 60-62, pi. 2, lig.s. 4^,5. 
Li/sis californicnsis Packard, 1922: 431, pi. 37, fig.s. 2-3: Sta- 

dum, 1973: pi. 2, fig. 12. 
Lysis dtiplicosta Gabb. — Saul and Alderson, 1981: 36, pi. 3, 

figs. 3^. Not. Lijsis dtiplicosta Gabb, 1864. 

Diagnosis: Medium to large Lt/sis, slicll eliiiigatc widi 
tine, usualK' \va\"\' eordlct.s, .sculpture (ilisolete on large 

Description: Medium to large (heiglit up to S4 mm), 
crt>pidulitorm, elongateK e.xpanded; .spire moderately 
high to low, approaching horizontal coiling, and approxi- 
mateh* 30 to 35% of total shell height; protoconch ap- 
proximately 1 to 1.5 whorls, low and smooth; teleoconch 
approxiniateK' 1.5 to 2 whorls, whorls subcarinate to 
rounded; periphen- near one-third whorl height; suture 
abutting just anterior to periphei")', descending; gro\\4:h 
line slightK" to moderately parasigmoidal, with sinus at 
posterior end of outer lip on large, smooth individuals; 
sculpture consisting of fine narrow cordlets, usualK' 
wa\"\\ alternating in strength, wath narrow interspaces; 
aperture elongate ovoid, its abapertural margin shaq^lv 
demarked and steeply descending; inner lip and col- 
umella flattened and expanded to form crescentic and 
moderateh" vvide deck submerged within aperture, pos- 
terior end attached to inside ot outer lip; inner deck 
margin concax-el\' cun'ed. 

S-\Titvpes: Oi Stomatia siiciensis CCS 5771, a-d. 

Lectot\pe (here chosen): Oi Stoinalid siiciensis COS 
5771, height GfS nun. 


CCS 5771d. 

Of Stoiiuitiii siiciensis CGS 5771a- 

Holobipe: 0{ Ufsis califoniicnsis UCMP 12287, 

ParaUpe: Oi' Lysis califoniicnsis UCMP I22SS. 

T\pe Locality: Stomatia siiciensis, Sucia Island. San 
Juan Co., Washington. Of Li/sis californicnsis, UCMP 
loc. 2167, Santa Ana Momitains, Orange Co., California. 

Figured Specimens: LACMIP 10495, 13389, from 
LACMIP loc. 26020; LACMIP 13390 from LACMIP 
loc. 10711; LACMIP 13391 from LACMIP loc. 10095; 
LACMIP 13392 from the Rosario Formation, (area 22) 
26 km east of coastline at elex'ation 200 m on west side of 
Caiion San Fernando, 15 km north oi Mesa San Carlos, 
approximately 32 km. southe;ist of El Rosario. Baja Cali- 
fornia, Mexico. 

Distribution: Upper Cedar District Formation, (area 
1) Denman Island, Gulf Islands, British Columbia; 
Lower Cedar District Formation, (area 2) Sucia Island, 
San Juan Co., Washington; Chico Formation, (area 7) 
Granite Bav, Placer Co.; Jalama Formation, (area 12) 
western Santa Ynez Mountains, Santa Barbara Co.; Chats- 
worth Formation, (area 16) Bell Canyon and Dayton 
Canyon, Simi HiEs, \'entura Co., Cahfomia; Ladd For- 
mation, upper Holz Shale Member, (area 18) Santa Ana 

Mountains, Orange Co., California; Rosario Formation 
(area 22) at Canon San Fernando, 26 km SE of El Ro- 
sario Baja Calilornia; Valle Formation, (area 23) 10 km 
north from Punta Abreojos, Baja Calilornia Sur, Mexico. 

Geologic Age: Late early Camp;mian and (>arl\' Maas- 

Discussion: The al)o\-e description was based on 107 
specimens. The deck of smaller specimens is ratlier nar- 
row and attaches to the base of tlie previous whorl. Small 
specimens are rounded, larger specimens more elongate. 
In larger specimens the shelf is broader and its posterior 
end attaches to the inside of the outer lip. Several speci- 
mens, especially tlie larger specimens, have an obtusely 
biangulate whorl al:)apieal to the periphen-'. Most speci- 
mens from Sucia Island are of small size and badlv 
weathered, but Whiteaves's largest specimen (1879: 129, 
pi. 16, fig. 4), here designated as lectotvpe, was indicated 
hv him to be "two inches and a half in length" (i.e., 66 
mm). The specimen from Bell C'an\-on (area 16) of Fig- 
ures 46-50, lacking its spire and with its outer lip bioki-n, 
is 75 mm high. 

The ribbing on larger specimens Ironi tlie Chatsworth 
Formation in Bell and Davton canyons, Simi Hills (area 
16) tends to become fainter toward the outer lip espe- 
cially on the posterior slope. The specimen figured (Fig- 
ures 40, 41) retains ribbing around tlie periphen', but the 
ribs fade posteriorly toward the aperture. Ileiglit range 
of the smoother specimens is 45 to 76 mm (incomplete), 
making these the largest specimens thus far found of 
Lysis. The largest of these specimens are larger than 
representatives of most species of CrepidiiJa; only Gran- 
dicrepiclula princeps {Cjonvad, 1856), attains a larger size. 

Genus Garznsin new genus 

Type Species: Garzasia diahla new species from the 
"Garzas Sand" Member oi the Moreno Formation, Stani- 
slaus Co., California; mid Maastrichtian. 

Description: Vei-y low to moderately higli spired. Ca- 
hjptraea-like shell with weak to strong spiral cords, ap- 
erture yei"y broadly e.xpanded and ffirming base of shell, 
shelf attached marginalK, surrounding the umbilicus, 
and spiralling into it. 

Discussion: Garzasia is most similar to Lysis but dif- 
fers fr(}m the latter in that the deck surrounds and spirals 
into an umbilicus somewliat as in Calyptnica but with a 
wider more open umbilicus. It thflers from Calyptraca in 
ha-ying a sturdier shelf attached marginally as in Trochita. 
It differs from Trochita (Figure 71) in having the shelf 
crescentic with an arcuate to slightly sigmoidal shelfal 
edge, the axis of spirialing ol its shelf off center, and 
external ribbing spiral rather than [)rotractiye. 

Etymology: The genus Garzasia is named for the 
"Garzas Sand" and Garzas Creek, Stanislaus Co., Cali- 

Garzasia intermedia (Cooper, 1894) 
(Figures 51-54) 

Page 130 

THE NAUTILUS, \'ol. 122, No. 3 

Figures 51-58. Species oi Cai~asia new genus. 51-54. Garzasia intermedia (Cooper, 1S94). 51-52. Flasto-leL-totvpe of CASG 
609, Point Loma Formation. 51. Apertural view, vertical dimension 21.5 mm. 52. Apical view, diameter 19 nnn. 53-54. Hypotype 
SDSNH 67149, SDSNH loc. .340.3. 53. .A.pertural view, vertical dimension 61.5 mm, diameter 64 mm. 54. Left-lateral view of spire 
above posterior part of aperture, height 20 mm. diameter 64 mm. 55-58. Gai-asia diahla new species. 55-57. Holotype LACMIP 
13393, LACMIP loc. 2258S. 55. Apertural view, vertical dimension 36 nmi, diameter 37 mm. 56. Abapertural view, vertical 
dimension 34 mm, diameter 37 mm. 57. Side view of spire above posterior pait of aperture, height IS mm. diameter 37 mm. 58. 
Paratype LACMIP 13394, LACMIP loc. 26353, slightly obhque left-lateral view (partial specimen), height 29 mm, diameter 9 mm. 

Stomatia intermedia Cooper, 1894: 46, pi. 3, fig. 43 [re- 
figured in Yates, 1903: pi. 3, fig. 43]. 
Lysis intermedia (Cooper). — Anderson, 1958: 170; Coan, 
1981: lfi.5, fig. 12 [reprint of Cooper's pi. 3]. 

Diagnosis: Large Gai~asia, haliotiform, earinate on 
periplieiy, scuIjDture elsewhere consisting of several low 
spiral cords separated by wide interspaces bearing up to 
10 spiral threads, shelf vers-' broad, with arcuate to 
sliglitlv sigmoidal margin. 

Description: Large size (height up to 37 mm, diam- 
eter 68 mm, same specimen); haliotiform v^dth very rap- 
idly expanding last whorl; protoconch imknowai; spire 
highly variable in elevation, ranging from 50 to 85% of 
total shell height; suture abutting below periphery; 
growth line prosocline; sculpture consisting of commonly 
five spiral cords; peripheiy earinate and bearing stron- 
gest cord; peripheral carina located anteriorly oi medial 
position on spire whorl but medially on last whorl; adapi- 
cal to medial carina two to three medium-strong cords 
with wide interspaces bearing up to 10 spiral threads; just 
abapical to medial carina several closely spaced and scaly 
medium strong cords; aperture circular; inner lip and 
columella flattened to slightly concave and expanded to 
form very wide crescentic shelf submerged within aper- 
ture; posterior end of slielf surrounds umbilicus and spi- 
rals into it. 

Lectotype: CAS 609 (formerly CSMB 13742), height 
6.4 mm, cUameter 22.6 mm (incomplete). 

Tvpe Locality: Point Loma, San Diego Co., Califor- 

Figured Specimen: 


SDSNH 67149 from SDSNH 

Distribution: Point Loma Formation, (area 19 and 20) 
San Diego Co., California. 

Geologic Age: Latest Campanian and early Maas- 
trichtian, Baculifcs hunacnsis zone. 

Discussion: The above description was based on sLx 
specimens, of which presei-vation is generally good to 

Cooper's (1894) figure 43 line drawing is actually a 
combination of three specimens. The lectotvpe (CASG 
609) is designated by us and is pluitograplicd liere (Fig- 
ure 51) for the first time. 

Garzasia intcnncdiii differs from G. ditihla new spe- 
cies in achieving larger size, having weaker carinae \wth 
the medial carina being most prominent, and having spi- 
ral threads in interspaces between carinae. 

Garzasia intermedia is somewhat similar to Sigapatella 
Lesson, 1830, some species of which [e.g., Sigapatella 
novaezelandiae (Lesson, 1830)] have spiral sculpture, an 
off-centered apex, and a well-developed "false" umbili- 

L. R. Saul Mid R. L. Squires, 2008 

Page 131 

cus. Tlie umbilicus ol G/ii'Zdsid appt-ars to lonn as the 
wideK' expauded, llatteued coluuiella is attached uiedi- 
alK' arouud the axis of coiliug loruiiug a hioad shellal 
area. In the holotspe of G. intcniwdia and some other 
specimens this area is an open, funnel shape, but in oth- 
ers the area is more or less iilled, resulting in sheK cs with 
differing "umbilical" dexelopment from a moderate to 
deep depression. The sheltal edge of G(n~zasia differs 
from that of Sigapatclla in being siguioidal latlici- lluiii 

A large specimen SDNHM fS7149/.34():5 has taint 
markings at each end of the shelf which rcscnibK' muscle 

Goi-zasia cliabla new species 
(Figures 55-58) 

Diagncsis: Medium sized, CahjptracaAike. with mod- 
erateK" high spire, having three to four e(|ual-strength 
carinate ribs with wide interspaces. 

Description: Medium to moderateh' large size (up to 
.30 mm in height), Cali/ptmcaAike shell with rapidlv ex- 
panding last whorl; spire moderately higli, approximately 
40 to 50% of total shell height; protoconch missing; te- 
leoconch appro.ximateK' 1.5 whorls, carinate; peripheiy 
near one-half whorl height; suture abutting at periphery; 
sculpture consisting generally of three to foiu" equal- 
strength and equall}' spaced carinae with veiy wide in- 
terspaces; aperture circular with its abapertural margin 
shai^pK- demarked and steeply descending; inner lip mar- 
gin slightlv signioidal; posterior end attached to inside of 
outer lip directly beneath suture; shelf spiraling into um- 

Holot)pe: L.\CMIP 13393, height 18 nun, diameter 
37 mm. 

ParaHpe: LACMIP 1.3394 from LACMIP loc. 26353. 

T\pe Locality-: LACMIP loc. 2258S. 

Distribution: Moreno Formation, "Garzas Sand" 
Member, (area 9 and 10) Merced and Stanislaus coun- 
ties. California. 

Geologic Age: "Mid" Maastrichtian. 

Etymology: The species is named for its occurrence in 
the eastern foothills of the Diablo Range, Merced and 
Stanislaus counties, California. The specific epithet clia- 
bla is used as a name in apposition. 

Discussion: The above description is based on eight 
specimens. Preservation of the known specimens of this 
species is poor. Except for the holotype, all the speci- 
mens are internal molds. 

This species is very similar to G. intermedia from 
which it differs in having a generally lower but more 
roundly inflated, spire that is apparently not variable in 
height, higher last whorl, much stronger sculpture, and 
the basal flange wTap into the aperture to meet the ap- 
ertural edge of the shelf. 


The presence of both line-ribbed and coarse-ribbed Ari- 
adnoria and Li/.vi.v suggests that species oi Li/sis may have 
been derived from Ariadnana. During the evolution of 
Lysis, the inner lip broadened (as did the columella) and 
apparently moved deeper into the aperture, thereby al- 
lowing more room for the foot to gi'asp haixl substrate. 
These changes, assumed to be a function of the rapid 
expansion of the aperture, were necessaiy in order that 
the aperture could accommodate a larger foot for attach- 
ment to a hard substrate. The resultant shell shape was 
crepiduliform. Modern trichotropids studied by Yonge 
(1962) in Puget Sound thrive on unstable shell beds. If 
some Cretaceous trichotropids resembled the modern 
trichotropids in (I) being filter feeders. (2) living on firm 
substrates, and (3) being protandrous hermophodrites. 
the)' would have had characteristics that allowed them to 
evolve toward tlie less vagile calyptraeids or crepidulids. 

The inner lip of trichotropids is homologous to the 
internal deck or shelf of calpvtraeids, and the develop- 
ment of the shelf in Li/sis seives as a pattern for the 
development of the flat shelf in Crepidida Lamarck, 
1799, and of the shelf in for the spiral shelf in 
CaJifptraea Lamarck, 1799. Trichotropid and crepidulid 
sculpture is predominantly spiral (i.e., in the direction of 
coiling), but some calyjjtraeid sculpture is radial or pro- 

The shell e.xterior of 7^//,s/.v suciensis group (i.e., Lijsis 
suciensis lineage) is fine-ribbed to relatively smooth; in- 
teriorly the columella and inner lip are flattened and 
broadened into a crescentic shelf which, as it lengthened 
posteriorly, came to resemble a shelf of Crepidida. Ex- 
cept for its large size. Lysis suciensis resembles the ear- 
liest species from the Pacific slope usually assigned to 
Crepidida: namely, 'Crepidida' pdeiiin (Gabb, 1864), 
which ranges from middle Eocene to Oligocene in Cali- 
fornia to Washington. Stewart (1927) and Hoagland 
(1977) slightly extended the range of C (Spirocrypta) 
pileinn by sviionvmizing with it the late earl\' Eocene 
Spirocrtfpta inornaia (Dickerson, 1916) and the middle 
to late Eocene Spiroen/pta diekersoni (Weaver and 
Palmer, 1922), but Yokes (1939) considered Spirocn/pta 
inornata of "Domengine" age to have "a fumiel-like pro- 
cess due to the upward cui^ving of the posterior portion 
of the septa" (Yokes, 1939, pi. 13. fig. 7) rather than the 
less apparent "umbilicus" of S. pdeum (see discussion 

Gabb (1864) originally placed 'Crepidida' pdeum in 
the invalid genus Cri/pta Humphrey, 1797 (a synonym of 
Crepidida), and subgenus Spirocrypta Gabb, 1864, of 
which 'C (S.) pileum is the type species. 'Crepidida' (S.) 
pileum, which is common in the Tejon Formation of 
middle Eocene age in Live Oak Canyon, Kern Co., Cali- 
fornia, has been figured several times (e. g., Gabb, 1864: 
pi. 29, figs. 233, 233a-b; Stewart, 1927: pi. 29, figs. 2-3; 
Anderson and Hanna, 1925: pi. 13, fig. 7; Clark, 1938: pi. 
4, fig. 19; Wenz, 1940: figs. 2660a, b [reprint of Stewart]. 
A large specimen from the late Eocene is figured by 

Page 1.32 

THE NAUTILUS, Vol. 122, No. 3 

Kleinpell and Weaver (1963, pi. 24, fig. 11). Weaver's 
(1943: 724, pi. 71, fig. 16) illustration of Cah/ptraea 
diegoana (Conrad) is a lapsus and is a posterior-end-up, 
apertural view of 'Crcpidiila' pileum. On page 356 he 
correctly lists tlie figure as Crepkhda pdcum. Gabb's 
(1864: pi. 29, fig. 233a) and Stewart's (1927, pi. 29, fig. 3) 
show the siielf Galib's figure is a tascimile and Stewart's 
is a photograph that has been reproduced in other dis- 
cussions of Spirpcnjpta (e.g., Wenz, 1940: 903, fig. 
2660a). Gabb's and Stewart's figures are based on lecto- 
ty}3e ANSP 4221, but, unfortunately, the shelf of this 
specimen is broken. Both figures create the false impres- 
sion that there is a sinus near both ends of the shelf and 
that the middle part protrudes and is concaxe. An addi- 
tional representative specimen (hypot\'pe LACIVIIP 
13395) was cleaned by the senior autlior and is illustrated 
in Figures 59-61. Its shelf (Figure 59), which is unbro- 
ken, is slightly sigmoidal and long on the left side (or 
anterior end) and shorter on the right side (or posterior 
end). There is also a slight con\e.\it\' of the somewhat 
sinuous shelf as it approaches the posterior end of the 
aperture and the shelf margin spirals over slightK' to form 
an indication of an umbilicus. In addition, the shelf is also 
narrower and closer to the shell margin cjn the left/ 

anterior and noticeabK' farther from the shell margin 
(deeper into the aperture) on the right/posterior. The 
shelf of "C." pileum thus, as noted by Gabb (1864), spirals 
inward toward the apex. Gabb's figure (pi. 29, fig. 233b) 
in part illustrates this, as does Figure 59. Although Stew- 
art (1927) s)'nonomized Spirocn/pfa with Crcpidida, 
Gabb's description of Spirocrypta recognizes this veiy 
important characteristic, which helps to distinguish Spi- 
rocn/pta from Crcpidida. In modem Crcpidida forni- 
raf/i. the shelf does not spiral into the whorl ape.x. 

Aperture/shelf features of Spirocn/pta pileum and S. 
inoniuia resemble those of the early Paleocene Spirogcd- 
cnis UimcUaria Finlav and Manvick, 1937, from New 
Zealand, in that the shelf of S. JameUaria is also narrower 
and closer to the shell margin on the left/anterior and 
noticeably farther from the shell inaigin (deeper into the 
apertiu-e) on the right/posterior. 

Our proposed evolutionary relationship between tri- 
chotropines and calvptraeids differs from any previous 
author's proposal. Htjagland (1977), for example, in her 
study o{ Crepidula Lamarck, 1799, rejected trichotropids 
as direct ancestors of calvtraeids and crepidulids. Hoag- 
land (1977) opined that although Tricliotropis Broderip 
and Sowerby, 1829, and Crcpiduhi had a common an- 

Figures 59-73. Fossil Calyptraeidae. 59-64. Cx)iiiparative Eocene Crepuliilii and Cah/ptraea species. 59-61. Spirocrijpta pileum 
(Gai)b, LS64), li^^jjotype LACMIP 1.339.5, LACMIP loc. 22.386, vertical dimension 3 mm, diameter 9 mm. 59. Apertural view. 60. 
Abapertnral view. 61. Lateral view. 62-64. ■Calijplraea' diegoana (Conrad, 1855), hyisotyiDe LACMIP 13458, LACMIP loc. 22.340. 
62. Apertural view, vertical dimension 29 mm. diameter 31 mm. 63. Apical vdew, diameter 31 mm. 64. Riglit lateral view, height 13 
mm, diameter 31 mm. 65-67. 'Calt/pfraea' aprria Solander in Brandrr, 17(S6, Inijotspe LACMIP 13.39fi, LACMIP loc. 7333. 65. 
Apertural view, vertical dimension 11 mm, dianiclcf 1 1 nmi. 66. Apical view, diameter II mm. 67. Right-lateral view, height 7 nun, 
diameter 11 mm. 68-70. Cah/ptraen i-liinrii\i.\ (Linnaeus, 1758), Recent, ty|oe-specics, livpotv'pe LACM 161651, Cherbourg 
(Manchc), Normandie, France. 68. Apertural view, vertical dimension 15 mm, diameter 14.5 mm. 69. Apical view, diameter 14. .5 
mm. 70. l-iigl It-lateral view, height 6 mm, diameter 14.5 mm. 71-73. Troeliifa tmcliiforum (Born, 1778), Recent, t)'pe-species, 
LACM loc. lot 75-41, Pumalin, west of IslaTelcon, Gulf of Corcovado, Cliiloc Province, Chile -intertidal 7L Apci jural view, vcitical 
dimension 40 mm, diameter .38 mm. 72. Apical viev^', diameter 38 mm. 73. Right-lateral view, height 18 mm. 

L. R. Saul 

R. L, Squires, 2008 

cestor. Tricltotroj)is lias no din't-l iclalidiiship to C.rcp- 
ichila because Trichotropis is lioreal, living in cokl and 
deep water and has rough sculpture, whereas Crcphliila 
had its origin in the Panamic region and is reiati\el\- 
smooth sculptured. Her implications that Trichotropis is 
a deep, cold-water gastropod and that Crepkluhi is a 
shcillow, warm-water gastropod are misleading. Motlern 
Trichotropis canccUata (Hinds, 1S43) is intertidal in 
.\laska, British Columbia, and Washington to depths of 
104 m off -\laska, 165 m off British Colvunbia, and 274 m 
off Washington (L.\CM Nhilacolog\- collection). In the 
southern part of T. cancclkitii's range, it is in cool- 
temperate, not boreal waters. The ([uestion is, however, 
not where trichotropids li\e now but where the\" were 
during the Late Cretaceous, when there were no polar 
ice caps and the subtropical and temperate zones were 
nnich wider. Trichotropids and calN'ptraeid-like gastro- 
pods co-e.\isted during the Late Cretaceous in the study 
ai-ea, and both forms li\"ed in relatively warm, shallow 
waters. .Although the Coniacian-Santonian faunas of 
northern Ciihiornia are noticeably cooler than Turonian 
faunas (Saul and Squires, 2008) oi that area, they would 
have been temperate. Hoagland (1977) mentioned that 
anatomical features of Trichotropis suggest affinities to 
cal\ptraeids, but she believed that similarities of life his- 
toid' and niche between Tricliotropis and Crcpichda aix' 
com'ergent. She suggested that Crcpichda is (leri\ed 
from some form of "calvptraeid stock" that, in turn, was 
deri\'ed from Trochita Schumacher, 1817, reported by 
Wenz (1940) to range from Eocene to Recent. She did 
not pro\ide an\' geologic time parameters as to when 
these deri\'ations took place, but indicated that Shimer 
and Shrock (1959) recorded the first caK-ptraeids as 
"low^er Cretaceous." The "lower" seems to be a minor 
lapsus; all printings of Shimer and Shrock from first 1944 
to last 1989 hst range oi CaJijptraea and Crcpichda as "U. 
Cret.-Recent" and Cmcibidum as "Tert. -Recent." Wenz 
(1940: 902), however, questionably included the peculiar 
looking genus Galcricuhis Seelev, 1861 from the Upper 
Creensand (Lower Cretaceous upper Albian) oi England 
in cal\ptraeids. It does not resemble any calyptraeid we 
have studied. Seelev (1861) named Crcpidida cooksoniae 
also from the Upper Greensand, hut Hoagland (1977: 
395) found it "unconvincing" as a calvptraeid. 

The Campanian to Maastrichtian Damesia Holzapiel, 
1888, of Europe and Tennessee, has been regarded as a 
cal\ptraeid bv some workers (e.g., Sohl, 1960), but Dock- 
en" (199.3) assigned Damesia to the pvramidelloids. 

Bandel and Riedel (1994) reviewed placement and 
content of Cal\traeoidea, and in comparison to Hoag- 
land's (1977) study, they arrived at a different familial 
content and different relationships between the famihes. 
Their CaKptraeoidea consisted of two families: Calyp- 
traeidae containing genera Cahjptraca, Cmcibidum, and 
Crcpidida; and Hipponicidae containing Cheilea, Hip- 
ponix. Neojanaca. and Thijlacus; but both families ex- 
cluded trichotropids. According to Bandel and Riedel 
(1994). all genera in Calyptraeidae lack a probosis and 

are obligalorN liller leeders. However, in Hawaii Ulbrick 
(1969) repoitcd algae grazing, in aildition to filter feed- 
ing by Cnicihidiiui spinosiini Sowerbv, 1824. 

Cretaceous trichotropines probably had several char- 
acteristics that lysines would have inherited from them: 
1) filter feeding, 2) living epifaunally, probably on a hard 
substrate, 3) brooding egg capsules from which plank- 
totrophic laiAae hatch, and 4) being protandrous her- 
maphrodites. Characteristics the lysines would pass on to 
cahptraeids. L\'sines" first recognizable difference from 
the trichotropines is the increase in expansion rate of the 
whorl, especially of the last whorl, resulting in an en- 
larged aperture in which the columella is broadened. The 
earliest broadening and enlargement were moderate and 
only become striking as geologically younger species be- 
gan to ha\'e a very distinctive moiphologv. Until con- 
nected with its ancestry, Li/sis was difficult to classify. 
The shell also liecame more flattened and limpet-like, as 
the aperture enlarged. 

Hoagland (1977) credited Crcpichda hochstcttcriana 
Wilckens (1922: 5-6, pi. 1, figs. 9a, b) as being the ear- 
liest Crcpidida because it was reported from the calcar- 
eous conglomerate stratigrapliic unit of the Lower 
"Amuri Group" in the "Amuri Bluff area, northeastern 
South Island, New Zealand (Wilckens, 1922: Warren and 
Speden, 1978). Modem speUing of "Amuri" is Haumuri, 
and the Cretaceous strata are referred to the Mata Se- 
ries. Woods (1917) reported that these strata at Haumuri 
Bluff contain the bivalves Inoccranms austrcdis Woods, 
1917 and Inoceramus pacificus Woods, 1917, which ac- 
cording to Wellman (1959) are limited to the Piripauan 
Stage of latest Coniacian to Santonian age. 

Warren and Speden (1978) noted "problems" with the 
early collections from this area but nevertheless, listed 
Maoiicnjpta hochstcttcriana (Wilkens, 1922) from the 
Campanian Okarahia Sandstone of the Mata Series. De- 
scribed conditions of collecting suggested a strong pos- 
sibility of co-mingling of material from different strata 
and that the only knowai specimens oiM. hochstcttcriana 
might not be from the Mata Series. In search of tlie t\pe 
specimen, we contacted three extraodinariK helpful 
New Zealanders: A. G. Beu, A. Grebneff and J. D. Stil- 
well. Their email communications (2006) indicated that 
M. hochstcttcriana is not from the Okandiia Sandstone, 
and is not of Cretaceous age. Beu found the t>pe speci- 
men (GNS TM2608) in the New Zealand Geological 
Sun-ey collections. Fortunately, the t\pe specimen was in 
a large enougli block of matrix to take a sample for mi- 
crofossil dating. He enlisted the aid of G. Wilson who 
dated the dinoflagellates as late Oligocene, at the oldest. 
Ian Raine, who looked at the rich spore-pollen assem- 
blage also from the sample, found Acacia pollen, which is 
not known earlier than Miocene in New Zealand. Mio- 
cene strata crop out above the Mata Series, and some of 
the original material sent to Wilkens was apparently from 
beach boulders derived from younger strata oyerl)ing the 
Cretaceous Mata Series. This Neogene age for Wilckens' 
species is much more likely considering that the shelf 

Page 134 

THE NAUTILUS, Vol. 122, No. 3 

covers nearly one-half of the aperture (Hoagland, 1977: 

Hoagland published more papers on Civpkhihi and, in 
19S6, she revised several items ot her 1977 paper l)ut did 
not design a new evolutionaiy pathway. We differ from 
Hoagland (1977) in that we believe it is difficult to go 
from the centrally oriented ape.x plan oi Trochita (Figure 
71) or Cahjptraea (Figure 68) with its spiraling shelf and 
get to Cirpidula with its eccentric spire and apparently 
unspiraled deck. Whether or not Lysis species are an- 
cestral to any modern Crepidiila, younger species oi Ly- 
sis achie\ed a crepiduloid form with a respectable shelf 
by wrapping the posterior end of the shelf onto the inside 
of the outer whorl. Furthermore, from this, the calyp- 
traeid form appears to have evolved with the develop- 
ment of an "umbilicus" in Gai'zasia intermedia making 
Cooper's specific name remarkably prescient. 

At about the Campanian/Maastrichtian boundaiy, de- 
velopment of the spiral shelf of Gai~asia resulted in a 
shelf that appears to be a pattern for development of 
shelves in Cahjptraea and perhaps Cnicihidiint. The very 
broad, depressed spiraling inner lip of Garzasia appears 
to pro\dde a likely pattern foi- development cjf the shelf of 
Cahjptraea or Cnicibidiiin, not for the more decklike 
shelf of Crepidiihi. 

The earliest reported Cahjptraea on the Pacific slope 
is 'Cahjptraea' diegoana (Conrad, 1855) which ranges 
from middle Paleocene to Oligocene and occurs from 
California to Washington and easternmost Russia 
(Squires, 1987). A representative specimen from the Te- 
jon Formation Eocene is showai in Figures fi2-64. Figure 
62 displavs the shelf of this species, which is similar to 
that of 'Cahjptraea' aperta Solander in Brander, 1766 
from the Eocene of Europe (Figures 65-67). Both of 
these species have often been referred to Trochita Schu- 
macher, 1817 (type species Turbo trochifonnis Born, 
1778), but their shelves (Figures 62 and 65) spiral from 
an off-center position roughlv a quarter of the diameter 
in from the apertiu'e edge. A so-called pseudoumbilicus 
at the upper end of the shelves in Figures 62 and 65 
could result from reduction of an unilnlicns such as that 
of Garzasia. The shelves of 'C chegoana and 'C aperta 
are narrower than shelves of Garzasia and expand across 
about one third of the aperture. They differ distinctly 
from that oi Trochita (Figures 71-73), which has a sturdy 
spiraling internal shelf extending from the centered axis 
of coiling to the outer shell margin, gi^'ing the impression 
of dividing the circular aperture in half. 

'Cahjptraea' diegoana does not appear to be a direct 
descendent of Garzasia. Its similarity to 'C aperta sug- 
gests an ancestor in the Old World Tethyan Sea and, like 
many of the other Early Cenozoic mollusks (Squires, 
1987, 2003), it probably arrived onto the Pacific slope via 
a circum-equatorial current. These species 'C diegoana 
and 'C aperta do not have shelves similar to that of 
Cahjptraea chinensis (Linnaeus, 1758), the type species 
ol Cahjptraea. In C. chinensis, the thin fragile shelf (Fig- 
ure 68) occupies about a quarter of the apertural circle. 

It arises from the apex with a folded-over edge that forms 
the umbilicus. At the open end of the umbilicus, the 
shelf edge abruptly veers counter to coiling direction and 
approaches the apertural margin at an acute angle. The 
total shelfal edge is sickle-shaped wdth a short handle 
(the umbilical edge) and a long cuived h\dde (the outer 
margin of the shelf^ This leaves a deep notch between 
the attachment of the shelf to the shell and a delicate, 
loliate shelf 

Trochita is present in modern eastern Pacific slope 
faunas from Mazatlan, Mexico to Valpariso, central 
Chile. It occurs in lower to middle Miocene strata as far 
north as the La Panza Range, San Luis Obispo Co., is 
present in the Keni River section, Kern Co. (Addicott, 
1970) and the Topanga Formation, Santa Monica Moun- 
tains, Los Angeles Co., California. It has a thick shell 
with protractive ribbing and, as mentioned above, a 
sturdy shelf. 


Both 'Crepichda' and 'Cahjptraea' have been reported 
from Campanian and Maastrichtian age strata from 
throughout the world. Presei'vation of most specimens 
makes identification of them problematical. Some of 
these gastropotls have proven to belong to other families 
or not to be of Cretaceous age. Others need verification. 
Our tally of occurrences is doubtless incomplete. 

Although stated above as "throughout the world," 
these gastropods are recovered from areas that were 
probably temperate to tropical. Verified lysines are all 
younger than Turonian and older than Eocene. Classing 
Spirogahienis as a Ivsine provides the only Paleocene 
record of this subfamily. Late Cretaceous occurrences 
suggest that the calyptraeids developed in several geo- 
graphic places from widely distributed trichotropids 
evoKing into Lysinae. An example of this is the evolving 
shape of Lysis. Although on the Pacific Slope caKptraei- 
form calyptraeoids developed from a crepiduliform ca- 
lyptraeoid, the calvptraeiform has been recognized more 
widely geographically, but not earlier than Coniacian. 

Cretaceous calvptraeiform calyptraeoids are more 
widely recognized geographically than are crepiduliform 
calyptraeoids. Europe has calvptraeiform occurrences 
but no crepiduliform occurrences. Africa has calyptraei- 
form occurrences in the north and crepiduliform occur- 
rences in the south. African occuri-ences ai'e close in time 
to those of the Pacific Slope. North America (exclusive of 
the Pacific Slope) has a veiy few reports along the Gulf 
Coast of calyptraeiform specimens. The Pacific Slope has 
both calyptraeiform Gai'za.sia and crepiduliform Lysis. 
South America has calyptraeiform species. Japan some- 
what mirrors tlie Pacific Slope. Its knowm Ivsine is of 
early Maastrichtian age, but the additional presence of a 
trichotnpid (Kase, 1990) similar to A. ohsiricta suggests 
that lysines were developing tliere roughlv synchronously 
with those along the Pacific Slope. New Zealand has no 

L. R. Saul and R. L. Squires, 2008 

Page 135 

R15 100 9=; 90 8S 80 75 70 (iS 60 55 50 45 40 35 30 25 20 15 10 
I ' ' ' I ' ■ ' ' I ' ' ' ■ I ■ ■ I , ■ . . I . , , . I , . . , I , , , . I , , ■ , I , , I , I . ■ I I . , , , I , . , , I , , , , I , , , , I , , . , I , , , . I 









E O t E N E 









Ariadnaria ■ 

Calyfitraea ' 












I Maoricrypta 

2 muscle scars 

? Crpndicrepidula 
? Crepldula 

■ Crepipatella 

Figure 74. Generalized proposed evolutionary trends ot calyptraeoids. Time scale after Gradstein et al., 2004. Galyptraeids and 
crepidulids probably e\'olv-ed from several trichotropids. Lysis or Lysis-like fossils of Coniacian to Maastrichtian age have been 
described from California, and from Campanian to Maastrichtian age from southern Africa and Japan. 1. Turriculate gastropods 
ha\ing gill capable of filter feeding, sedentary' adult life on hard substrate. Probably capable of copulation, protandrism, and brooding 
ot \oung. 2. Broadening ot columella and inner lip. Enlarging final whorl. 3. Attacliment of posterior shelf end to interior of outer 
lip to develop crescentic shelf 4. Some species developed "umbilicus" in spiraling slielf as in Gai-asia. 5. "Umbilicus" closed or nearly 
so in Eocene 'Crepidula' and mo\ing toward more central position in more circular base. 6. Broad shelf spiraling from near center 
of base. Shelf edge nearly straight. 7. Spiral shell with low spire, round base. Shelf edge extremely sigmoidal. °^ = Pleistocene. Stages 
abbreviated are Turonian, Coniacian, Santonian, Maastrichtian. 

known Cretaceou,s h^sines but does have the v'oungest 
known calvptraeiform lysine. 

EUROPE: Crepidula mytiloidea Bellardi and Mich- 
elotti, 1840 from Villavemia near Tortona Italy was listed 
among Nomina Dubia bv Hoagland (1977) as being from 
the Cretaceous. Bellardi and Michelotti were describing 
a Tertian' fauna. Because the specimen and illustrations 
are so small, Hoagland could not verih' that this species 
was a Crepidida. 

Calyptraea cretacea (crOrbignv, 1842: 390, pi. 234, 
figs. 1-3) [Infiindibulum] was e.xamined by Kollmann 
(2005) who determined that d'Orbigny's material was 
from Campanian of Charente-Maritime, southern 
France, and that the specimen identified as C. cretacea 
bv Delpev (1942: 165, fig. 1) was from Maastrichtian of 
southwestern France. Poor preservation of d'Orbigny's 
t\"pe caused Kollmann (2005: 172, pi. 18, fig. 18) to refer 
to it as "Calyptraea s. lato cretacea (d'Orbignv, 1843), 
species dubia." Delpev's illustration shows no shell. Koll- 
mann considered both specimens to be only Calyptraea 
sensu lato. These specimens expand their whorl diameter 
much less rapidly and have much higher spires than 
specimens of Lysis and Garzasia from the Pacific Slope 
of North America. 

Calyptraea depressa Delpey, 1942 (p. 165, fig. 2) from 

Maastrichtian southwestern France, has a spiraling shelf 
similar to tliat of Eocene 'Calyptraea' aperta e.xcept that 
its shelfal margin is concavely arcuate betvveen rim and 
whorl center. 

Kollmann and Odin (2001: 446, pi. 1, figs. 18-19) re- 
corded Cahjptraea sp. of Maastrichtian age from south- 
western France, but the preserx-ation of the specimens 
appears to be too pcjor to allow positive generic identi- 

AFRICA: As figured, 'Calyptraea' houci (Fenin- 
quiere, 1912: 10-11, pi. 1, figs. 7-11) from the Maas- 
trichtian of Tunisia externally resembles some forms as- 
signed to 'C aperta. The shape of the shelf of "C houei 
shown in figure 9 is difficult to determine, but appears to 
have a straight edge and probably resembles that of 'C 
aperta . 

Galcrus libyca Quaas, 1902, was described from upper 
Maastrichtian/ possibly Danian strata {Exogyra overwegi 
beds) from the Ammonite Hills, Great Sand Sea, Egvpt 
(Quaas, 1902: 238, pi. 25, figs. 26-29). It has been re- 
ported also from the Congo (Dartevelle and Brebion, 
1956: 29-30, pi. 1, figs. 9-10), and from Libya, Egypt, 
Congo, and Madagascar (Bandel and Riedel, 1994: 339- 
340, pi. 7, figs. 2-3). Bandel and Riedel reported Quaas' 
original specimens lost in World War II and figured the 

Pa^e 1.36 

THE NAUTILUS, Vol. 122, No. 3 

exterior of a subsequentlv collected specimen of 'Calyp- 
traea" libijca which resembles "C apei-ta. They did not 
figure the shelf, but described it too briefly as "a flat 
spiral shelf like that of modern Cali/ptraca." 

A Crepklida chain was reported In- Brebion (1956) in 
describing Crcpidida congolcnsis Brebion, 1956, from 
the upper Campanian of the Congo, Africa. This Ahican 
species resembles a Lijsis more than it does a modem 
Crepidula, in that C. congolensis has coiling similar to 
Lysis and a depressed inner lip that barely wraps onto 
the labral side of the aperture. Lysis? congolensis (Bre- 
bion, 1956: 89, fig. 1; pi. 1, fig. 7a, 7b) is most similar to 
L. jalamaca in shape and sculpture but appears to have 
much finer ribbing tlian L. j(damaca. 

'Calyptraea' priDiogcnita Kiel and Bandel (2003: 460, 
fig. 4.14—4.16) and Li/sis capcnsis (Rennie, 1930) illus- 
trated by Kiel and Bandel (2003: 460, fig. 6.1-6.2) are 
from the upper Santonian/lower Campanian, Umzamba 
Formation. 'Calyptraea' priinogenita was described from 
a single worn and broken specimen. Its ribbing (except 
on last quarter- whoi'l) is protractive as in TrocJiita, and it 
has a thick shell as does Trochifa. Its whorl shape is more 
trochiform than in Trochita or Cah/ptraca, it consists of 
more whorls than a Trod t da or a Calijptraea. the last 
\\h()rl lacks the notalile enlargement of a Trocliita or a 
Cali/ptraca. and unlike Trochita or Cah/ptraea, it has a 
small open umbilicus and "the columellar hp bears a 
strong plate" (Kiel and Bandel, 2003). 

Lysis capcnsis is \e\y similar in shape and sculpture to 
L. duplicosta , but its inner lip seems narrower and more 
similar to tliat of L. niickci/i. Lt/sis diiplicosla is generallv 
lower spired and has a broader imiei' lip/cokunella. 

Rennie (1945: 50, 116, pi. 3, fig. 10) figured a Calyp- 
traea sp. from the Upper Cretaceous Senonian of An- 
gola, Africa. A more precise age is unknowai and, al- 
though the shape and angle of sutiire in the figure re- 
semble Cah/ptraea. the base and aperture are not 
illustrated, thereby making generic assignment indeter- 

NORTH AMERICA (exclusixc of the Pacific Slope): 
Cnicihuhiiu? sp. of Sohl (1960: pi. 10, fig. 21) is an im- 
mature, incomplete specimen from the uppi-r part of the 
Ripley Formation (Maastrichtian) in Mississippi. Sohl in- 
dicated that its incomplete shell suggested a close rela- 
tionship to Cntcibidum, and that it definitely appeared to 
belong in the Calvptroidea. The specimen is too incom- 
plete to determine its genus, its similarit)' to Cnicihuliim 
could be a result of the way it is broken, but its spire does 
suggest Cal)ptraeoidea. 

Sohl (1960) classed Thi/lacns cretaceous Conrad, 1860 
in Capuhdae, but Dockeiy (1993) moved it to Calyp- 
traeidae and Bandel and Riedel (1994) included Con- 
rad's species in Hipponicidae. It apparently lived at- 
tached to the columella within the apertuic of empty 
gastropod shells. It differs from Lt/sis and Garzasia in the 
way Thylacns muscles were attached. At the end of ju- 
venile coiling and beginning of expansion of the last 
whorl. Thi/laciis deposited left and riglit prongs instead 

of modifving the inner lip/columella into a shelf as in 
Lysis. Sohl {I960: pi. 10, fig. 4) and (Dockeiy (1993: 
pi. 18, figs. 1 and 4) provided good illustrations of the 
early development which does seem more hipponicid 
than calvptraeid. ^\'hen developed to niaturitx a horse- 
shoe-shaped muscle scar resulted. Dockery's (1993) 
specimens were from the Coffee Sand of Campanian age 
and Sohfs (1960) from the Riplev Formation of Maas- 
trichtian age. 

SOUTH AMERICA: 'Calyptraea' aperta (Solander in 
Brander, 1766), a European Eocene species (see Figures 
65-67 for a representative late Eocene specimen from 
the Paris Basin, France), was identified bv Olsson (1944: 
248-249, pi. 9, figs. 10-13) from northern Peni. Olsson's 
placement of the Tortuga fossil beds below his Radiolite 
sandstones with Bacnhtcs suggests a Maastrichtian age. 
As he provided only exterior \dews and no description of 
the shelf, this identification needs further verification. 
Specimens identified as 'Cah/ptraea' aperta from Europe 
and the American Gulf Coast range widely as to whorl 
height and sculpture which varies from smooth to spi- 
nose. The Peruvian specimens increase in diameter at a 
slower rate and thev have more strongK impressed su- 
tures than Eocene specimens, suggesting that the Peru- 
vian specimens are probabK' not C. aperta. 

Cah/ptraca lacvis (Philippi, 1887) from Maastrichtian 
of central Chile was described as Trochita lacvis Philippi 
(1887: 92, pi. 11, fig. 3; referred to Galeropsis by W'ilck- 
ens (1904: 195-196, pi. 17, figs. 9a, b) because of its high 
spire, and to Calyptraea (Trochita) by Bandel and Stin- 
nesbeck (2000: 763-764, pi. I. fig. C). Bandel and Stin- 
nesbeck provided a view of the exterior only, and de- 
scribed the shelf as "a flat concave shell like that of mod- 
ern Cah/ptraca." Although "fiat concave shell" might 
partially describe a Trochita shelf, it is not an acurate 
description of tlie shelf in C(di/ptraca chincnsis (Figure 
68). Wilckens (1907: 13, pi. 3,' fig. 6) also reported'Cw- 
lyptraea aft. lacvis Pliilippi of Maastrichtian age from 
soutliern Patagonia, but the pi'esenation of the specimen 
appears to be too poor to allov\' positive generic identi- 

C(dyptraca pdeolus d'Orbigny, 1841 was indicated by 
Hoagland (1977: 354) to have been recorded from Lower 
Cretaceous strata of Argentina by von Ihering (1907), but 
the species was listed by FerugHo (1937: 187) from the 
Patagoniano (Tertian) of Lago Argentino, Argentina. 

JAPAN: As noted earher. Lysis iziiniiensis Kase, 1990 
has been described from the earliest Maastrichtian of 
Japan. It resembles the group of Lysis duplicosta and 
appears from the illustrations (Kase, 1990: figs. 2.11- 
2.12) to be most similar to L. duplicosta in strength of 
sculpture and development of innerlip/eoluniellar expan- 
sion. Kase (1990) also recorded a trichotropid similar to 
Ariadnaria obstricta liom the Izumi Group in Japan. 

NEW ZEALAND: The early Paleocene (Danian) 
Spirogalaenis Finlay and Manvick, 1937, from New Zt'a- 
land is Lysis-like, especially as to growth line, and diis 
simikiritv reriuires further studv to determine whetlici' it 

L. R. Saul and R. L. Squires, 2008 

Page 137 

results from common ancestn-, con\'ergent evolution, or 
parallel e\'olution, Finla\' and Mannck (1937) assigned 
Spirogalcni.s to famih' CaKptraeidae and based their ge- 
nus on Spirogalerus lamellaiia Finlav and Man\'ick, 
1937, u-hich resembles L. sucicnsis except that L. siicien- 
sis lacks the "pseudounibilicus" described for Spirogal- 
erus. Finlav and Man\ick (1937) and Boshier (1960) 
opined diat Spirogalcnis lamcUaria could represent die 
e\'olutionai"\' link between crepidulids and calyptraeiform 
SigapatcIIa Lesson. 1830. 

Classification of Spirogalcnis has been inconsistent. 
\\'enz (1940) made it a subgenus oi Cahjptraea Lamarck, 
1799. Beu and Mawvell (1990) made Spirogalcnis a sub- 
genus of die calN-ptraeid SigapatcIIa Lesson, 1830, but 
Stilwell and Zinsmeister (1992) separated SigapatcIIa 
from Spirogalcnis because the latter has a strongly ex- 
erted spire. Collin (2003a) noted drat SigapatcIIa has a 
shell and anatomy similar to die cal)'ptraeid genera Ca- 
hiptraca, Trochita Schumacher, 1817, and Zcgalenis 
Finlaw 1926. Collin (2003b) considered SigapatcIIa to be 
monoph\"letic but did not mention Spirogalcnis. Mar- 
shall (2003) recognized SigapatcIIa (=Zegalcnis) as a dis- 
tinct genus based on its shelf edge being broadly and 
e\enh' conca\'e instead of sigmoidal. This is a veiy dif- 
ferent shelf than that of Cali/ptraea chincnsis (Figure 
68). The differences between Spirogalcnis and Sigapa- 
tcIIa are similar to those found between Li/sis and Gar- 
zasia. and the New Zealand forms also record an e\-olu- 
tionaiA- pattern of enlargement of the last whorl and the 
aperture, \\'ith broadening of the columella/inner lip area 
into a shelf \\ithin a hmpetiform shell. Thus, these two 
genera were probablv not links from actual crepidulids, 
but represent links from Ivsines to calvptraeids. 


\\'e mention again the extraorclinan' assistance of the 
Ne\\' Zealanders in documenting a Tertian' age for a 
specimen mistakenly recorded as Cretaceous. Malcolm 
G. Laird, University of Canterbuiy, Christchurch, New 
Zealand, and Jeffrey D. Stilwell. Monash University, A. 
Grebneff, Otago Universitv, Dunedin, New Zealand, and 
A. G. Beu, GNS Science, New Zealand, provided help in 
researching the age of the New Zealand Cretaceous 
units. Thev recruited micropaleontologists Graeme Wil- 
son, GNS Science, New Zealand, who provided an age 
range based on the diatoms, and Ian Raine, GNS Sci- 
ence, Ne^\" Zealand, who recognized the Acacia pollen 
and reported its maximum age in New Zealand. Ray- 
mond Graham, Roval British Columbia Museum, Victo- 
ria, Canada pro\ided infonnarion on the occurrence of 
Lysis diiplicosta in the Gulf Islands area. Lindsey 
Groves, LACM Malacology, provided specimens of Ca- 
hjptraca chincnsis and Trochita trochifonnis and assis- 
tance in finding several obscure references. We also 
thank an anon\inous re\iewer and S. Kiel of University of 
Leeds. L'.K. for improvements in the text. 


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Localities are Lt\CN!lP. unless othenvise noted. .\11 quadrangle maps 
are U. S. Geological Sun'e\' maps. Bracketed numbers are areas shown 
on Figure 1. 

[4] CASG 61794. [= CAS 1346-A]. In conglomeratic sandstone 1.6 km 
(1 mi.) aboxe moutli of Hiding Creek, North Fork Cottonwood Creek, 
Ono Quadrangle (1.5 minute, 1952), Shasta Co., California. Budden 
Can\on Fomiation, Bald HiUs VIember. Late Albian. 

7333. Le Faxel. Paris Basin, France, Late Eocene (Bartonian Stage). 

[IS] 1009-5. [=CIT 83]. Fine sandstone just above shale, sectionline 
fence gate on old road 0.4 km (1/4 mi.) W of Schulz Ranch, 122 m (400 
ft.) S. of nordieast comer of section 19, T. 5 S, R. 7 W, El Tore 
Quadrangle (7.5 minute. 1968), soudi side of Williams Canyon, Santa 
-Ana Mountains. Orange Co.. Cahfoniia. CoU.: B. N. Moore, 1 Januar)', 
1926. Ladd Fomiation. uppennost Holz Shale Member. Early Cam- 

[16] 10711. [=CIT 11.58]. Appro-ximately 1.84 km (1.5 mi.) due west of 
Los ,\ngeles-^>ntura Co. line on die boundaiy (extended) between T. 
1 X and T. 2 X, nordi bank of Bell Canyon, southeast slope of Simi 
Hills, Calabasas Quadrangle (7.5 minute. 1952). X'entura Co., Califor- 
nia. CoU.: \\". P. Popenoe. 18 Julv. 1935. Cliatsworth Formation. 
Middle Campanian. 

[3] 10757. [=CIT 1593]. Massi\e sandstones in bed of North Fork Bear 
Creek, appro.ximatelv 777 m (2550 ft.) soudi and 533 m (1750 ft) east 
of northeast comer of section 5. T. 31 N, R. 1 E, Wliitmore Quadrangle 
(15 minute. 1956). Shasta County, CaUfomia. CoU.: W. P. Popenoe and 
\V. M. ToveU, 12 Sept.. 1941. Redding Formation, Bear Creek Sand- 
stone. Coniacian. 

[5] 10846. [=C1T 1014]. Concretions in sandstone, right bank ot Chico 
Creek about 1.6 km (1 mi.) upstream from the httle bridge across creek 
below Mickev house and about 4.16 km (2.5 mi.) N6°W of 14-mile 
house on Humboldt Road. NA\- Vi, SE 1/4 of section 1. T. 23 N, R. 2 
E. Paradise Quadrangle (15 minute. 1953), Butte Co., CaUfomia. CoU: 
\V. P. Popenoe and D. W. Scharf. 16 August. 1931. Cliico Fomiation, 
top of Ponderosa \\'a\ Member. Late Coniacian or early Santonian. 

[23] 11944. Approximately 10 km X of Punta Abreojos (SW of San 
Ignacio). in first ridges X of arroyo diat crosses Punta Abreojos road 

(cUrt) just S of Campo Rene turnoff, approx. 2 km NW of road. Approx. 
2-3 fall up ravine, liiU .5 km to east is pachydiscid loc, Viscaiano Penin- 
sula, Baja Ciilifoniia Sur, Mexico. CoU.: R. Demetrion, 1987, 1989. X'iiUe 
Formation. Middle? Campanian, with Hoplitoplaceirticeras? 

[14] 14310. About 450m nortli and 70m west of lookout at summit of 
Warm Springs Mountain at elev. of 1052 m (3450 ft.), at base of Kirby's 
(1991, M, A. Tliesis) measured section no. 2, Wami Springs Mountain 
Quadrangle (1958), Los Angeles Count)', Califomia. Coll.: M. X, Kirb\', 
Bas;il San Francisquito Formation. Late Maastrichtian, 

22340. Gritt\' conglomeratic sandstone lenses in fine-grained sandstone 
and shales, east side Grapevine Canyon about 0.8 km (0.5 mi.) south of 
its north end, about 0.4 km (0.25 mi.) east of, and 30.48 m (100 ft.) 
above tlie abandoned highway roadbed on east side of canyon, at about 
648 m (2100 ft.) contour, about 91 m (300 ft.) nordiwest of TejoiV 
granite fault contact. Localit)- is 304S m (10.000 ft.) N24°W of 3174 ft. 
BenchMark at old Fort Tejon, Tejon Quadrangle, Kem County, Cali- 
fomia. Coll: W. P. Popenoe, 9 December, 1946. Tejon Formation. 
Middle Eocene. 

22386. Prominent shell bed at crest of ridge on east side of Live Oak 
Canyon, about 0.4 m (0.25 mi.) south of its mouth, T 10 N, R 19 W, 
Pastoria Creek Quadrangle, 7.5', 1958, photorevised 1974, Kern 
County. Califomia. CoU.: W. P. Popenoe, 13 March, 1947. Tejon For- 
mation, Metralla Sandstone Member. Middle to upper Eocene ("Tejon 

[10] 22588. About 2/3 of the wav to the top of a gulh' on soudiwest 
slope ol a noitliwest-trencUng hiU on south side of Garzas Creek, where 
tlie creek enters the San Joaquin Plain, approximately 610 m south and 
183 m west of northeast corner of section 19, T 8 S, R 8 E, Howard 
Ranch Quadrangle, 7.5', 1953, photorevised 1971, west side of San 
Joaquin Valley. Stanislaus Co., CaUfomia. CoU.: W. P. Popenoe and T. 
Susuki, April, 1950. Moreno Formation. "Garzas Sand" member. 
Middle Maastrichtian. 

[4] 23464. [PRl] Up small creek from Suli\'an Ranch Rd. crossing, and 
1.28 km (0.8 mi.) noitli of ranch, near Gas Point Rd., 701 m (2300') N 
75°E from mouth of Huling Creek, 579 m (1900') S, 488 , (1600') E of 
NW comer of section 16, T. 30 N, R. 6 W, Ono Quadrangle ( 15 minute, 
1952), Shasta Co., CaUfomia. CoU.: P. U. Rodda and M. A. Muiphy, 
May 1955. Budden Canyon Formation, B;Jd HUls Member, unit iv in 
Matsumoto, (1960). MidcUe Cenomanian, probably Tunilites costatus 

[5] 23617. Fossil in hard, blue-gray concretion in gray-weathering buff 
sandstone approximately 15.2 m (50 ft.) below highest conglomerate, 
approx. 0.8 km (0.5 mi.) upstream from Mickey house on west side ot 
Cliico Creek, 1.52 iii (5 ft.) above stream, 716.2Sni (2350 ft.) noitli. 
609.6 ni (2000 ft.) west of soudieast comer of section 1, T. 23 N, R. 2 
E, Paradise Quadrangle (15 minute, 1953), Butte Co., California. CoU.: 
R. B. Saul, 14 August, 1955. Chico Formation, top of Ponderosa Way 
Member. Late Coniacian or early Santonian. 

[5] 23639. In concretions in massive, greenish-gray sandstone, east 
bank of Chico Creek, west of meadow with large flat-topped, lava block 
at north edge near road, 373.38 ni (1225 ft.) soudi and 292.6 m (960 ft.) 
west of norfieast comer of section 23, T. 23 N, R. 2 E, Paradise 
Quadrangle (15 minute, 1953), Butte County, noitliem Califomia. Col- 
lectors: L. R. Saul and R. B. Saul, 20 August, 1952, Chico Formation, 
lowermost part of Ten Mile Member Earlv Campanian. 

[12] 24122. Fine- to coarse grained buff sandstone; 76.2 ni (250 ft,) 
noiili of jeep trail in Jalama Canyon; elevation 190 m (625 ft.), 6.58 km 
(4.11 mi.) east and 1.1 km (0.69 mi.) south of Jalama Ranch Headquar- 
ters; 0.93 km (0.58 mi.) west and 0.66 km (0.41 mi.) north of southeast 
comer of topo, Lompoc HiUs Quadrangle (7.5 minute, 1959), Santa 
Barbara Co., Califomia. Coll.: D. Dailey. August, 1959. Jalama Forma- 
tion. Late Campanian-early Maastrichtian. 

[12] 24128. Dark gray conglomerate in first small canyon east of Ra- 
niajal Canyon, elevation 167.6 m (550 ft.), 0.54 km (0.34 mi) soudi, 3.25 
km (2.03 mi.) east of Jalama Ranch Headquarters, 1.22 km (0.76 mi.) 
north 4.27 km (2.67 mi.) west of soudieast comer of Lompoc HiUs 
Quadrangle (7.5 minute, 1959), Santa Barbara Co., CaUfomia. Coll.: D. 

Page 142 

THE NAUTILUS, Vol. 122, No. .3 

Diiiley, August 19.58. Jalama Formation. Late Campanian-early Maa.s- 

[12] 24237. Medium-grained, buff, arkosic sandstone, 396.2 m (1300 
ft.) nordi of Jalama Creek, elevation 160 m (525 ft.), 0.48 km (0..30 mi.) 
south, 3 km (1.S8 mi.) east of die Jalama Ranch Headquarters, 1.28 km 
(O.SO mi.) north, 4.59 km (2.S7 mi.) west of southeast comer of Lompoc 
Hills Quadrangle (7.5 minute, 1959), Santa Barbara Co., California. 
Coll.: VV. P. Popenoe, September 1938. [alama Formation. Late Cam- 
panian-early Maastrichtian. 

[6] 24340. Penz vicinity, conglomerate beds cropping out just below a 
drainage canal, southeast side of new Oro\ille Hw)', about 1.2 km (0.75 
mi.) northeast of intersection with Pentz-Magalia-Oroville road, 426.7 
m (1400') S., 182.9 m (600') W of the northeast corner of section 36, T. 
21 N, R. 3 E, Cherokee quad. (7.5 minute, 1949), Butte Co.. California. 
Coll.: W. P. Popenoe, 1960, Chico Formation, Musty Buck Member. 
Early Campanian. 

[8] 24349. [ = USGS M8601 and USGS M8745]. In place? large angular 
block of sandstone surrounded by sand at shorehne in covelet on north 
side of elongate seaward-pointing rock; approx. N30°W of Pigeon Point 
lighthouse, just south of Bolsa Point, Pigeon Point Quadrangle (7.5 
minute, 1952), San Mateo County, California. Coll.: L. R. and R. B. 
Saul, October 11, 1960. Pigeon Point Fm. Middle Campanian. 

[16] 26020. [=CIT 11.58]. Hard sandstone slabs in hne-grained sand- 
stone, cropping out on high bare clifi, north bank of Bell Canyon, just 
east of mouth of large gully, and 152.4 m (500') S, 2743.3 (9o6o') west 
of northeast corner of section 4, T. 1 N, R. 17 W, Calabasas Quadrangle 
(7.5 minute, 19.52), Simi Hills, Ventura Co., California. Coll.: W. P. 
Popenoe, 11 Feb., 1972. Chatsworth Formation. Middle Campanian, 

[10] 26353. Approximately 1.2 km south ol Garzas Creek, 671 m south 
and 114 m of northwest comer of section 20, T 8 S, R 8 E, Howard 
Ranch Quadrangle, 7.5', 1953, photorevised 1971, Stanislaus County, 
California. Coll.: R. B, Stewart and W, P. Popenoe, 1944. Moreno 
Formation, "Garzas Sand" member. Late early to early late Maastrich- 

[11] 26370, Reworked fossiliferous Turonian blocks in upper Campa- 
nian conglomerate lens in shale, northeast side of Cooper Canyon, 
approx. 411.5 m (1350') n, 670.6 m (2200') W of soutlieast comer of 
section 2, T. 21 S, R. 14 E, Alcalde Hills Quadrangle (7.5 minute, 
1969), Alcalde Hills, Fresno Co., Califomia. Coll.: J. Alderson, 18 June, 

1977. Panoche Formation, "Alcalde Shale" Member. Late Turonian 
(with juvenile Subpiionocyclus sp), 

[18] 26951, Small sandstone lens approx. 6.1 m (20 ft) above road in 
roadcut on north side of Silverado Truck Trail, 274,3 m (900 ft.) south 
of northeast corner of section IS, T. 5 S, R. 7 W, El Toro Quadrangle 
(7,5 minute, 1949), Orange Co., Califomia. Coll.: A. A. Almgren, 4 
Dec. 1981. Ladd Formation, uppermost Upper Holz Shale Member. 
Late early Campanian. 

[17] 26967. Small exposure of coarse-grained, poorly sorted sandstone 
at bottom of northwest-flowing tributary to main fork of Garapito 
Creek, 449.6 m (1475 ft.) and" 2835 m '(9380 ft.) east of nortliwest 
comer of section 5, T. 1 S, R. 16 W, Topanga Quadrangle (7.5 minute, 
1952, photorevised, 1981), Santa Monica Mountains, Los Angles 
County, Califomia. Coll.: J. M. Alderson, 31 Dec, 1981. Tuna Canyon 
Formation. Coniacian. 

[3] 28717. South Cow Creek Valley, sandstone associated with con- 
glomerate, lower South Cow Creek Valley, about 152.4 m (500 ft.) 
downstream from old bridge site across creek, and about 1.6 km (1 mi.) 
due west ot liuilchngs on Hunt Ranch, NE 1/4 of section 17, T. 31N, R. 
2W, Millville Quadrangle, Shasta Co., Califomia. Coll.: W. P. Popenoe, 
27 Oct., 1971. Redding Formation, Oak Run Conglomerate Member of 
Haggart, 1986. Late Santonian. 

[19] SDNHM 3403. Taylor Made Golf Facilitx :it Salk Drive and Col- 
lege Blvd.. elevation 451 m (148 ft.) iilong College Blvd., lat, .33°8'25' 
N, long, 117°16'56' W. [in general = SDNHM :3402, 3404, 4071, 4073], 
Carlsbad Research Center, Site 29, San Luis Rey Quadrangle 7.5' 
(1968), San Diego Co., Cahfomia. Coll.: B. O. Riney, 26 April, 1987. 
Point Lonia Formation, Late Campanian/?earl\' Maastrichtian, 

[19] SDNHM 3405. Carisbad Research Center, 1.6 km north and 2.7 
km west of southeast corner of San Luis Rey Quadrangle 7.5' (1968), 
indicated area along west side of College Blvd. starting about 0.32 km 
trom intersection with El Caniino Real extends southward for 0.15 km, 
San Diego Co., Califomia. CoU,: B, O, Riney, 1987. Point Loma For- 
mation. ?Late Campanian/early Maastrichtian. 

[18] UCMP 2167. 3,2 km (2 mi.) N WW of BM 1271, Corona Quad- 
rangle (1902), at a gate about 0.8 km (0.5 mi) below Modjeska Springs 
in Williams Canyon, Santa Ana Mountains, Orange Co., Califomia. 
Ladd Formation, uppermost Holz Shale. Middle'? Campanian, 

THE NAUTILUS 122(3): 143-150, 2008 

Page 143 

Feeding behavior, phylogeny, and toxinology oi Conns fa rviis 
Reeve, 1843 (Gastropoda: Neogastropoda: Conidae) 

Samuel S. Espino 

Institute of Chemistn.' 

Unh'ersitv of tlie Philippines 

Diliman, Quezon Citv 1101, 



Department of Chemistn' 

Um\ersit\' ot the Philippines in the Msayas 

Miag-ao,' Iloilo, PHILIPPINES 

Alan J. Kohn 

Department of Biolog)' 
University of Washington 
Seattle, WA 98185 USA 

James A. Villanueva 

Institute of Chemistry 
University of the Philippines 
Diliman, Quezon City 1101, 

Frank M. Heralde III 

National Institute of Molecular Biolog\' 
and Biotechnolog)- 
Uni\ersit\' of the Philippines 
Diliman. Quezon Cit\- 1101, 

Patrice Showers Comeli 

Department of Biology 

University of Utah 

Salt Lake City, UT 84112 USA 

Gisela P. Concepcion 

Marine Science Institute 
University of the Philippines 
Diliman. Quezon Citv 1101, 

Baldomero M. Olivera' 

Department ol Biolog\ 

Uni\ersit\- of Utali 

Salt Lake City, UT 84112 USA 


Ameurfina D. Santos 

National Institute ot Molecular Biolog)' 
and Biotechnology 
University of tlie Philippines 
Dihman, Quezon City, PHILIPPINES 


The Indo-Pacific cone snail Conns fnrvus Reeve, 1843, is 
shown to be moUuscivorous, based on four Unes of evidence: (1) 
Specimens ofConusfurriis maintained in aquaria were directly 
obser\"ed to attack and devour other gastropods, including 
odier cone snails: (2) Its radular teeth share moiphometric 
characteristics with other known mollusci\'orous Conns and dif- 
fer from those of vermivorous and piscivorous species; (3) Mo- 
lecular ph\logenv places Conus fnrvus within a clade of mol- 
luscivorous species; (4) Sequences of two peptide toxins are 
most similar to conotoxins previously characterized from mol- 
lusci\'orous Conus species; one of these closely related peptides 
was previously shown to block molluscan L-t\pe Ca channels. 
Similar to moUuscivorous species such as Conns oniaria and C, 
lictoriae, C. fnrvus varies widely in shell shape and color pat- 
tern, even within its relatively restricted geographic range. 

Additianal Keyioords: Conoidea, snail-hunting cone snail, radu- 
lar morphology.', conotoxins, RT-PCR, phvlogeny. 


Cone snails are venomous marine gastropods (Kohn, 
1959; Rockel, Kom, and Kohn, 1995), and the major 
biologically active components of their venoms are the 
conotoxins (01i\'era, 2006). tvpicallv small (12-35 amino 

^ Author for correspondence 

acid residues), highly constrained peptides. Many cono- 
toxins discriminate between closely related molecular 
isoforms of a particular ion channel family (Terlau and 
Olivera, 2004); this property makes them valuable tools 
in neuroscience, and confers their therapeutic potential. 
The estimated -100,000 different pharmacologically ac- 
tive peptides present in li\ing Conns venoms comprise a 
very' substantial neuropharmacological resource. The sys- 
tematic study of conoto.xins is a seemingly daunting task 
considering the enormous number oi peptides. As die 
conotoxin sequence database expands, it is becoming 
clear that certain U'pes are likely to be produced by spe- 
cies that specialize on particular prey and that belong to 
particular Conus clades (Olivera, 2002, 2006). Thus, es- 
tablishing both the prey and the phylogenetic relation- 
ships of Conus species provides an important guidepost 
for searching for conoto.xins vvitli certain t\iies of physi- 
ological activities. 

Traditional cone snail taxonomy is based on shell mor- 
phology; this has proved difficult because of the absence 
of resolution and possible convergence of shell charac- 
teristics. None of the intragenic classification schemes 
primarily based on shell shape and sculpture, shell color 
patterns, and radular moiphology has received wide- 
spread acceptance (Duda. Kohn, and Palumbi, 2001). 
Thus, these traditional criteria need to be complemented 
bv other t\'pes of data. 

The specific focus ot this stucK', Conus fun us ("the 

Pase 144 

THE NAUTILUS, \'ol. 122, No. 3 

dark cone") is iiniLSiial in several respects. Althougli it is 
found over a relatively restricted area oi the tropical 
Indo-Pacific (almost all specimens have been collected in 
the Philippines, where it is quite connnon in some lo- 
calities), its shell shape and color pattern \aiY widely 
(Figure 1). In most localities, the t\pical lorm of Cdiuts 
furviis has a browai and white shell. However, in the 
Southwestern Philippines, a remarkable diversity of 
forms occurs, particularly along the island of Palawan 
and in the Sulu Sea (Figure 1). As new localities are 
e.xplored, additional variations are being discovered. In 
the comprehensive treatment of Indo-Pacific Conns spe- 
cies by Rockel et al. (1995), the number of taxa synony- 
mized with Coims furvus was probablv among the high- 
est for any species. The authors discussed about a dozen 
distinct forms which they regard as geographic variations. 
Rockel et al. (1995) concluded that Conus furvus "ap- 
pears to be characterized bv isolation oi local popula- 
tions." In general, the occurrence of sinistral specimens 
in an\- Conus is exceedingly rare; another notable feature 
of C. furvus is that most sinistral Conus specimens col- 
lected in the Pliilippines belong to this species (an ex- 
ample is showTi in Figure 1). 

In a stud\" using the calmodulin intron sequence as a 
ph\logenetic marker, Duda and co-workers (Duda et al., 
2()i)l;' Duda and Kohn, 2005; Duda and Palumbi, 2004) 
grouped C. fuiuus wdth C. littcratus in their phvlogenetic 
tree. They inferred that the hvo species di\erged onlv at 
the beginning of the Pliocene; since C. litteratns is a 
well-established vermivorous species that ieeds on eapi- 
tellid polychaetes (Kohn and Nybakken, 1975), wdth a 
fossil record going to the Lower Miocene, this strongly 
suggested that C. furvus is a vermivorous Conus. 

Purchasers of Conns fnixms in the Guimaras Islands 
public market, in the Philippines, make a soup bv boiling 
the snails with tomatoes, onions, and lemongrass. Onlv 
the portions of C furvus near the foot are eaten; appar- 
ently the sections near the hepatopancreas have a bitter 
taste and grainy texture. An alternative culinaiy use of 
the snails is as an ingredient for a local dish prepared 
with noodles. \'arious other snails are typically sold with 
it: C. radiatns, a piscivorous species, is the most common 
other Conus found for sale with C. furvus at this locality. 

In this paper we present data on prev capture beliavior 
bv C. furvus. the moiphometric characteristics of its 
radular teeth, a phvlogenetic analysis based on standard 
mitochondrial sequences, and the first conotoxin se- 
quences obtained from this species. These data consis- 
tently support the conclusion that Conus furvus is a mol- 
luscivorous species. 


Specimen Collection and Dissection: Specimens of 
C. furvus for DNA extraction were [iiu'chased in a public 
market in Guimaras Island, Philippines. The snails were 
dissected to remove the hepatopancreas for total geno- 
mic DNA extraction and the \'enom duct foi- RNA ex- 

traction. A few specimens were maintained in an 
aquarium. Specimens for radular tooth analyses were 
collected in Coron Harbor and Bobok Island, Palawan, 
Philippines, and presen'ed in alcohol. The shells are 
deposited in the Field Museum of Natural Histoiy, 
Chicago, FMNH 300120, 300121, and 300122. The body 
of specimen No. 300121 is presei"ved in the Bemice P. 
Bishop Museum, Honolulu, Hawaii, BPBM 24S751. 

Analysis of Radular Teeth. Tlie I'adular sacs of four 
specimens were dissected and 4-6 teeth from each 
mounted permanently in polvAinyl lactophenol medium 
on slides. Barb and blade lengths were measured from 
the tip of a tooth as described bv Kohn et al. (1999), 
using an ocular micrometer and Nomarski differential 
interference contrast optics. 

Genomic DNA Extraction: The total genomic DNA 
was extracted using the Xanthogenate DNA Extraction 
Protocol (Tillet and Neilan, 2000), a method based on 
the use of the polysaccharide solubilizing chemical xan- 
thogenate. The mediod is non-toxic and requires no en- 
zymatic or mechanical steps to break cells. 

RNA Extraction: Frozen \enoni duct in RNAZater 
(Ambion, Austin, Te.xas) was thawed in ice prior to RNA 
extraction, and a sample of the tissue (10-15mg) was 
homogenized in 1 mL Trizol reagent (Invitrogen, Carls- 
bad, California). The total RNA was extracted according 
to the mannfactin'er's recommendation. 

Aniplificalioii of the .Milochondrial 12S and 16S 
rRNA and mtCOI Genes: For PCR amplification, 
primer pairs designed to hybridize to a segment of 12S, 
IfiS rRNA genes and mtCOl genes were used. The 
primer sequences used for tlie amplification are listed in 
Table 1. PCR amplifications of the three mitochondrial 
genes were carried out in a 10 |ji.L reaction mixture con- 
taining IX reaction buffer (200mM Tris HCl, 500mM 
KCl, pH 8.4), 0.7 mM of each dNTPs, 0.7 niM of both 
primers, 0.4 unit Taq DNA Polymerase, 3.0 mM MgCU, 
and approximatelv 50.0 ng of template DNA. PCR am- 
plification was carried out for 40 cycles. Denaturation 
was carried out at 94 °C, annealing at 48°-55 °C and 
extension at 72 °C. The PCR products were visualized on 
1% agarose gel containing 0.1 mg EtBr/mL. Multiple or 
smeared PCR products were further purified using the 
WIZ Prep DNA purification kit (Promega, Madison, 
Wisconsin), othei"wise, the terminated reaction was di- 
rectly sequenced. These sequences have been deposited 
at GenBank. 

12S Accession Numbers: inuniiniUs EUfiS2274, anU- 
cus EU682275, handanus EUfi82277, dalVt EU6822S1, 
c'piscopatus EU682283, fliivus EU794315, furvus 
EU682284, geographus EU794316, kintoki EU794317, 
littcratus EU794318, magus EU794319, uuirrru:>rens 
EUfi82288, luonachns EU794320, ohsnirus EU794321, 
oniaria EU6822S9, Irxlllr EU68229fi, tulijui EU794322, 
Virgo EU794323; IfiS .Accession numbers: rnninirnlis 

Espino et al., 2008 

Page 145 

Figure 1. Some of the different \ariations of Conns funus. The brown colored specimens shown in the bottom row. center, are 
similar to those used for this research. Xote the sinistral specimen in the lower left hand corner. 

Page 146 

THE NAUTILUS, Vol. 122, No. 3 

Table 1. Primer pairs used for the aniplitication of the 12S, 16S rRNA gene and mtCOl gene. 




Upstream primer 
Dowmstreani primer 

5' .AGAG(C/T)G(.VG) 








EU682299, aulicus EU794324. haiuhnuis EU794325, 
dalli EUn7S935. episcopates EU()78937. flavus 
EU794326, funus EU6S2301, ^co^raplms EU794327, 
kintohi EU79432S, littcratus EU794329, mao^iis 
EU078939, nuirmorciis EU794330, }iioiiac]ius 
EU078938, obsainis EU794331, omaria EU794332. tex- 
tile EU078936, tidipa EU794333, vir^o EU794334. 

RT-PCR of M and O-superfainily Conotoxin 
Genes: Single-stranded cDNA was s)iithesized Irom 
the total RNA extracted from the snail's venom duct 
using; oligodT primers and Superscript II (Invitrogen, 
Carlsbad, Calilornia) and iollowing the manufacturer's 
protocol. Double-stranded cDNA was synthesized using 
primers targeted to the consen-ed regions of the M- and 
0-superfamil\- genes. The product of the sxiithesis was 
visualized in a 1% agarose gel containing EtBr (O.I mg/ 
mL). The terminated reaction containing the PCR prod- 
uct of the correct size (~0.4Kb) was used for the cloning 
reaction. The cloning reaction followed the manufactur- 
er's protocol for the TOPO-TA cloning kit (Invitrogen 
Life Technologies, Carlsbad, California). Tlie clones 
were screened for inserts of the correct size and were 
cultured on a selective medium and incubated overnight 
with shaking (200 i-pm) at 37 °C. The plasmids from 
these clones were extracted and secjuenced. 

Phylogenetie Analysis: 12S and IfiS rliXA sequences 
were aligned using ClustalX, vl.8 (Jeanmougin, Thomp- 
son, Gouy, Higgins, and Gibson, 1998). The alignments 
were refined by eye using VlacClade 4.08 ( Aladdison and 
Aladdison, 2001). Trees were optimized using the indi- 
vidual rRNA alignments and the concatenated align- 
ments (presented herein). To account for the complexity 
of sequence evolution, final anaKses were restricted to 
model-based maximum likelihood (PAUP4bl0, (Swof- 
ford, 2002)) and Bavesian inference (Huelsenbeck and 
Ronquist, 2001). Each method used maximum likelihood 
parameters describing sequence evohition that were op- 
timized with a GTR+I-hG model that includes six pos- 
sible substitution Upes (GTR) and allows some sites to 
be invariant (I), allows across-site rate heterogeneit)' (G) 
and allows imequal base frequencies. 

The Bayesian analysis ran for 100, 000 generations 
(sampled eveiy 100 generations) with the first 25,000 
generations discarded as burn-in trees. Two MCAICMC 
runs (metropolis-coupled Monte-Carlo markov-chain), 
using four chains each, were used to thoroughlv explore 
tree space. Convergence of the likelihoods was judged 
adequate by monitoring the MrBayes output describing 
the average standard error of the difference (ASED) in 

split frequencies between the two nms and by comparing 
plots of the tree log-likehhood trees from generation 
50,000 to 100. Bv the last generation, the ASED was less 
than 0.004; the plot of likelihoods versus generation had 
stabilized. P'urthermore, the PSRF (Potential scale re- 
duction factor) reached 1.00 for the total tree length and 
for each model parameter. 


Feeding Observations: Individuals oi Cu)uis fitrvus 
were fretjuently obsei'ved to prey on other cone snails in 
the aquarium. Most obsen-ations were made with C. ra- 
diatits as the prev. Upon introduction of prey into an 
aquarium tank with C. ftirvus, inmiediate behavioral re- 
sponses were obsened both from predator and prey. The 
prey immediatelv crawled awav irom C. fiimus and at- 
tempted to buiy itself in the sand. At almost the same 
time, C. fiimis pointed its siphon toward the retreating 
snail and crawled in its cUrection. While still relativelv far 
away from the prey, C. fuixus extended its proboscis, and 
inserted it into the body of its pi'ey with one jerking 
motion. Immediately the prey was obsei'ved to stop mov- 
ing. Conns fnrvns then maneuvered the innnobilized 
prev so that its aperture was turned upwards, away from 
the sand, after which C fnrvns commenced feeding on 
the uptuiiied C radiatns. 

Analysis of Radular Teeth: Radular tooth length 
ranged from 6.4 to 8.9% of shell length (mean = 7.5%) in 
the four specimens examined. Each tooth is armed widi 
a barb that extends for 5% of the tooth length from the 
tip (range 4.6-5.4%), and a blade that extends for 9% of 
the length fioni the tip (range 8.9-9.3%i). This moipho- 
metric data clearly identifies the teeth as those of a mol- 
luscivorous species, as noted bv Nvbakken ( 1990; see also 
Kohn, et al., 1999; Nishi and Kolin, 1999). However, 
these workers did not present details of C. furvus tooth 
moiphometn', wiiicli differs marketlK from previously 
examined molluscivorous Conns species. Nishi and Kohn 
(1999) divided the eleven species thev studied into three 
groups based on both discrete and quantitative charac- 
ters. The former, i.e. presence of one barb, one blade, 
and continuous serration clearly place C. fnrvns in Group 
A, with C. arancosns, C. baiuJanns. and C. mannoreus. 
Groups B and C have teeth with two barbs and no blade 
(Table 2). However, C. fnrvns differs he mi the Group A 
species in its quantitative characters. Its teeth are more 
than twice as long relative to shell length, thus matching 
the iiiniilx IS of Group C rather than Group A (Table 2). 
The (', firms teetli also differ from Group .-X and more 

Espino et ;il., 2008 

Page 147 

Table 2. Compari.son oi'Coiitis fiiixiis radular teeth with those of other moUuscivorous species according to qualitative (discrete) and 
quantitati\e (continuous) characters. N, Number of specimens measured. Qualitative characters: Bl, First barb; B2, Second barb; BL, 
Blade; SE, Serration; +, present; -, absent; C. Continuous; D, Discontinuous. S^, Shell length; T^, Tooth length; BIl, Barb length; 
Bl. Blade length; SEl, Serration length; BAw Base width; SH„ , Shaft width .\\\ dinicnsinns are in mm. Data on Groups A. B, and 
C from \ishi Mid Kolin (1999). 


Quantitative characters 

Group Species 



B2 BL 









C fiimis 



- + 








A C- arancosus 
C. handanus 
C. manuoiviis 




























0.05 0.04 0.06 

(0.03-0.08) (0.03-0.05) (0.05-0.09) 

0.74 0.05 0.03 

(0.62-0.81) (0.04-0.08) (0.(J2-0.05) 

C C. episcopatus 1 1 
r omaiia 9 

(' pcnnaccuK :30 

D 0.07 0.05 0.08 

(0.07-0.08) (0.04-0.05) (0.07-0.09) 

0.68 0.07 0.05 

(0.67-0.69) (0.06-0.08) (0.04-0.05) 

closeK' resemble Groups B and C with respect to all 
four other quantitati\'c characters that the\' share (Ta- 
ble 2). 

PCR .\niplifieali()ii and Plivlogenetic Recon.struc- 
tion: A ph\ logeiietic tree based on t\\'o standard mark- 
ers. 12S and 16S rRXA was constructed as described 
under Methods. Coiuis fiirvits clustered on the same 
branch \\ith species prexiously sho\\ai to ieed on other 
mollusks (Figure 2). In contrast. Conns fu reus was well- 
resohed from C. litteratus, the worm-hunting species 
postulated pre\iousl\" to be C. funus's closest relati\e. 
The h\-pothesis that C. fiiniis is closer to the C.littcratiis 
than to die other mollusk huntins: snails is siffliiticantlv 
rejected (p<0.0001) using Kishino-Hasegawa (Kisliino 
and Hasegawa. 1989) and Shimodaira-Hasegawa (Shimo- 
daira and Hasegawa, 1999) tests comparing the respec- 
ti\e log-hkeliliood scores. The hypodiesis that C. fiirvus 
and C. UUeratus are sister species is signifieantlv rejected 
b\- these tests using the 128 and 168 data alone. Because 
of die great \ariation of forms presently assigned to Co- 
nns fuiTus. we also obtained a "bar code" sequence (COl 
gene) from die specimens analy-zed, which is deposited 
in GenBank. 

Mapping Radular Tooth Characters to the Molecu- 
lar Phylogenetic Tree: \h.iiplionietric d;ita on i;idn- 
lar teeth were available for seven of die nine moUuscivo- 
rous species on the molecular tree (Figure 2), and the 
\alues of the sexen quantitati\'e morphometric characters 
for diese species shouni on the right side of Table 2 were 
entered into a discriminant function anahsis. A phyloge- 
netic tree generated from the pro.ximit}' matrix calculated 
from the centroids of the first two canonical \ariates 
(thev accounted for 99.6% of the variance) (Figure 2B) 

was completelv congruent with the conotoxin gene-based 
tree (Figure 2.\). 

O- and M- Superfaniily Genes fi-oni Cfiimm: Primer 
pairs designed to hybridize to the conserved regions of 
the O- and M-superfamily conotoxins successfully ampli- 
fied conotoxin genes from the C. furvns venom duct. 
Translation of the amplified sequences resulted in amino 
acid sequences clia]-acteristic of the canonical arrange- 
ment of conotoxin precursors. Both oi the sef|uences 
have a hydrophobic N-terminal signal sequence, which 
is highlv consen'ed among conoto.xins belonging to the 
same superfamih', an inten-ening pro region, and the 
mature region at the C-terminus. 

An O-superfamily conotoxin (FrT.l) trom C. fiirviis 
was successfully amplified and sequenced, and a 264- 
base sequence containing an open reading frame coding 
for the was obtained. A 302-base sequence 
coded for a second conotoxin from Cfui'Dus (FrS.l) that 
belongs to the M-superfamilv was also cloned and se- 

The O-snperfamiK' conotoxin trom C. fun us (Fr7.1) 
was stiikinglv similar to an co-conoto.\in isolated from 
Conus textile (Fainzilber et ak. 1996), wTxVII (Figure 
:3A). Alignment of the amino acid sequences shows 
82.1% similarit)'. toTxXTIA has previously been showni to 
target L-ty^De Ca channels in mollusks; given the high 
sequence similarity, FrT.l is likely a voltage-gated Ca 
channel antagonist. A comparison of the M-superiamily 
conoto>dns from C. fuixus (FriB.l) with M-superfamily 
conotoxins from C. textile (Tx3.5) and C. marmorcus 
(Mr3.5) also shows a high degree of similarit)' (Figure 
3.B). Alignment of Fr3.1 with Tx3.5 sliows the serjuences 
to be 84.4% identical while Fr3.1 aligned with Mr3.5 
shows the sequences to be 67.2% identical. The 

Page 148 

THE NAUTILUS, Vol. 122, No, 3 




C. ammiralis 


■ C. dalli 

■C. textile 



EC. a, 
— C. 

C. aulicus 

'. episcopanis 
C. omaria 



C. handanus 


■ C. furvits 

C. ammiralis 

C. textile 

C. episcopatiis 

C. omaria 

C. bandanas 

C. marmoreus 
C. fiirvHS 


■ C. f lavas 


H 50 


• C. geographiis 

C. ohscaras 

C. tulipa 




('. magus 
— C. kintoki 

10 units 


■ C. lilteratas 

■ 05 substitutions/site 

Figure 2. A. Optiiiiiil iiiiLxiiiiuiii likelihood tree from a Bayesian analyses of 12SrRNA and IGSrRNA sequences estimating the 
evolutionary relationship among some Conns species. The uppermost clade, from Conns ammiralis to C. fnrvus, comprises nioUus- 
civorous species. Branch labels are Bavesian support x'alues summarizing confidence in each split. B. Neighbor-joining UPGMA tree 
from analysis of radnlar tooth moiphometn", based on euclidean distance matrix of centroids of canonical variates 1 and '1 determined 
from discriminant hmction anahsis of T radnlar tooth variables, measured as in Nishi and Kohn (1999). 

Fr.3.1 sequence clearly belongs to the in-3 branch of the 
M-conopeptide supertamilv, which is characteristic of 
mcjUuscivorous Conua species (Coipuz ct al.. 20().5). The 
two C. furvus conotoxin sequences are most similar to 
conotoxins from otliei" molluscivorous species, consistent 
with the behavioral obsenations made above and with 
the plu'logenetic tree shown in Figure 2. 


In the central Philippines, Conns jiimis occurs on rocky 
intertidal habitats. At low tide, individuals can be found 
in depressions on the substrate, together with otlier mol- 
lusks; this is where C. furvus is generally collected by 
fishermen in Guimaras Island, to be sold with other com- 
mercial shellfisli at the public market. Conus furvus was 
formerly abundant along the shores of Guimaras Island, 
but an oil spill in the area in August 2006 threatened the 
marine life; C. furvus became much more scarce at this 
locality after the spill, but the populaticni appears to be 

From the results presented above, we coiichidr that 
Conus furvus is a molluscivorous species. This conclu- 
sion is based on direct obsei-vation of C furvus attacking 

gastropod pre\', on nior[ih(inu'tiic characteristics of the 
railular teeth that are shared witli other molluscixorous 
Couus. by the high degree of sequence similarity be- 
tween toxin precursors in the venom duct of Conus fur- 
vus and previously described conotoxins from other mol- 
luscivorous Conus, and bv its phylogenetic proximity to 
them based on molecular markers. A previous molecular 
residt suggesting that C. furvus might be vermivorous 
(Duda et al., 2001; Duda and Kohn. 2005) is probably 

An unusual feature of tlie Conus furvus attack on C. 
niiiiiiliis is diat the prey was apparently injected with 
venom only once; multiple stinging was never obsen-ed. 
All other molluscivorous Conus species previously ob- 
sei'ved attack prey using inulliple injections of venom 
before starting to feed on the iiinnobilized prev (Yoshiba, 
19S3, 1987; Kohn, 200.3). 'flic leason for this 'behavioral 
difference is imclear, but the \enom composition could 
well reflect it. A fmther biochemical characterization of 
Conus furvus venom is in progress (S. Espino and G. 
Concepcion, unpublished results). 

The phylogenetic relationship bet\yeen C. fuivus nud 
other Conns species evaluated for 12S and 16S rRNA 
gene sequences (Figure 2) is consistent with C furvus 

Espino et al., 200S 

Paee 149 


Mr3 5 


Mature toxins 








Figure 3. A. Aliijnment ot the toxin resjion ot FrT.l from 
Conns fimiis and coTxMI from Conns textile (GenBank Nuc. 
Ace. No. DD012T70). B. Alignment of FrS.l from Conns ftir- 
vns with otlier M-superfamilv precursors highlighting the con- 
seived amino acids. Tx3.5 and Mr3..5 were from Conns textile 
and Conns mamiorens, respectively (Coipuz et al., 2005). The 
predicted matin'e toxins are eJso shown; the asterisk denotes an 
amidated C-terminus. 

being more closeh' related to the mollusk hunting Conns 
species tlian to C. litteratus (Bayesian clade support = 
98), a \'ermivorous species. The several clades of mollus- 
civorous Conns probably have a common ancestor. Using 
the tree based on molecular data in Figure 2A, the well 
supported groups can be referred to, using the subgenera 
of Marsh (1964) as clade names) as the Conns clade (with 
Conns mannoreiis as the type), the CijUnder clade, (witli 
Conus textile as the t\'pe), and the Dariocomis clade 
(with Conus onmria as the U'pe). Although Conns furous 
belongs within the larger mollusk hunting clade, includ- 
ing tliese three groups, its position within the clade re- 
mains unresolved. Gi\'en its lack of a tented pattern, it 
has an unusual shell pattern for a molluscivore. However, 
some specimens of smaller, usuall)' tented species such 
as Conns barbieri and Conus victoriae occasionally have 
plain brown shell \'aiiants, not unlike t\|:)ical specimens 
of Conus furvus. 

A more comprehensive molecular analysis of moUus- 
civorous Conus, including C. furvus. has recently been 
carried out; these results will be presented elsewhere. 
Additional phvlogenetic markers, such as ITS-2 se- 
quences, were used for this study. This has led to greater 
refinement of die phyiogenetic tree and supports the 
basic conclusion that C. furvus is a molluscivore. 


Tills work was supported in part by GM48677 from the 
U.S. National Institutes of General Medical Sei"vices (to 
BMO). The support of the University of the Philippines 
in the \isayas for a fellow'ship award to S.E. to pursue 
graduate studies in Chemistrs' is gratefully acknowl- 
edged. XSF Grant 0316338 partly supported the re- 
search of AJK. The late J.E. Norton contributed the C. 
fuixus specimens for radular tooth moi-phometry, and 
Dr. M. Nislii assisted in the data analysis. 


Coipuz, G. P., R. B. (acobsen, E. C. Jimenez, M. Watkins, C. 
Walker, C., Colledge, et al. 2005. Definition of the M- 
conotoxin superfamily: characterization of novel peptides 
froin molluscivorous Conus venoms. Biochemistiy 44(22): 

Duda, T. F., Jr. A. J. Kohn, and S. R. Palumbi. 2001. Origins of 
diverse feeding ecologies within Conns, a genus of ven- 
omous marine gastropods. Biological Journal of the Lin- 
nean Society 73" 391-409. 

Duda, T.F., Jr. and A. J. Kohn. 2005. Species-level phylogenog- 
raphy and evolutionaiy history of the h)perdiverse marine 
gastropod genus Conns. Molecular Phylogenetics and 
Evolution 34: 257-272. 

Duda, T. F., Jr. and S. R. Palumbi. 2004. Gene e.xpression and 
feeding ecolog)': evolution of piscivoiv in the venomous 
gastropod genus Conus. Proceednigs: Biological Scieneces 

Fainzilber, M., J. C. Lodder, R. C. van der Schors, K. W. Li. Z. 
Yu, A. L. Burlingame, et al 1996. A novel hydrophobic 
omega-conotoxin blocks molluscan dihydropyridine- 
sensitive calcium channels. Biochemistn' 35(26): 8748- 

Huelsenbeck, J. P. and F. Ronquist. 2001. MRBAYES: Bayes- 
ian inference of phyiogenetic trees. Bioinformatics 17: 

Jeanmougin, F., J. D. Thompson, M. Gouy, D. G. Higgins, and 
T. J. Gibson. 1998. Multiple sequence alignment with 
Clustal X. Trends in Biochemical Sciences 23: 403^05. 

Kishino, H. and M. Hasegawa. 1989. Evaluation of die maxi- 
mum Hkehhood estimate of the evolutionaiy tree topolo- 
gies from DNA sequence data, and the branching order in 
homiiioidea. Journal of Molecular Evolution 29: 170-179. 

Kohn, A. J. 1959. The Ecolog)' of Conns in Hawaii. Ecology' 
Monograph 29: 47-90. 

Kohn, A. J. 2003. The feeding process in Coinis victoriae. In; 
Wells. F. E., D. I. Walker, and D. S. Jones (Eds.) The 
Marine Flora and Fauna of Dampier, Western Australia, 
Volume 1. Western Australian Museum, Perth, pp. 101- 

Kohn, A.J., M. Nishi, and B. Fernet. 1999. Snail spears and 
scimitars; A character analysis of Conns radular teeth. 
Journal of Molluscan Studies 68; 461-t81. 

Kohn, A. J. and J. W. Nybakken. 1975. Ecolog)-- of Conns on 
eastern Indian Ocean fringing reefs; diversity of species 
and resource utilization. Marine Biolog)' 29: 211-234. 

Maddison, D. R. and W. P. Maddison. 200L MaClade version 
4.08. Sinauer Associates; Sunderland, Massachusetts. 

Nishi, M. and A. J. Kohn. 1999. Radular teeth of Indo-Pacific 
molluscivorous species of Conns: A comparative analysis. 
Journal of Molluscan Studies 68: 483^97. 

Nybakken, J. 1990. Ontogenetic change in the Conns radula, its 
form, distribution among die radula t)pes, and signifi- 
cance in systematics and ecology. Malacologia 32; 35-53. 

Olivera, B. M. 2002. Conns venom peptides: reflections from 
the biolog)' of clades and species. Annual Review of Ecol- 
ogy and Systematics 33: 25-47. 

Olivera, B. M. 2006. Conns peptides: biodiversity-based dis- 
coveiy and exogenoniics. Journal of Biological Chemistn' 
281(42); 31173-31177. 

Rockel, D., W. Korn, and A. J. Kohn. 1995. Manual of the 
Living Conidae. Vol. 1, Indo-Pacific Region. Verlag 
Christa Henimen, Wiesbaden, 517 pp. 

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of Log-Likelihood with Apphcations of Phvlogenetic Tillet, D. and B. Neilan. 2000. Xanthogenate Nucleic Acid Iso- 

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cal Reviews 84: 41-68. (Abstract) [in Japanese] 

THE NAUTILUS 122(3):151-154, 2008 

Page 151 

Two new species oi Mitrella (Gastropoda: Neogastropoda: 
Columbellidae) from the lower Miocene Chipola Formation of 
northwestern Florida 

Richard Duen- 

Post Office Box 1055 
Okeechobee. FL 34973 USA 


Two new species oi Mitrella from the lower Vliocene Chipola 
Foniiation of are described. Mitrella haijcsonim new species 
and Mitrella plu/llisae new species are found primarily in 
Chipola Formation exposures along Farley Creek in northwest- 
ern Florida. USA. Placement of the new species in die genus 
Mitrella is tentati\e. 

Additional Keywords: Neogene, ultraviolet light, gastropod. 


Species assigned to the genus Mitrella are wadely distrib- 
uted in tlie warm and shallow waters of the Recent seas 
of tlie world (Gardner, 1947. 1948). The exact origin of 
Mitrella is not clear, but some of the earliest represen- 
tatix'e of tlie genus from the southeastern portion of 
North America appeared in the Eocene (MacNeil and 
Dockeiy III, 1984; Palmer, 1937). Maury- (1910) de- 
scribed diree Mitrella (as Astijris) from the Chipola For- 
marion. These were originally deposited in the Cornell 
Uni\ersit\- collection at Ithaca, New York, and now de- 
posited at PRI. With more specimens than Maury had at 
her disposal, Gardner (1947) described 13 species and 
t\vo subspecies of Mitrella from the Chipola Formation. 

Mitrella have been found in all exposed facies of the 
Chipola Formation on die Chipola River, Tenmile Creek 
and Farlev Creek, in tlie Chipola River drainage in Cal- 
houn Countw Florida. Thev are also present in the 
Chipola Formation e.x|iosed in the lower bed at Alum 
Bluff in Libertv Count\-, Florida. The two new species 
described herein have been collected only in the Chipola 
Formation exposures in the Chipola Ri\'er drainage, pri- 
niarilv at Farlev Creek. \'okes (1989) stated the Chipola 
facies along Farley Creek \\4iere the two new species are 
most prevalent is a bi\alve-ricli niihohd Hme-sand with 
manv calcareous algae and coral heads, and assumed it 
was a shallow back-reef environment. 

Some specimens were photographed under ultraviolet 
Hght (Ur\") to facihtate visuahzation of color patterns. In- 

stitutional aL)bre\iatiuns are: UF: Florida Museum of 
Natural Histoiy (FLMNH for locality records), Univer- 
sity of Florida, Gainesville; PRI: Paleontological Re- 
search Institution, Ithaca; ANSP: The Academy of Natu- 
ral Sciences, Philadelphia; AMNH: American Museum 
of Natural Histon', New York. 


Superfamily Buccinoidea Raiinesque, 1815 
Family Columbellidae Swainson, 1940 
Genus Mitrella Risso, 1826 

Diagnosis: Small to veiy small, smooth, fusiform 
shells. Sculpture, if present, of incised spiral hues. Spiral 
sculpture is generally restricted to tlie anterior end near 
the base: axial sculpture is, with few exceptions, entirely 
absent. The outer lip is dentate in adult specimens (Di- 
agnosis according to Campbell [1993] and Keen [1971]). 

? Mitrella hayesomm new species 
(Figures 1-3, 10) 

Description: Shell fusiform, narrow. Height ut liolo- 
type 19.8mm. Protoconch mammillated, with about t\vo 
smooth, rounded whorls, the second expanded, with no 
perceptible protoconch/teleoconch transition. Teleo- 
conch with ten slightlv convex, shiny whorls with micro- 
scopic irregular spiral grooves, a.xial sculpture lacking. 
Spire elevated with somewhat concave sides. Suture dis- 
tinct. Aperture elongated, less than half the length of the 
entire shell. Outer lip sinuous, varicose externally, mar- 
gin thin and sharp. Basal lip extending slightly beyond 
pillar. Posterior canal produced and thickened externally. 
Denticles present on inner surface of outer lip, ten to 12 
in number, weak on anterior third, strongest on central 
third, frequently absent on posterior third. Parietal wall 
with two axial lidges, outer lidge weak with small raised 
beads reflecting underlying sculptiu'e on pillar. Inner 
ridge prominent with median notch, adapical half of 
ridge strong, abapical half tapering and completelv chs- 
appearing abapically. Base of pillar with about nine 

Pag-e 152 

THE NAUTILUS, \'ol. 122, No. 3 

R. Duerr, 200S 

Paee 153 

oblique groo\'es with a idiiikK'iI siiiiiinit lu'tween 
groin es. Dark band about one third width ol pcnnltiniate 
whtirl appears below suture on last threi' whorls ol ho- 
lot\pe. Band has irregular narrow, obli(|ne, lighl colored 
lines, some forming u- or \-ligni'es. 

Type Material: llolotxpe: UF 119655, height 
IQ.Sinm, width 5.6nun: Paratvpes: ANSP-IP 81324, one 
specimen; AMNH-Fl 43312,' one specimen; PRI 8382, 
one specimen; Diegel-Duerr collection, one specimen. 
All from t^pe localit\'. , •-.,,. 

T>pe Locality: FLMNH locality Farlex' Creek 07 
(CA022), FarleV Creek east of SR 275, Calhoun County, 
Florida (to protect privacy rights of landowners, specific 
localit\- information is available onlv to qualified re- 
searchers upon written request to the author or the In- 
vertebrate Paleoutolog\' Dixision of the FLMNH), 
Chipola Formation. 

Distribution: Chipola Formation along Teumile and 
Farle\- Creeks and the Chipola River, Calhoun County, 


Etymology': Named to honor the forestn'-oriented 
Ha\es faniih- who ha\-e graciously granted the author, 
and others, permission to collect on tluir piDpeit)'. 

Remarks: The species currently placed in MitrcUa 
comprise a complex group and may have been diiterently 
assigned to die genera A?;rt, Asti/ri.s, Nitidelh by different 
authors (Keen, 1971). The taxonomie position of M. 
haijcsonim is questionable and its current assignment to 
Mitrella is tentative. Mitrella hai/csoruin has unique 
characters, such as the bulbous second whorl of the pro- 
toconch, extended slightly concave spire and wide sipho- 
nal canal extending below the pillai", and cannot be con- 
fused with any other knowii Mitrella. Althougli no speci- 
mens oi Mitrella dalli (Maun-, 1910), the species in the 
Chipola Formation closest in form to M. hai/csonim, 
were a\ai]able for studv, Maun's figure ot M chilli (1910: 
pi. 6, fig. 2) indicates a smaller shell (12 mm), a shorter, 
stouter spire, and a narrower anterior canal than M. 
hai/esomm. The nearest European fossil congener of Mi- 
trella haijesorum is MitrcUa (MacnireUa) nasspicles 
(Grateloup, 1827) (Figures 4—6) from the earlv Pliocene 
of Italw which is larger, wider at the midbody, and has a 
narrow anterior canal. The maximum height of all speci- 
mens of Mitrella haijesonim examined is 19.93 mm, 
minimum height is 18.17 mm. Exposure of M. haije- 
sonim to ultra\iolet light (Figure 10) reveals a fluores- 
cent pattern of axial flammules in addition to the narrow 
band below the suture on the anterior whorls of the new 

species. Seven spt'c iiiu'us ol the most common Mitrella 
in the Chipola Formation, Mitrella iselvia Gardner, 
1947, a more robust species than M. Iiai/e.soniin, were 
examined under UV light and revealed fine filamentous 
lines covering all teleoconch whorls. Three specimens of 
Mitrella aseina Gardner, 1947, a shorter species than M. 
haijesorum. exhibits, under UV light, narrow axial bars 
nmning from suture to suture, offset in alignment from 
prexious whorls, willi lour per w horl on most teleoconch 

r' Mitrella plu/lli.sae new species 
(Figures 7-9, 11) 

Description: Shell small, hisilornr Height 7 mm, 
width 2.2 irmi. Protoconch with two smooth whorls, sec- 
ond whorl enlarged. Teleoconch witli five smooth, con- 
vex whorls, without axial stulpture. Spire sides shghtly 
concave. Suture impressed. Apeiiure less than half the 
length of entire shell. Outer lip with slight vaixx, margin 
thin and shaq:), usually dentate witliin. Thin parietal wash 
present. About seven impressed olilicjue grooves cross 
base of pillar, separated by narrow liands with rounded 
summits. Faint round spots, eight on last whorl of holo- 
type, appear on last three whorls. Spots about one- 
quarter the height of penultimate whoil in diameter rest 
just above median line of the whf)rls. 

Tyjie Material: Holot)q3e, UF 119656, height 7.(.) nun, 
width 2.2 nun; ParatyjDes, ANSP-IP SI325, one speci- 
men; AMNH-FI 43313, one specimen; PRI 8383, one 
specimen; Diegel-Duerr collection, one specimen. All 
from t)rpe locality. 

Type Locality: FLMNH locality Farley Creek 07 
(CA()22), Fai-lev- Creek east of SR 275, Calhoun Count)', 
Florida, (to protect privacy rights of landowners, specific 
locality information is a\ailable only to qualified re- 
searchers upon written reijnest to the author or the In- 
vertebrate Paleontolog\' Dixision of the FLMNH), 
Chipola Formation. 

Distribution: Chipola Foinuilion along Tenrnile and 
Farley Creeks and the Chipohi Hixer, Calhoun Count)', 

Etymology: Named for Plnllis Diegel, the author's 
companion and a knowledgeable conchologist and pale- 

Remarks: As witli Mitrella hai/esonim, the assignment 
of i\/. plnjllisae to Mitrella is tentatix'e. A cursoiy inspec- 
tion would indicate Mitrella phijllisae to be a dwarf M. 
haijesonim. Closer examin;ition reveals that M. phijllisae 

Figures 1-11. — Mitrella species. 1-3. Apertural, lateral, and abapertural viexvs of the holotyjDe oi' Mitrella luiijesoniin new species, 
UF 1196.5.5, height 19.8 mm, width 5.6 mm, Burdigalian Miocene. 4-6. Apertural, lateral, and abapertural xiexx's of Mitrella nassoides 
(Grateloup. 1827) UF 119657, height 25.5 mm, xxidth 8.6 mm. Zaiiclean Pliocene, from Liguria Comity, Ceriale, Italy, for comparison 
xxitli M. haijesonim. 7-9. Apertural. lateral, and abapertural xiews of the holotspe oi Milnllu yilujlhsae new species, UF 119656, 
height 7.0 mm. width 2.2rnm, Burdigalian Miocene. 10, 11. Mitrella exposed to U\' light. 10. Ilolotxpe oi' Mitrella haijesonim (same 
specimen as Figure .3) showing U\' exposed pattern. 11. Holotxpe of Mitrella phi/lhsae (same specimen as Figure 9) shoxving UV 
exposed pattern. 

Page 154 

THE NAUTILUS, Vol. 122, No. 3 

differs from the much larger M. hayesorum by the 
shorter spire, the shghtly more impressed suture, and 
fewer oblique incised lines on the base of the shell. A 
series of dots about 1 mm in diameter are visible on M. 
phijUisac encircling all teleoconch whorls, one-third the 
length of the whorl below the suture, eight on the last 
whorl. E.\-posure to UV light reveals (Figure 11), in ad- 
dition to the larger dots, a field of minute dots covering 
the entire teleoconch, somewhat reminiscent of the pat- 
tern on the Recent Mitrclln ocellata (Gmehn, 1791) from 
the western Atlantic. Also, the anterior portion of the 
outer lip of M. plujUisac is less developed than that of A'/. 
hayesontm, wliich is wider and extends bevontl the pillar. 
The height of all specimens of M. phijllisae examined 
varies less than 1.0 mm, from a maximum of 7.4 mm to 
a minimum of 6.S mm. 


The author extends his appn-ciation to Roger Poitell, 
FLMNH, for pnniding the plates and connnenting on an 
earlier draft of the manuscript. Digital images were pre- 
pared by Sean Roberts, FLMNH. Thanks also to Marc 
Grigis, Belgium, for donation of a specimen of MitreUa 
nassoicles. to Dr. Bernartl Landau, Portugal, for encour- 
ageinent and advice, and to Burke and Brooks Hayes 
and also Archie and Vicki Whittington for permission to 
collect on their respective properties in North Florida. 
Pamela McBride provided secretarial assistance. I am in- 

debted to two anonymous reviewers whose comments 
and suggestions improved the manuscript. 


Campbell, L. D. 1993. Pliocene molliKsks from die York-town 
and Cliowen River Formation in Virginia. Virginia Divi- 
sion of Mineral Resources, Pulilication 127: vii + 173 pp., 
43 pis.. 

Gardner, J. 1947. The molluscan fauna of the Alum Bluff 
Group of Florida. Fait VIII. Ctenobranchia (remainder), 
Aspidobranchia and Scaphopoda. United States Geologi- 
cal Survey. Professional Paper 142-H: i-ii + 493-638, pis. 

Gardner, J. 194S. Mollusca from the Miocene and Lower 
Pliocene of Virginia and North Carolina. Part 2. 
Scaphopoda and Gastropoda. United States Geological 
Sun-ey. Professional Paper 199-B: i-iii +179-279, pis. 24- 

Keen, A. M. 1971. Sea Shells of TropiciJ West America. Marine 
mollusks from Baja California to Peru. Second Edition. 
Stanford Universitv Press, Stanford, California: .xiv + 1064 
pp., 22 pis. 

Mac Neil. F. S. and D. T. Docken- III. 19S4. Lower Oligocene 
Gastropoda, Scaphopoda, and C^ephalopoda of the Vicks- 
Imru; Group in Mississippi. Mississippi Bureau of Geology. 
Bulletin 124: 41.5 pp., 72 pis. 

Maun, C. [. 1910. New Oligocene (Miocene) shells from 
Florida. Bulletins of American Paleontology 4(21): 119- 
164, pis. 18-26. 

Vokes, E. H. 1989. An oveniew ol the Chipola Fonnation, 
northwestern Florida. Tulane Studies in Geolog)' and Pa- 
leontolotrv 22: 13-24. 

THE NAUTILUS 122(3);155-165, 2008 

Page 155 

Catalogue of the t\pe material of molliisks deposited at the 
Zoology Museum, University of Costa Rica 

Fresia Villalobos-Rojas 
.\na. G. Guzman-Mora 

Escuela de Biologi'a 
Universidad de Costa Rica 
Apdo. 2060, S<\n Pedio 
San lose, COSTA RICA 

Yolanda E. Cainacho-Garcia' 

Esfuela de Biologia and Museo de Zoolntji'a 
Uni\ersidad de Costa Rica 
Apdo. 2060, San Pedro 


This catdogiie compiles the basic data for mollnsk tvpe mate- 
rial deposited at the Zoolog\- Museum of the University of 
Costa Rica. It includes 62 holohpes and 151 paratypes, repre- 
senting 6S species and 41 genera. The species, authors, date of 
publication, bibliographic references, type locality, catalogue 
number, number of specimens, state ol presen'ation, and re- 
lated infonnation are indicated for each taxon. In some cases, 
remarks on die current taxonomic status and other useful in- 
formation are provided. For all species, except for opistlio- 
branchs. the shell of either the liolot\-pe or one of the paratypes 
is illustrated. 

Additional Keijwords: Holotvpe, parat)pe, MoUusca, molKisk 


The Zoology Museum of the University of Costa Rica 
(MZUCR) was founded in 1960 by UCR professors 
Douglas C. Robinson and Wilhain Bussing. The collec- 
tion consists of both invertebrates and vertebrates, and 
includes birds, fishes, mammals, reptiles, amphibians, tu- 
nicates, decapods, sipunculids, sponges, corals, and mol- 
lusks, among other groups. The specimens come from a 
wide \'ariety of research projects, and donations from 
national and international researchers, as well as amateur 
naturalists. Most of the material is preserved in a wet 
collection (70% ethanol), but there is also a dry collec- 

The Mollusk Collection at the UNivERsm' of 


Former UCR curator Carlos \'iIIalobos and a great num- 
ber of other researchers started the mollusk collection at 
the Museum in 1964. Since then, UCR staff and visiting 
scientists have continuously added to the collection. In 
addition, a significant amount of material has been do- 

^ Author for correspondence 

nated to the Museum. Some examples include the col- 
lections of malacologists George Richard, from Univer- 
sity of La Rochelle, France (mainly specimens of the 
family Conidae and other material from the Indo- 
Pacific), Kristie L. Kaiser, associated with the Santa Bar- 
bara Museum of Natural Histor)' (material from Cocos 
Island, Costa Rica), and tlie Latin American collection of 
independent scientist Dwight W. Taylor (including 
terrestrial and freshwater mollusks as well as type 
material); and amateur naturalists Michael Montoya (a 
valuable collection of marine mollusks from the Me- 
soamerican region, Cocos Island, and the Caribbean), 
Mary Yost (marine mollusks from Guanacaste, Costa 
Rica), Robert Nishimoto and others in 1969 (marine 
mollusks from Puntarenas, Costa Rica), and Jerry Well- 
ington in 1972 (marine mollusks from both the Pacific 
and the Atlantic). In 2005, the collection increased in size 
considerablv, having absorbed the mollusk collection of 
the Costa Rican National Biodiversity Institute (INBio), 
which closed its Malacology Department in September 
2004. Currently, the collection is comprehensive, in- 
cluding more than 37,670 lots and 216,587 specimens 
representing the inolluscan fauna of the Paciiic Coast 
of North, Centra], and South America (however, there 
is also material from the Indo-Pacific, France, and 

The collection is divided into two main categories: a 
dry collection consisting mainly of tlie shells oi bivalves 
and gastropods, and a wet collection comprising cepha- 
lopods, gastropods, bivalves, polvplacophorans, and ter- 
restrial and Ireshwater mollusks preserved in alcohol. 

Considering that this collection is the most important 
in the country and perhaps in all of Central America, and 
due to the recent increase in the amount of type mate- 
rial, it is important to compile this information and pub- 
lish a catalogue that addresses in detail the literature 
available for the type material, its location, and illustra- 
tions of the shell of each type represented. We hope to 
facilitate the future work of taxonomists and help locate 
type material, contributing in this respect to their re- 

Page 156 

THE NAUTILUS, Vol. 122, No. 3 

F. N'ilkilobos-Rojas et al., 2008 

Page 157 

Type Collection 

This catidogiie iiicludes 62 luilotxprs aiul 151 parahpt^s, 
representing 68 species and 41 genera. Ail hpes are 
listed alphabeticalK b\- species epitliet. Photographs of 
opisthobranch tvpes are not included, although they are 
listed in this catiilogue. For each t;L\on, the following 
information is included: the name ol the species cited 
exactly as it was published in the original description, the 
author(s), date oi publication, bibliographic references, 
tspe locality catalogue number, nmnber of specimens, 
state of presenation, and related information (SEM 
stubs, microscopx slides, egg masses of the specimens). 
In some cases, remarks on the ciu'rent taxonomic status 
and other useful information are provided. We also in- 
dicate whether the specimen is presei-ved complete (in- 
cluding the shell and soft parts) or \\licn onl\- th(^ slicll 
rem;iins in the dn' collection. 

In a few cases, the information in the original publi- 
cation about the location of the t\pe material is not ac- 
curate, and we clarify tliis in the remarks. For each spe- 
cies, either the holotspe or one of the paratvpes is pho- 
tographed, and the figure number is properly indicated 
beside the specimen. 

.Although the original publications on some t\pe ma- 
terial (Noumea regalis Ortea, Cabailer and Moro, 2001; 
Dcntimargo an^ouauta Espinosa and Ortea, 2002; Tico- 
cijstiscus ibcria Espinosa and Ortea, 2002; Cratena piii- 
taensis Ortea, Cabailer, and Espinosa, 200.3; Millcria 
rifmica Ortea, Cabailer, and Espinosa, 2003; Phidiana 
adiuncta Ortea, Cabailer, and Moro, 2004; and Costoa- 
nachis cascahulJoi Espinosa and Ortea, 2004) stated tliat 
the material was deposited at INBio, in fact, tlie material 
\\as ne\er deposited there or at the University of Costa 
Rica. i\lthough attempts have been made to clarify the 
location of this material with the authors, none has been 
successful. Abbreviation used in the text: m.a.s.l. = 
meters abo\^e sea level. 


academica, Okenia. Camacho-Garcia and Gosliner, 
2004: 431-138, figs. 1-3. T\pe locality- Playa Tamarindo, 
Tempisque Consen-ation Area, Puntarenas, Costa Rica 
(9°03'58" N, 85°51'08" W), m depth. HOLOTiTE: 
MZUCR-INB000311S102 (specimen), MZUCR- 
1XB00037649S8 (radula and jaw, SEM stubs), 
PARATi'PE; MZUCR-IXB0001496648. San Miguel, 
Reser\'a Natural Absoluta de (Jabo Blanco, Tempisque 
Conservation Area, Puntarenas, Costa Rica (9°.34'5.3" N, 
85°08'26" W), m depth. 

aeci, Philinopsis , Ortea and Espinosa, 2001b: 41, pi. 
2C, ■3A. T\pe localitx"; Punta Mona, Manzanillo, Linion, 

Costa Rica (9°6'37" N, 82°66'08" W), 6 m depth. 
MZUCR-INB0()03721981. Same locality data as holo- 
t)pe. Remarks: Paratope said to be deposited at the In- 
stituto de Oceanologia, Habana, Cuba, but was deposited 
at INBio. 

alfiopivai, Plcsiocijstiscus. Espinosa and Ortea, 2002: 
102-106, figs. 1-3, pl.l. Ty^je locality: Manzanillo, 
Union, Costa Rica (9°39" N, 82°,39" W), 20-24 m depth. 
HOLOTYPE: MZUCR-INB0003754713 (shell. Fig- 
ure 21). 

anulatuH, janoJus. Camacho-Garcia and Gosliner, 
2006: 129.5-1305, figs. 1-7. Type localit)': Isla Ballena, 
Parque Nacional Marino Ballena, Puntarenas, Costa Rica 
(9°06'24" N, 8.3"43'35" W), 6 m depth. HOLOTYPE: 
MZUCR-INB0001495772 (specimen), MZUCR- 
INB()0()3764958 (SEM stub with jaw), MZUCR- 
INB0003764987 (SEM stub with radula), MZUCR- 
INB0003765()66 (slide preparation with labial plate). 
PARATYPES: MZUCR-INB0003764915 (1 speci- 
men). Same locality data as holotype; MZUCR- 
INB0003836171 (1 specimen), SW side of Isla Plata, 
Guanacaste, Costa Rica (10"26'4S" N, 85°48'20" W), 10 
m depth. 

arleyi, MdaneUa, Espinosa, Ortea and Magana, 2001: 
123-124, fig. 3. T\pe localit^': Punta Mona, Manzanillo, 
Limon, Co,sUi Rica (9°37" N,'82°37" W), 10-12 m depth. 
HOLOTYPE: MZUCR-INB0003138455, (shell. Fig- 
ure 1). 

awapa, Doto, Ortea, 2001a: 21-23, pi. 2G, fig. 9. T>pe 
locality: Punta Mona, Manzanillo, Limon, Costa Rica 
(9°37'' N, 82°37" W), 10 m depth. HOLOTYPE: 
MZUCR-INB0001497507 (specimen and egg mass). 

beatrix riopejensis. Helicina {"Gemma"), Richling, 
2004: 30.3-308, figs. 13.3-138. T\pe localit\-: SW of Liv- 
eiqDOol (about 24 km of Puerto Limon) along Rio Peje, 
Limon, Costa Rica (90°55'46" N, 83°13'1.5" W), 1135 
m.a.s.l. HOLOTYPE: MZUCR-1NB00.3542625 (shell. 
Figure 30). PARATYPE: MZUCR-INB0003542626 
(shell). Sann- locality data as holotspe. 

boeckereli, Alcadia (Microalcadia), Richling, 2001: 
6-7, figs. 9-12. Tvpe locality: Guanacaste National Park, 
about 10 km S of Santa Cecilia, Volcan Orosi, near 
field station Pitilla, beginning of Sendero Orosilito, 
primaiy forest, Guanacaste, Costa Rica (10°59'18" N, 
85°25'.34" W), 700 m.a.s.l., HOLOTYPE: MZUCR- 
INB00034049S0 (shell. Figure 24). Remarks: species was 
assigned to Alcadia in Richling. 2004; 374—377, figs. 257- 
262; pre\iouslv in Helicina. 

bramale, Hoplodoiis, Faliey and Gosliner, 2003; 198- 
201, figs. 17E, 28-30. Type localit>-: Puerto Escondido, 
Manuel Antonio National Park, Puntarenas, Costa 
Rica (9°23' N, 84°0S' W), m depth. HOLOTYPE: 

Figures 1-12. 1. Holot\pe of McUancla cirh'i/i. 2.1 iiiiii. 2. Holot\pe oi MelaneUa zugnigac. 1,1 iiini. 3. Holot\pe ot Tiiphora 
orteai. 2.0 mm. 4. Parat\pe oi Chicoreux (Plu/Unnotiis ) eversonii. L37 mm. 5. Parat\pe oi Tijphisopsis carobkogluiulae. length 19 mm; 
diameter. 11mm. 6. Parat\pe oi Mitrella loisae, 4.9 mm. 7. Holotspe oi Hi/aliiui rhicoi. 9.1 mm. 8. Holot\pe of Pniniim caluiitdcn.sis. 
12.5 mm. 9. Holohpe oi Pninum chumi, 18.2 mm. 10. Holotype of Pniiuiin liohnuiiic. 19. .5 mm. 11. HolotApe ot Pniniim lizaiioi. 
13.2 mm. 12. Holohpe oi Vokaiinn socoac, 11.8 mm. 

Page 158 

THE NAUTILUS, Vol. 122, No, 3 


INB0003572306 (6 .specimens), Punta Uvita, Marino 
Ballena National Park, Puntarenas, Costa Rica (9°08' N, 
83°45' W), 0-2 m depth; MZUCR-INB0001498.550 (1 
specimen), m depth. Same localitv data as the other 

bribri, Gibbenila, Espinosa and Ortea, 200U: 10(1- 
101, pi 1, fig. 4. Type Locality: Punta Mona, Manzanillo, 
Limon, Costa Rica (9°38' N, 82°37' W), 10-15 m depth. 
HOLOT^'PE: MZUCR-INB0003;349908 (shell. Figure 
16). PARATYPES; MZUCR-INB()()().3:349905, MZUCR- 
INB0003349906, MZUCR-INB0003349907 (1 specimen 
each, shells). Locality data for all paratypes same as ho- 

caballeri, Philinc. Ortea, Espinosa and Moro, 2001: 
38-40, figs. 9-10, pi. 2B. T>'pe locality: Punta Mona, 
Manzanillo, Limon, Costa Rica (9°37' N^ 82°37' W), 9 m 
depth. HOLOTYTE: MZUCR-INB00()3138445. 

cabecar, Doto, Ortea, 2001a: 34-37, figs. 16-17, pi. 
2M. Type Locality: Punta Mona, Manzanillo, Limon, 
Costa' Rica (9°.39' N, 82°37' W), 20 m depth. 
HOLOTYPE: MZUCR-INB000.3449742. 

cahiiitaensis^ Pniiiuin, Magaiia, Espinosa and Ortea, 
2003: 122-124, figs. 1, 2A-B, pis. lA, 2A. Type Locality: 
1 km E from Puerto Vargas Station, Cahuita National 
Park, Limon, Costa Rica"(9°43'33" N, 82°48'31" W). 
HOLOTiTE: MZUCR-INB0003718203 (shell. Figure 
8). PARAT\TE: MZUCR-INB0003718204 (shell). Same 
localit\' data as holotvpe. 

caribetica, Gibbenila, Espinosa and Ortea, 2002: 
113-114, fig. 10, pl.l. Type locality: Punta Mona, Man- 
zanillo, Limon, Costa Rica (9°37' N, S2°37' W), 8 m 
depth. HOLOTYPE: MZUCR-INB0003718205 (shell. 
Figure 17). 

carolskoglundae, Typhisopsis, Houart and Hertz, 
2006: 56-58, figs. 17-25, 47-49, 59, 63. Type localit\': 
Playas del Coco, Guanacaste, Costa Rica (10°55'53" N, 
85°69'81" W), 24-37 m depth, on mud bottom. 
PARATYPE: MZUCR-6153, (shell. Figure 5). Boca de la 
Honda, Veraguas, Panama (7^27' N, 80°51' W), in white 
sand. Remarks: The coordinates and the collecting local- 
ity of this paratype appear to be incorrect since these 
coordinates plot inland. 

chicoi, Ht/alina, Espinosa and Ortea, 1999a: 167-169, 
figs. lA-C, 2A, 3A-D. Type locality: Manzanillo, Lim<')n, 
Costa Rica (9°38' N, 82°39' W), 5-12 m depth. 
HOLOTYPE: MZUCR-INB0003350839 (shell. Figure 
7). Remarks: Paratype said to be deposited at INBio in 
the original publication, but never sent to INBio or the 
University of Costa Rica. 

chiquitica, Oligyra, Richling, 2001: 1-2, figs. 1- 2. 
Type Locality: 9 km W of Matina, a little stream up the 
Rio Barbilla from the crossing of the road Siquirres to 
Limc'jn, along a tributaiy of Rio Barbilla, Limon, Costa 
Rica (10°03'29" N, 83°22'24" W), 70 m.a.s.l. 
HOLOTYPE: MZUCR-INB0003404977 (soft parts 
and shell. Figure 32). PARATYPE: MZUCR- 
INB000.3404981 (.shell and soft parts). 

chiimi, Pniniini. Espinosa and Ortea, 2000: 107-108, 
figs. 8-9. T)pe locality: Manzanillo, Limon, Costa Rica 
(9°38' N, 82°39' W), 10-15 m depth. HOLOTYPE: 
MZUCR-INB0003349912 (shell. Figure 9). 
INB0003349914, MZUCR-INB0003349915 (1 specimen 
each, shells). Locality data for all paratypes same as ho- 

convenientis, Eubranchus. Caballer and Ortea, 2002: 
81-85, figs. 2, 3, 7 pi. IB. Type locality: Manzanillo, 
Limon, Costa Rica (9°38' N, 82°39' W), m depth. 
HOLOTYPE: MZUCR-INB00():35768.32. 

corcovadensis, Cnjptostrakon, Cuezzo, 1997: 1-8, 
figs. 1-14. Type locality: Corcovado National Park, Si- 
rena Station, Sendero a Rio Los Patos, Puntarenas, Costa 
Rica (8°30' N, 83°.35' W), 10 m.a.s.l. HOLOTYPE: 
MZUCR-INB0()01468087 (shell. Figure 37; SEM stub 
with radula). PARATYPE: MZUCR-INB0001468080 
(shell and soft parts). Remarks: Paratype locaHty data 
same as holotyjje. In the publication, the holotype and 
paratype catalogue numbers mistakenly written 
INB00046S087 and INB000468080, respectively. 

costaciibensis, Janoliis, Ortea and Espinosa, 2000: 
80-83, figs, 1-2. Type localit^': Miramar, N coast of La 
Habana,'Cuba (2:3°7'21" N," 82°25'10" W), 20-25 m 
depth. PARAT\TE: MZUCR-INB00014974:32 (1 speci- 
men), collecting locality Manzanillo, Limon, Costa Rica 
(9°38' N, 82°39' W), 20-25 m depth. 

cruzmoralai, Dviitiimirt^o, Espinosa and Ortea, 2000: 
110-113, fig. 11, pi. 2. Type locality: reefs from Manza- 
nillo, Limon, Costa Rica'(9°38' N, 82°37' W), 12-15 m 
depth. HOLOTYPE: MZUCR-INB000,3.349903 (shell. 
Figure 14). 

ctirere, Doto, Ortea, 2001a: 17-18, fig. 7, pi. 2C. T)pe 
locality: Puerto Viejo, Limon, Costa' Rica (9°38' N, 
82°39' W), 6 m deptli. HOLOTYPE: MZUCR- 

destinyae, Ctitliona, Hermosillo and Valdes, 2007: 
119-124, figs. IC, 4, 5. Type locality: La Godomia, Zi- 
huatanejo, Guerrero, Mexico (17°37.854' N, 10r33.562' 
W). PARATYPES: MZUCR-INB00031 18106 (4 speci- 
mens), Plava A\-ellanas, Guanacaste, Costa Rica 
(10°13.583' N, 85°.50.433' W). 

duao, Doto, Ortea, 2001a: 28-30. figs. 12- 13, pis. 2A, 
2K. T\pe locality: Punta Mona, Manzanillo, Limon, 
C;osta'Rica (9°37' N, 82°37' W), 10-19 m depth. 

echandiensis, Helicina, Richling, 2004: 271-277, figs. 
77-84. Type localit\-: La Amistad National Park, Las Al- 
turas Sector, Southern Cordillera de Talamanca, S of 
Cerro Echandi, Campamento Echandi, Puntarenas, 
Co.sta Rica (09°01'33" N, 82"40'12" W), 2840 m.a.s.l. 
HOLOTYPE: MZUCR-INB0003542520 (soft parts 
and shell. Figure 26). PARATYPES: MZUCR- 
INBO00:3574()64 (I specimen, soft parts and shell), 
MZUCR-INB()()();3542521 (1 specimen, soft parts and 
shell), MZUCR-INB0003428246 (19 specimens, shell 
and soft parts). Locality data for all paratypes same as 

F. Villalobos-Rojas et al., 2008 

Page 159 

eUzabethae, Adrana. Ortea and Espinosa, 2001e: 61- 
64, fig. 17. T\pe locality: in front of Gandoca beach, 
Limon, Costa Rica (9°36' N, 82°35' W), 10-15 m depth. 
HOLOTYPE: MZUCR-INB0003449558 (shell. Figs. 

escondida^ Helicina ("Gemma"), Richhng, 2004: 348- 
357, figs. 210-218. T>'pe locaht>': 9 km W of Matina, a 
little stream up the Rio Barbilla from the crossing of the 
road from Siquirres to Limon, along a tributary of Rio 
Barbilla, Limon, Costa Rica (10°03'29" N, S3''22'24" W), 
70 m.a.s.l. HOLOTYPE: MZUCR-INB0003542623 (soft 
parts and shell. Figure 27), PARATYPE: MZUCR- 
INB0003542624 (soft parts and shell). Locality data 
same as holohpe. 

espinosai, Ancula, Ortea, 2001b: 49, pi. 2D. Type 
localit\': Punta Mona, Manzanillo, Limon, Costa Rica 
(9°37''N, 82°37' W), 9 m depth. HOLOT\TE: MZUCR- 
I\B0003 188764. 

eugeniae, Elijsia, Ortea and Espinosa, 2002: 130-133, 
figs. 1-2, pi. lA. Ty-pe locality: Manzanillo, Limon, Costa 
Rk-a (9°38' N. 82°39' W), 16 m depth. HOLOTiTE: 

eversonii, PhijIIonotiis, D'Attiho, Myers and Shasky, 
1987: 162-164, figs. 1-2. Type Locality: SW side of Isla 
Manuelita. Isla del Coco, Costa Rica (5°33' N, 87°03' W, 
66 m depth. PARAT\TE: MZUCR 4934 (shell, Fig- 
ure 4). 

gandocaensis, Rissoella, Ortea and Espinosa, 2001a: 
36, pi. 2A. Type locality: Punta Mona, Manzanillo, 
Limon, Costa 'Rica (9°37' N, 82°37' W). 9 m depth. 
HOLOTYPE: MZUCR-INB0003323831. Remarks: shell 
destroyed, only dry soft body present, not figured here. 

genecoani, Plesiocijstlscus, Espinosa and Ortea, 2000: 
96-97, fig. 1, pi. 1. Type locahty: Manzanillo, Limon, 
Costa Rica (9°39' N, S2°37' W), 25 m depth. 
HOLOTYPE: MZUCR-INB0003349916 (shell. Figure 
22). PARAT\TES: MZUCR-INBOOO.3,349917 (2 speci- 
mens, shells). Locality data same as holot\'pe. 

goslineri, Cijlichnella , Valdes and Camacho-Garcia, 
2004: 459-497, figs. 4D-F, 5. Type locahty: Sector Playi- 
tas, Golfo Dulce, Puntarenas, Costa Rica (8°44'19" N, 
83°21'57" W), m depth. HOLOTYPE: MZUCR- 
INB0001497964 (shell. Figure 23). PARATYPES: 
MZUCR-INB0003718957 (15 specimens, sheU and soft 
parts, lateral gizzard plate and radular teeth in SEM 
stubs). Localitv^ data same as holotype. 

hojarasca, Alcadia (Microalcadia), Richling, 2001: 
■5-6, tigs. 6—8. Type locality: cordillera de Tilaran, about 
9 km N of Santa Elena, near Mirador Gerardo, Guana- 
caste, Costa Rica (10°22'19" N, 84°48'25" W), 1450 
m.a.s.l. HOLOT\TE: MZUCR-INB0003404979 (shell. 
Figure 25). Remarks: species was assigned to Alcadia in 
Richhng, 2004: 370-374, figs. 249-255; it was previously 

holandae, Pnimtm, Espinosa and Ortea. 1999b: 17.5- 
176, iio; IH. T\pe locahty: Cayos Limon, Islas San Bias, 
Colon, Panama (9°.33'ob" N, 78°53'30" W), 
HOLOTYPE: MZUCR-5750 (shell. Figure 10). 

PARATYPE: MZUCR-INB0003350838 (sheh), Punta 
Uyita, ManzaniUo, Limon, Costa Rica (9°38' N, 82M1' 
W). Remarks: According to the original pubhcation the 
holotype was deposited at the Instituto de Oceanologia 
de la Habana, Cuba, but in fact it was deposited at IN- 
Bio. The paratype was erroneously assigned the cata- 
logue number INBI0CR100150149S in the original pub- 
hcation. Also, tlie locality for the paratype, Punta Uvita, was 
misspelled in die original publication as Punta "Ubitas." 

inbiotica, Trapania, Camacho and Ortea, 2000: 317- 
321, figs. 1-3. T)pe locality: San Miguel Station, Reserva 
Absoluta de Cabo Blanco, Tempisque Conservation 
Area, Puntarenas, Costa Rica (9°34'49" N, 85°08'28" W). 
1 m depth. HOLOTYPE: MZUCR-INB00015008S9. 

iugula, Data, Ortea, 2001a: 26-27, fig. 11, pi. 2J. Type 
locality: Punta Mona, Limon, Costa Rica (9°39' N. 
82°37' W), 25 m depth. HOLOTYPE: MZUCR- 

kekoldi, Doto, Ortea, 2001a: 18-21, fig. 8, pi. 2 E-F. 
Type locality: Punta Mona, Limon, Costa^Rica (9°38' N, 
82°37' W), 9 m depth. HOLOTYPE: MZUCR- 
INB0003449587 (soft body and egg mass). 

ZeopoMoi, Eiibranchiis, Caballer, Ortea and Espi- 
nosa, 2001: 5.5-56, fig. 14, pi. 2E. T\pe locahty: Punta 
Mona, Manzanillo, Limon, Costa Rica (9°38' N, 82°37' 
W), 6 m depth. HOLOTYPE: MZUCR- 

lizanoi^ Pninum, Magana, Espinosa, and Ortea, 2003: 
124-126, figs. 2C, 3, pfs. IB, 2 B. T\pe locahty: Baliia 
Junquillal Wildlife Refuge, Golfo de Santa Elena, 
Guanacaste, Costa Rica. HOLOTYPE: MZUCR- 
INB000.3481195 (shell. Figure 11). PARATYPES: 
MZUCR-INB0003481192 (1 .specimen, shell, including a 
slide with radula), MZUCR-INB0003476231 (1 speci- 
men, shell and soft parts). 

loisae, Mitrella, Pitt and Kohl, 1979: 467-468, figs. 
2A-B, 3A-B. Type locality: N side of Punta Coralillo, 
Bahia de Caldera, about 20 km S of the city of Puntar- 
enas, Puntarenas, Costa Rica (9°54' N, 84°44' W). 
PARATYPES: MZUCR 2363 (2 specimens, sheU, Fig- 
ure 6). 

magagnai, Dendrodoris. Ortea and Espinosa, 2001d: 
52-53, fig. 13, pi. 3E. Type locality: Manzanillo, Limon, 
Costa Rica (9°38' N, 82°39'' W), 20 m depth. 

manzanilloensis, Polijcera, Ortea, Espinosa, and Ca- 
macho. 1999: 161-163, fig. 3. Type locality: Manzanillo, 
Limon, Costa Rica (9°38' N, 82°39' WJ, 8 m depth. 
HOLOT^TE: MZUCR-INB0001496124. 

marioi, Gibbenda, Espinosa and Ortea, 2000: 101- 
102, fig. 5, pi. 1. Type locahty: coral reefs of Manzanillo, 
Limon, Costa Rica (9°39' N, S2°39' W), 30 m depth. 
HOLOTYPE: MZUCR-INB0003371977 (a shde prepa- 
ration with radula). 

millenae, Cuthona, Hermosillo and Valdes, 2007: 
124-128, figs. ID, 6, 7. Type locality: Los Arcos, Bahi'a 
de Banderas, Jalisco, Me.xico (20°32.8.55' N, 105°17..340' 
W), 19 m depth. PARATYPE: MZUCR- 
INB0003836263, Playa Real, NE Punta Roble, Guana- 

Page 160 

THE NAUTILUS, Vol. 122, No. 3 

F. X'illalobos-Rojas et al.. 20()S 

Page 161 

Figures 24-32. 24. Holot)pes of Alcadia (Microalcadia) hoeckcreli, 1.1 mm. 25. Alcadia (Microalcadia) liojarasca, 2.4 mm, 26. 
Helicina ("Tristramia") echandiemls, 7.2 mm. 27. Helicina ("Gemma") escondida, 6.2 mm. 28. Helicina ("Gemma") monteverdensis, 
6.6 mm. 29. Helicina ("Gemma") talamancensis , 9.2 mm. 30. Helicina ("Gemma") beatrix rinpejensis, 7.8 mm. 31. Helicina ("Tris- 
tramla") punctisulcata cuericiensis, 5.9 mm. 32. Oligijra chiquitica, 4.8 mm. 

Figures 13-23. 13. Holohpes of Voharina i/olandae, 6.2 mm. 14. Dentimargo cnizmoralai, 2.5 mm. 15. Dcntiinargo zaidettae. 
2.5 mm. 16. Gibbenda bribri, 11.0 mm. 17. Gibbenda caribetica, 6.0 mm. 18. Gibbenda sierrai, 2.1 mm, 19. Gibbcnda ubitaensis, 
1.8 mm. 20. Granulina minae, 1.0 mm. 21. Plesiocysticiis alfiopivai. 1.9 mm. 22. Plesioct/sticus genccoani, 2.2 mm. 23. CijUchnella 
goslineri, 4.5 mm. 

Page 162 

THE NAUTILUS, Vol. 122, No. 3 

F. \'ill;dobos-Rojas et al, 2()()S 

Pase 163 

caste, Costa Rica {10°23.2()0' N, 85^50.733' W), 5 m 

minae, Gnniiiliiui. Espinosa and Ortea, 2()()(): 103, fig. 6, 
pi. 1. T\pe localiU': Punta Mona, Manzanillo, Liinon, Costa 
Rica i9°37' N. .Siz°37' W"), 12-15 m depth. IK)LOT\"PE: 
M7A'CR-IXR(KK)3349904 (shell. Figure 20). 

monteverdensis, Hcliciiia ("Gcniiiia"), Ricliliiig, 
2004: 334-34S, figs. 183-199. T)pe localit)': Cordillera 
de Tilaran, near Montexertle, Zona Protectora Arenal- 
Montexerde, Resena Biologica Bosque Nuhoso Mon- 
te\erde, Sendero Bosque Nuboso, Puntarenas, Costa 
Rica (lO^lS'OS" N, 84°47'41" W), 1550 m.a.s.l. 
HOLOTYPE: MZUCR-INB0003542627 (shell Figure 
28). P.-VRATtTE: MZUCR-INB0003542628 (shell). Lo- 
calih" data same as holotspe. 

orteai, Ci/crcv. X'aldes and Camacho-Garcia, 2000: 
445^56, figs. 1-5. T\pe locality': Plava Cabuya, Cabuya, 
Cobano, Puntarenas,' Costa Rica (9°39' N, 85=11' W), 
m depth. HOLOTYPE: MZUCR-INB0()01500521. 
PARATYPE: MZUCR-INB00()15()0644. Eocalitv data 
same as holotspe. 

orteai, Tripljora, Espinosa, 2001: 21-22, fig. 7. T\pe 
locality": Pimta Mona, Manzanillo, Limon, Costa Rica 
(9°37" N, 82°37" \V). 10-15 m depth. HOLOTYPE: 
MZUCR-INB0003138795 (shell. Figure 3). 

osae, Joninna. Camacho-Garcia and Gosliner, 2008: 
165-167. figs. 21-22. Tvpe locality: Plava Gallardo, 
Golfito, Osa'^Consen-ation Area, Costa Rica'(8°38'33" N, 
S3°13'40" W), m depth. HOLOTYPE: MZUCR- 
INB0003701453, MZUCR-INB000379944() (radula and 
jaw, SEM stubs). 

pacifica, Cliiapaplii/sa. Ta\lor, 2003: 170-171, fig. 
168, pi. 8.8. T\pe Iocalit\; Rio Tenorito, Hacienda 
La Pacifica, Guanacaste, ' Costa Rica (10°29.02' N, 
89°09.58' W). 100 m.a.s.l. PARATYPES: MZUCR- 
INB00033S2239 (10 specimens, shells, Figure 33). 

proranao. Data. Ortea, 2001a: 23-25, fig. 10, pi. 2H. 
T\pe localit\ : Punta Mona, \hmzanillo, Limon, Costa 
R'ica (9°37''N, 82°37' W), 9 m depth. HOLOTiTE: 

punctisulcaia cuericiensis, Hcliciiia. Richling, 2004: 
277-283, figs. 87-96. Tvpe locality: Cordillera de Tala- 
manca, Estacion Cuerici, 4.5 km E of \'illa .VUlls, Sen- 
dero al Mirador, Cartago, Costa Rica (9°33'28" N, 
83°40'1.3" W), 2700 m.a.s.l. HOLOTYPE: MZUCR- 
IXB0003542622 (shell. Figure 31). PARATYPE: 
MZUCR-IXB0003.542.541 (shell and soft parts). Other 
localitx- data same as holotxpe (09°33'19" N, 83°40'13" 
W. 2600 m.a.s.l.). 

sanctijohannis, Mai/abiiia, Taylor, 2003: 102-104, 
fig. 85, pis. 3.5, 3.6. Txpe localit)': Barra del Colorado, 
Limon, Costa Rica ■a0°46.37' N, 83°35.27' W). 
PAKAT\TES: MZUCR-IXB00033S2237 (10 specimens. 

shell), MZUCR 69-01 (10 specimens, shell. Figure .34). 
Locality data tor all paraty[5es same as liolot)pe. 

selva, Ercolania, Ortea and Espinosa 2001c: 45—47, 
fig. 11. Tvpe localit)-': Manzanillo, Limon, Costa Rica 
(9°39' N, S2°39' W),0 m depth. HOLOTYPE: MZUCR- 
INB00()3 13832. PARATYPE: MZUCR- 
INB0(J03449624. Localitv data same as holotyije. 

sierrai, Gibhenda, Espinosa and Ortea, 2000: 99-100, 
fig. 3, pi. 1. T)pe locality: Punta Mona, Manzanillo, 
Limon, Costa Rica (9°38' N, 82°37' W), 10-15 m depth. 
HOLOTYPE: MZUCR-INB0003.349909 (shell. Fig- 
ure 18). 

sinusdulcensis, Trojiiiuiiild, Taylor, 2003: 110-111, 
figs. 91-94. Type locality: small stream in pasture 3 km 
SE of Golfito, Puntarenas, Costa Rica (8°36.6S' N, 
83°8.48' \V). PARATYPES: MZUCR-INB0003382246 
(5 specimens, shells. Figure 36). 

socoae, Volvaiina, Espinosa and Ortea, 1999b: 171- 
172, figs. IE, 2D-E. Tyj^e locality: Manzanillo, Limon, 
Costa "Rica (9°39' N, 82°45' VV), 1.0-1.5 m depth. 
HOLOTITE: MZUCR-IXB0003350887 (shell. Figure 
12). PARATYPE: MZUCR-INB0003350888 (shell).' Re- 
marks: in the original publication the holotype and 
paratvpe wei'c eri'oneouslv assigned the catalogue num- 
bers iNBI0CR001496127'and INBI0CR1001496128, re- 

talamancensis, Hcliciiia ("Gemma"), Richling, 2001: 
3-5, figs. 3,4. Tvpe localitv: Fila Costena, N of Bajo Bo- 
nito, N of Rio C'laro, Puntarenas, Costa Rica (8"44'41" N, 
83°02'09" VV), 980 m NN. HOLOTYPE: MZUCR- 
INB0003404978 (shell. Figure 29). PARATYPES: 
MZUCR-IXB0001494642''(8 specimens, shell), 
MZUCR-INBO()01487761 (2 specimens, shell and soft 
parts), 3.5 km from Escuela de Llano Bonito Carretera a 
San Vito, Puntarenas, Costa Rica (8°44'27" N, 83°02'04" 
W), 840 m m.a.s.l., MZUCR-INB0001494509 (1 speci- 
men, shell and soft parts), 3.5 km XE from Escuela de 
Llano Bonito, Puntarenas, Costa Rica (8°44'54" X, 
83°02'04" W), 920 ni m.a.s.l. Remarks: species was as- 
signed to Helicina in Richling, 2004: 308-318, figs. 140, 
142-148; it was previously Oh/gira talamancensis. The 
paratvpe MZUCR-IXB()0()3389580 (1 specimen, shell, 
localitv data same as MZUCR-IXB00()1494509) is said to 
be deposited at INBio but is not present. 

temphquensis, Joninna. Camacho-Garcia and Gos- 
liner, 2008: 167-170, figs. 2.3-24. Tv^x' localit)': Cabo 
Blanco, Costa Rica (9°34'50" X, 8.5°08'26" W), m 
depth. HOLOTYPE: MZUCR-IXB0003542376. 
PARAT^TES; MZUCR-IXB0003542377 (2 specimens). 

tempinciiiensis, Mai/abina, Taylor, 2003: 109-111, tig. 
90, pi 4.4. 'I\pe locality: edge of marshes, 100 m E oi W 
end of airsti'ip, Palo Verde Xational Park, Guanacaste, 
Costa Rica (10°20.68' X, 85°20.60' W). PARATYPES: 

Figures 33-39. 33. Paratvpe oi Chiapaphijsa pacifica, 6.1 mm. 34. Paratope ot Mtn/ahiiia saiictijuhannis, 6.7 mm (UCR 69-01). 35. 
Paratvpe oi Maijabina tempisqiiensis, 7.5 mm. (INB00033S2244). 36. Paratype oi Tropinauta sinusdulcensis, 5.5 mm. 37. Holotype 
oi Cn/ptostrakon corcotadensis. 4 mm. 38. Holotvpe oiAdrana clizabcthae. (left valve dorsal view) 32.5 mm. 39. Ilolotvpe oiAdnina 
elizabethae, (left valve ventral view) 32.5mm. 

Pase 164 

THE NAUTILUS, \'ol. 122. No. 3 

MZUCR-INB()00:3382244 (10 .specimens, shell. Figure 
35), MZUCR 70-01 (10 specimens, shell). 

tica, Fiircilla. Espinosa and Ortea, 2000: 105-107, fig. 
7, pi. 1. Type locality: coral reels oi Manzanillo, Limon, 
Costa Rica (9°39' N, 82°39' W), 30 m depth. 
HOLOTYPE: MZUCR-INB000337197fi (including a 
slide with radula). 

tica, Mexichromis, Gosliner, Ortea, and Valdes, 2004; 
589-593, figs. lA, 2, 3. Type locality: Bajo del Diablo, 
Isla del Cafio, Puntarenas, Costa Rica (8°42'04" N, 
83°53'20" W), PARATYPES: MZUCR-INB()()1486639 
(2 specimens). 

iibitaensis, Gibbcnihi. Espinosa and Ortea, 2000: 97- 
99, fig. 2, pi. 1. Type locality: Punta Uxata, Manzanillo, 
Limon, Costa Rica (9°38' N,'82°41' W), 12-15 m depth. 
HOLOT\TE: MZUCR-INB00()3349902 (.shell. Figure 
19). Remarks: the T\pe localitw Punta Uvita, was mis- 
spelled in the original publication as "Ubita." 

i/olandae, Volvdrifiii. Espinosa and Ortea, 2000: 108- 
110, iig. 10, pi. 1. T\pe localitx': Manzanillo, Limon, 
Costa Rica (9°39' N, S2°39' W), 10-25 m depth. 
HOLOT^'PE: MZUCR-INB0003350843 (shell. Figure 
13). PARATiTES: MZUCR-INBOO()3350841 (1 speci- 
men, shell), MZUCR-1NB0003350842 (1 specimen, 
shell). Locality data same as lujlohpe. Remarks: accord- 
ing to the original publication, the paratype MZUCR- 
INB0003350841 was deposited at the' Instituto de 
Oceanologi'a de la Habana, Cuba, and the paratype 
MZUCR-INB0003350842 was deposited at Museo de 
Ciencias Naturales de Tenerife, Canary Islands; how- 
ever, both are deposited at MZUCR. 

zaidettae, Dcntimarop, Espinosa and Ortea, 2000: 
113-114, fig. 12, pi. 2. Type locality; Punta Mona, Man- 
zanillo, Linion, Costa Rica (9°38' N, 82°37' W), 12-15 m 
depth. HOLOTYPE: MZUCR-INB0003349911 (shell. 
Figure 15). PARATYPE; MZUCR-INB()()()3349910 (1 
specimen, shell). Locality data same as holoty'pe, Re- 
marks: According to the original publication the paraty'pe 
was deposited at the Instituto de Oceanologi'a de la Ha- 
bana, Culia, but in tact it is deposited at MZUCR. 

zugiiigae, Mchinella. Espinosa, Ortea and Magaiia, 
2001: 26-27, hgs. 8C, D. Type locality; Punta Mona, 
Manzanillo, Lim^on, Costa Rica (9°37' N, 82°37' W), 10- 
15 m depth. HOLOTYPE; MZUCR-INB0003138456 
(shell. Figure 2). 


We are vety grateful to Alvaro Morales and Ingo Werth- 
mann tor their logistic suppoit during the elaboration of 
this paper. The photographs were taken at the facilities 
of Centre de Investigacion en Ciencias del Mar y Lim- 
nologia (CIMAR) at the University of Costa Rica. Eu- 
gene V. Coan and David Riit\ill made constructive com- 
ments on this mamiscilpl. 


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una iiueva especie de Eii.hrinicliii\ Forl)fs, lrS34. A\icen- 
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Camacho-Garcia, Y. and T. Gosliner. 2004. A new species of 
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the zephyrinid nudihranch genus]aiwhis (MoUusca; Nudi- 
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THE NAUTILUS 122(3): 166-170, 2008 

Pao-e 166 

Dilemma japonicum new species (Bivalvia: Anomaloclesmata: 
Poromyiclae): A new record of the genus from the 
Northwest Pacific 

Takenori Sasaki 

The Universit}' Museum, 
The University of Tok\o 
7-3-1 Hongo, BunkAO-ku, 
Tokyo 113-0033 

Jose H. Leal 

The Baile\ -Matthews Shell Museum 
P. O. Box 1580 
Sanibel, FL 33957 USA 


The fourth species of the bivalve genus Dilemma Leal, 2008. is 
described from disarticulated valves collected off central Japan. 
The discovery of these specimens represents a significant range 
extension for the genus to the Northwest Pacific. The new 
species is distinguished from the other three known species by 
its surface sculpture, shape ot escutcheon, and hinge. The pro- 
dissoconch, indicative of lecithotrophic development, and shell 
microstructure, widr outer homogeneous and inner nacreous 
layers, are illustrated for the first time for a species of the 

Additional Keywords: Septibranchia. Poromxoidea. Japan, 
nacreous lax'er. pi'odissocondi 


unusual genus of 

Dilemma is a recently estabhshed 
Anomalodesmata (Leal, 2008). The first described spe 
cies of the genus, D. inexpcctatum (Crozier, 1966), was 
dredged from bathyal depths off northern New Zealand 
and originally assigned to Corciihim. family Cardiidae 
(Crozier, 1966) due to general similarities in shell shape. 
However, its original familial allocation was questionable 
from habitat and depth alone, because Corcidum species 
inhabit shallow lagoons of tropical waters and utilize pho- 
tosvnithesis-derived energy by symbiotic dinoflagellates 
(Farmer et al., 2001). Subsequent discoveiy of obviously 
related species with preserved soft parts allowed Leal 
(2008), using details of the macroanatomy, to demon- 
strate (1) that Corcuhim inexpectatum and the other, 
then newly found species are related and desen'ed 
grouping under a new genus, (2) that the new genus 
should be included in the Poromyidae, and (3) that spe- 
cies of Dilemma are carnivores. In fact, camivon' is a 
feeding habit that is common in the Anomalodesmata, a 
group of bivalves mostly present in the deep sea. 

Three species were originally (Leal, 2008) assigned to 
the genus: (1) Dilemma fnimarhernonim Leal, 2008, 

from off Key West Florida, 229 m, (2) D. spectralis Leal, 
2008, from off Vanuatu, 950-961 m, and (3) D. inexpec- 
tatum (Crozier, 1966} from off Three Kings Islands, New 
Zealand, 805 m. Their geographic distribution is disjunct, 
but shell moipholog)' is surprisinglv similar between the 
species of Florida and Southwest Pacific. 

Duiing a research cniise off central Japan, in 2000, die 
senior audior collected small unpaired \";il\es oi Corcidiim- 
likc bivalves, but its locality (240-273 m) was obviously 
too deep for Coreiihim. In addition, the inner shell layer 
is iridescent, indicating presence of a nacreous layer. 
This conchological feature alone precluded inclusion of 
this new species in the Cardiidae. However, lack of soft 
parts prevented a better resolved familial or supra- 
familial placement of the new taxon. The publication by 
Leal (2008) alkm'ed allocation of the unknown bi\alve in 
that newly established genus of the Porom\idae. In this 
paper, we describe diis most interesting new species and 
repoit a new record of the genus from the Northwest 


Four unpaired \al\es were sorted from sediments 
dredged from southeast of Kamogawa, Chiba Prefecture, 
Japan, with a biological dredge (RA'T..\nsei-MARU, cmise 
KT-00-05, statio'n 1, 240-273 m, 34°59.963' N, 
140°27.159' E- 35°00.020' N, 140°28.427' E, May 17, 
2000). All specimens were dead and disarticulated. The 
shells were coated with platinum and vanadium and pho- 
tographed with a scanning electron microscope (Hitachi 
S-2250N), folloudng a standard method (see Geiger et 
al, 2007). The holotype and two paratvpes are deposited 
in the Department of Historical Geolog\' and Paleontol- 
ogy, The University Museum, The Universits' of Tok"\'o 
(UMUT), and one paratvpe in The Bailev-Matthews 
Shell Museum, Sanibel, Florida (BMSM). 

T. Sasaki and l.H. Leal 2008 

Pac;e 167 

Figures 1— t. Dilcinniii jajionirum new species. 1. Lateral view. 2. Posterior view. 3-4. Interior view. 1-3. Left valve. 4. Right 
val\e. 1. Holotxpe, UMUT RM 29689. 2. Paratope L UMUT RM29690, 3. Paratyise 2. UMUT RM2969L 4. Parat)pe 3, BMSM 
1T9S3. Abbrexiations: ct = cardinal-like tooth; e = escntcheon; k = keel; It = lateral tooth; p = projection in hinge; s = socket. 


Superfamilv Poroimoidea Dall, LS86 
Family Poromyidae Dall, 1886 
Genus Dilemma Leal, 2008 

T>pe Species: Dilemmii fnimnrkcrnonim Leal, 2008, 
b\ original designation. 

Diagnosis: Sliell compressed anteroposteriorlv and 
expanded lateralK'; lateral outline of articulated valves 
cardioid; umbones projecting dorsally and located ante- 
riorly; sharp oblique carina dividing anterior and poste- 
rior regions; maximum growth a.\is having ca. 30° against 
anteroposterior a.\is; hinge a\is short with cardinal-like 
tootla and socket in each valve; lateral tooth reduced, 
present only in right valve; ligament external, double- 
lavered; lithodesma absent; shell interior lined with 
sheet-like nacreous laver. See Leal (2008) for anatomical 

Dilemma japonicum new species 
(Figures 1-12) 

Diagnosis: Posterior region ol shell sculptured by 
rough, lamellate, thin, commarginal ribs; surface irregu- 
larl\- punctate on anterior region but puncta radially ar- 
ranged on posterior region; escutcheon not distinctly de- 
marcated; prominent projection present in posterior re- 
gion of hinge. 

Description: Slu^ll tliin, fragile, compressed antero- 
posteriorK'; shell height larger than shell length. Anterior 
and posterior regions clearly demarcated b\ shaqi keel 
(Figure 1. kj: anterior region narrower and less inflated 

than posterior. Surface of anterior region smootli, flat- 
tened in macroscopic view, but microscopically punctate 
(Figure 5). Surface of posterior region sculptured by sev- 
eral weak radial ribs (Figure 3), and more prominently so 
by dense commarginal ribs, also pimctate in enlarged 
view, but puncta arranged neath' in radial direction un- 
like on anterior region (Figure 6). Umbo angulated by 
keel and rib along posterior margin (Figure 7); umbonal 
cavity deep; umbo involute (Figure 3). Prodissoconch 
simple, flattened, sliield-like, ca. 200 |xm in lengtli, with- 
out prominent sculpture or division into prodissoconch I 
and II (Figure 8). Hinge provided with single cardinal- 
hke tooth and socket on each valve (Figures 3-4, 7, ct, s); 
cardinal-like tooth located anterior to socket on left valve 
(Figure 3), theii- position reversed on right valve (Figure 
4). Lateral toodi small, present only on right valve (Fig- 
ure 4, It). Single shaip projection (Figures 3-4, 7, p) 
prominent on posterior side of hinge in similar position 
and size in right and left \'alves. Ligament and outlines of 
muscle scars not obsei-ved in disarticulated valves. Shell 
margin consisting of two layers, outer homogeneous 
structure and inner nacreous structure (Figure 9-10, 
OL, ID. In outer la)'er elongate granules arranged ver- 
ticalK' but lacking clear boundary. In inner layer suboval 
tablets fusing in growth region (Figure 11), numerous 
sheet-like layers forming nacre (Figure 12). 

Type Material: Holotyj^e, 4.7 mm (height: SH) x 2.0 
nim (length: SL), UMUT RM29689 (Figure 1); Paratype 
1, 5.6 mm (SH) x 2.4 mm (SL), UMUT RM29690 (Fig- 
ure 2); ParaHpe 2, 6.0 mm (SH) x 3.0 nuu (width), 
UMUT RM29691 (Figure 3); Paratvpe 3, 9.3 mm (SH) x 
4.0 mm (SL), BMSm' 17983 (Figure 4). 

Pase 168 

THE NAUTILUS, Vol, 122, No. 3 

Figures 5-8. Dilciiuiiii japoiiiiinn new species. 5-6. Detail of surface sciilptiiie, 5. Aiiteiior slope, (>. Posterior slope. 7. Detail of 
hinae. 8. Prodissoconch. 5-6. llolot>pe, UMUT RM29fi.S9. 7-8. Parat\pe 2, UMIJT RM29691. Abbreviations; ct = cardinal-like 
toodi; p = projection in hinge; pd = prodissoconch; s = socket. 

Type Locality: off Kamogawa, Chiba Pre- 
fecture, Japan', 240-273 m, 34°59.963'-35°()0.020' N, 
140°27.159'-140°2S.427' E. 

Distribution: Known onK liiini tlic t\pe localitw 

Etymology: The .species epithet recognizes the coun- 
try ot the type locality. The epithet japoniciim agrees in 
gender with the name Dilemma, a late-Latin neuter noun 
derived from the Greek. Of the previously named species 
in the genus, inexpectatiim is a neuter gender epithet, 
fntmarkernonun has a genitive ending that is not to be 
influenced by the gender of the genus, and spectmli.s is 
an epithet originallv used hv Leal (200S) as a noun in 


The inclusion ol the new species in the genus Dihinnui 
is well supported by the presence of the diagnostic shell 
characters oi the genus: anteroposteriorly compressed 
shell, cardioid outline in articulated valves, shai"p carina 
dividing the shell into die anterior and posterior regions. 

cardinal-like tooth and socket in each valve, and nacreous 
shell interior. Obviously there is no other group having a 
combination of these characters in bivalves. 

From the three previouslv kmov\ii species ol the genus. 
Dilemma inexpcciatum (Crozier, 19fi6) from northern 
New Zealand is most similar to the new species in having 
rough commarginal sculpture in the posterior region. 
However, the two species ditter in two characters: the 
rows of pores are absent and the escutcheon is distinct in 
D, inexpectatiim. As summarized in Table 1, the four 
species in the genus can be distinguished among them- 
selves by four shell chai^acters: sculpture in the posterior 
region; the presence or absence ot small pits; the dis- 
tinctness of the escutcheon; and the presence or absence 
of the posteiior projection of the hinge. The paired pos- 
terior projections are present only in D.Japonieiim; they 
are not part oi the hinge teeth system, because they do 
not aiiiculate as a tooth and a siK'ket between the right 
and Icll \al\cs. Uidcntunatelv, the (jutlines of muscle 
scars were unclear in the specimens ol the new species. 

The original description of Dileitima mentioned 
"Shell apparently nacreous internally . . ."' (Leal, 200S: 3), 
but did not oiler any detailed descriptions or discussions 

T. Sasaki and ].H. Leal, 200S 

Pasre 169 

Figures 9-12. Dilemma japonicum new species. Sliell microstructure. 9-10. (Jbsened at small broken part ot shell margin, vertical 
section of homogenous structure in outer layer (OL) lined by nacreous structure in inner layer (IL). 11-12. Obhque (11) and 
horizontal (12) \ie\vs of nacreous laver at inner shell surface. Parat)-pe 2, UMUT RM29691. 

on tlie shell microstnjcture ot the three originally in- 
cluded species. The description of the inner nacreous 
shell la\'er in the present stud\' confirms its presence in 

A shell consisting of outer homogeneous and inner 
nacreous layers (Figure 9-12) supports inclusion of 
the new species in Anomalodesmata (Tavlor et al, 1973; 
Prezant, 1998). The niicrostiiicture of the outer la\er is 

somewhat similar to a simple prismatic structure in that 
elongate granules are arranged veitically. The outer layer 
is identified as a homogeneous stmcture, because, imlike 
t}'pical prismatic stmcture, ciystals lack shaqj bound- 
aries. It should be observed, however, that environmen- 
tal changes can alter otherwise organized shell micro- 
stiTJCtures to appear irregular, smoothed, and homoge- 
neous (R. Prezant, pers. comm.). 

Table 1. Shell characters and distribution of the four Dilemma speci 

D. fniiaarkernonim 

D ^peelralis 

D iiiexpeetaliiin 

D. Japoiiieiim 
new species 

Sculpture in posterior Dense, coarse growth 

region lines 

Small pits on exterior Absent 


Escutcheon Not clearK" separated 

Projection posterior Absent 

to beak 

Distribution Off Kev West Florida, 
USA'. 229 m 



.\ot clearK" separated 

Oft \'anuatu. 

Southwestern Pacific, 
9.5t)-961 0] 

Rough commarginal 

ribs and grooves 

ClearK defined 

Off Three Kings 

Islands, New Zealand, 
S0.5 m 

Rugose, lamellate 

commarginal ribs 

Not clearK separated 

Off central Japan, 
Northwest Pacific, 
24(V27:3 m (dead) 

Pase 170 

THE NAUTILUS, Vol, 122, No. 3 

The prodissoconch oi Dilemma japonicum new species 
indicates non-planktotrophic, lecithotrophic develop- 
ment. It measures ca. 200 |JLm in length (Figure 8), a size 
that fits into the predicted size range (135-230 |jL,m) of 
prodissoconchs of bivalves with lecithotrophic develop- 
ment (Ockelmann, 1965; Jablonski and Lutz, 1983). The 
developmental modes of the other species of the genus 
are unknown. 

Although two of the other live-collected species of 
Dilemma were found attached to hard substrata, the mi- 
crohabitat of the new species is unknown. The bottom 
sediment from the type locality contained numerous 
dead shells and other biogenic fragments such as those of 
bryozoans and sponges. Because most bivalves were dead 
and disarticulated, large part of samples in the dredge 
haul was inferred to be transported and accumulated. 
Specifying the actual microhabitat of the species is an 
interesting target for future sampling. 

The description of Dilemma japoniciim extends the 
geographic range of the genus to the Northwest Pacific, 
in addition to previous records from oif Florida and the 
Southwestern Pacific (off Vanuatu and New Zealand). 
The genus has a broad geographic range, extending 
across the Panama land bridge and Eastern Pacific bar- 
rier. Future sampling of deep-sea hard substrates may 
yield additional records of Dilemma species from other 
locations in the Pacific and in other oceans. 


The authors are indebted to the reviewers, Riidiger 
Bieler, Robert S. Prezant, and Richard C. Willan, for 
their comments and suggestions. Richard Willan and Rii- 
diger Bieler helped with etymologies and confirmed the 
gender of the name Dilemma. The samples of the new 
species became available with the kind assistance of Dr. 
Suguru Ohta (formerly Ocean Research Institute, Uni- 

versity of Tokyo), other colleagues on board and crew of 
WW Tansei-Maru (Japan Agency for Marine-Earth Sci- 
ence and Technology, formerly Ocean Research Insti- 
tute, University of Tokyo) during the cnaise KT-00-05. 


Crozier, M. A. 1966. New species and records of MoUusca from 
off Three Kings Islands, New Zealand. Transactions of the 
Royal Society of New Zealand, Zoology 8: 39^9. 

Fanner, M. A, W. K. Fitt, and R. K. Trench. 2001. Moqshology 
of the symbiosis between Corculum carclissa (MoUusca: 
Bivalvia) and Symbiodiniuni corculonim (Dinophyceae). 
Biological Bulletin 200; 336-343. 

Geiger, D. L., B. A. Marshall, W. F. Ponder, T. Sasaki, and 
A. Waren. 2007. Techniques for collecting, handling, and 
preparing small molluscan specimens, Molluscan Re- 
search 27(1); 1-49 

Jablonski, D. and R. A. Lutz. 1983. Lai-val ecology of marine 
benthic invertebrates; paleobiological implications. Bio- 
logical Reviews 58: 21-89. 

Leal, J. H. 2008. A remarkable new genus of carnivorous, 
sessile bivalves (MoUusca: Anomalodesmata: Poromyidae) 
with descriptions of two new species. Zootaxa 1764: 1-18. 

Ockelmann, W. K. 196.5. Developmental types in marine bi- 
valves and their distribution along the Atlantic coast of 
Europe. In: Cox, L. R. and Peake, J. F. (eds.) Proceedings 
of the First European Malacological Congress, London, 
1962, Conchological Society of Creat Britain and Ireland, 
and the Malacological Society of London, London, 
pp. 25-35. 

Prezant, R. S. 1998, Subclass Anomalodesmata, In; Beasley, 
P, L„ Ross, C, J, B, and Wills, E, (eds,) MoUusca: The 
Southern Synthesis, Fauna of Australia, Vol, 5, Part A, 
SCIRO PubUshing, Melbourne, pp, 397-249, 

Taylor, J, D„ W. D. Kennedy, and A. Hall. 1973. The shell 
structure and mineralog)' of the Bivalvia. II. Lucinacea- 
Clavagellacea, Conclusions. BuUetin of die British Mu- 
seum"(Natural History). Zoology 22: 2.5.5-294. 

THE NAUTILUS 122(3): 171-1' 


Page 171 

On the genus Heteroschismoklcs Ludbrook, 1960 (Scaphopocla: 
Gadilida: Entalinidae), wdth descriptions of two new species 

\'ictor Scarabino 

Departement S\stcinatimic et Exolutioii 
Museum national d'Histoire naturelle 
UMS Taxonomie-Collections CP51 
55 iiie de Buffon 
75005 Paris, FRANCE 

Carlos Hennquc Soares Caetano 

Departaincaitt) do Zuologia, 

Universidade Federal do Estado do Rio de 


Av. Pasteiu-, 45S 

Rio de Janeiro, 22.290-240 BRAZIL 


Heteroschismoides is a deep-sea genus characterized by shells 
being beh\'een 10 and 20 mm as adults, sculptured by 9 to 10 
prominent primary' ribs and a unique deep irregular apical fis- 
sure on dorsal side, considered until now to include a single 
species: Dentaliiiin subtcifissum Jeffreys, 1877. During revision 
of material from se\'eral expeditions carried out by the Museum 
National d'Histoire Naturelle, Paris, hvo new species were 
identified and are here described: H. meridionalis new species 
and H. antipodes new species In addition, designation of die 
lectohpe of H. subterfisstis is proposed, as well as new records 
for this species in tlie nordieastem Atlantic ocean are given. 
Heteroschismoides meridionalis new species is closely related 
to H. subteifissus, but die first has a smaller ma.ximum diameter 
of shell and smaller apical aperture diameter. Heteroschis- 
moides antipodes new species from Solomon Islands is smaller 
dian odier two species for both, shell length and lissure exten- 
sion. The distance of point of maximum curvatuie from the 
apex in H. antipodes new species is located nearer to the apex 
than in H. meridionalis new species and H. subterfissus. The 
results here obtained considerably enlarge the geographical 
distribution of the genus and suggest a worldwide batliyal and 
ab\ssal distribution for diis genus. 

Additional Keywords: Heteroschismoides siibteijissus, Het- 
eroschismoidinae, tusk shell, lectotype, new species, Brazil, 
Solomon Islands, deep-sea 


The genus Heteroschismoides was proposed b\' Lud- 
brook in 1960 to include the deep sea species Dcntalium 
subterfissum Jeffreys, 1877 (for further supraspecific in- 
formation see Steiner and Kabat, 2001). The most re 
markable character of die genus is a unique, deep and 
irregular apical lissure at die dorsal side of shell not present 
among other Scaphopoda (Chistikov, 1982; Scarabino, 
1995). Since Chistiko\- (1982), this genus has been con- 
sidered as monospecific and restricted to the Atlantic 
Ocean. During tlie revision of material from several e.x- 
peditions carried out bv the Museum national d'Histoire 

naturelle, Paris, in the northeastern and southwestern 
Atlantic Ocean (Brazil) as well as off Solomon Islands in 
the Pacific, a number of specimens corresponding to the 
genus were identified. In a preliminan,- sorting, Brazilian 
specimens were placed under H. subteifissus. but the 
unexpected finding ol representatives in the tropical Pa- 
cific, decided us to undertake the present revisiorl. Here, 
we propose the lectotype of Heteroschismoides subter- 
fissus and describe two new species, extending consider- 
ably the geographical distribution range ol the genus. 


The material was collected during several expeditions car- 
ried out bv the MNHN and Ifremer, known as BiOGAS I to 
XII (1972-1981), Thalassa (1970-73) and Incal (1973): 
in the Gulf of Gascony; Abitlaine (1982): off Portugal 
and Spain; NoR.ATLANTE (1969): large coverage of the 
North Atlantic; Bi./vgoRES (1971): off Azores; Bio\'EMA 
(1977): Vema Trench; MD55 (1987): Southeastern Bra- 
zil; and Salomon 2 (2004): off Solomon islands. 

In \iew to better define the species, a morphometric 
analysis was attempted based on selected undamaged 
shells of the three species. Shell measurements were 
taken according to Shimek (1989), Steiner (1999) and 
Steiner and Linse (2000), including length (L); maximum 
diameter (Max), that is in this case equal to the anterior 
aperture diameter; nniximuni cun'ature (Arc); distance 
of point of maximum cui'vature from the apex (Larc); 
apical aperture diameter (Apd). Besides those measure- 
ments, we also estimated the extension of the apical fis- 
sure (Fiss) located at dorsal side of shell. To assure in- 
dependence among variables, we performed a prelimi- 
naiy correlation analysis among all variables with strongly 
correlated variables (r > 0.70) being excluded. The non- 
parametric Kruskal-Wallis test was emploved to examine 
for differences of each moi"phometric parameter (un- 
transformed data) between species. Dunn's multiple 
comparisons test was used a posteriori to assess signifi 
cant differences between species. A multi\'ariate approach 

Paee 172 

THE NAUTILUS, Vol. 122. No. 3 

was carried out utilizing Discriminant Function (DF) 
Analysis to integrate all nioiphonietric data in a single 
anaKsis. To perform this anah'sis, we standardized the 
nioq^hometric data following Romesburg (1984). 

Institutional abbreviations used in tlie text ai"e: BMNH: 
The Natiu-;il Histoiy Museum, London; IFREMER: Insti- 
tut Franij-ais tie Recherche pour I'E.xploitation de la Mer: 
MNR}: Museu Nacional, Rio de Janeiro, Brazil; MNHN 
Museum national d'Histoire Naturelle, Paris; USNM 
National Museum oi Natural Histow, Smithsonian Insti- 
tution, \\'ashington, DC, US.A.. Other abbre\iations are: 
CP: beam trawl; DC: '"Charcot" dredge; stn: station; Iv: 
live-collected; dd: shell only. 


Order Gadilida Starobogato\', 1982 
Suborder Entalimoqiha Steiner, 1992 
FamiK' Entalinidae Chistikov, 1979 
Subfamily Heteroschismoidinae Chistikov, 1982 
Genus Heteroschismoides Ludbrook, 1960 

Type Species: DentdUuui suhtcifissu}ii Jeffreys, 1877 
(by original designation). Recent, northeastern Atlantic 

Description (Modified fi-om Scarabino, 1995: 302): 

Shell up to 20 nuu length, slightly cuived, regnlarlv ta- 
pering. Translucent grey when the animal is alive, 
opaque to polished white or cream when empt}'. Longi- 
tudinally sculptured by 9-10 primaiy ribs that reach the 
anterioi- aperture or vanish towards it. Intercostals spaces 
convex or straight, smooth or finely striated longitudi- 
nally bv 8 to 14 lines. Secondaiy ribs, predominanth- a 
single one, can be present in each intercostal space. Apex 
with a long and wide irregulai- fissm-e on dorsal side earlv 
obsen-ed in late embiTonic stages and juvenile shells. 
Section polygonal, more notorious at the central zone of 
shell, fading or not towards the slightly laterally com- 
pressed thin oral aperture in adult specimens. Shell frac- 
tures and repairs are frecjuentK' obsened, as well as 
twisting of shell. 

Radul.V: Rachidian tooth with anterior margin 
rounded and lateral margins thick; laterals with sharp 
pointed primaiy cusps and 4 important denticles; mar- 
ginal slightly curved with conspicuous lateral processes, 
better obsen-ed in light microscope. 

Di.stribution: Recent, Atlantic Ocean and tropical Pa- 
cific Ocean, batlnal-abvssal. 

Remarks: Scarabino (1995: 302) mentioned bv mis- 
take ■"10-12 primaiT ribs"; in the light of new information 
this numlier is here corrected as 9 to HI piiniai'v ribs. 

Molecular data of H. subtcifisstis has been published by 
Steiner and Dreyer (2003).^ 

Hctemscltismoiclcs subtciji.\sus (Jeffreys, 1877) 
(Figures 1-6, 15) 

Dcittaliuin subteifissum Jeffreys, 1877: 154; 1883: 660, pi. 49, 
fit^. 3; Watson, 1879: 516; 1886: 10, pi. 1, fia;. 10; Pilsbiy 
and Sliaqj, 1897: 61, pi. 7, figs. 15-19; Warc=n 1980: 53. 

Dentalium (Hetewschisma) suhteifissum: Henderson, 1920: 58. 

Dentaliinn (Dentalium) suhteifismm: Nickles 1979: 47, fig. 5a b. 

HctcroscJiismoides subteifissus: Chistikov, 1982: 675, figs. 3, 5; 
Steiner 1998: 78; Steiner and Rabat, 2001: 446; 2004: 651. 

Description: Sliell up to 9.2 mm length, slightly 
cun'ed, regularly tapeiing. Translucent shiny grey, pres- 
enting fractures and repairs. Longitudinal sculpture of 
nine primaiy notorious ribs and a single secondaiy one in 
between, more notorious in the dorsal half, all reaching 
the thin oral aperture. Intercostals spaces convex, finely 
striated longitudinalK' by eight lines. Apex with a 2.3 mm 
long, wide, irregular fissure on dorsal side. Trans\'ersal 
section polygonal, slightK' compressed laterally, less ap- 
parent at apex. 

Type Material: Lectotype, here selected (see remarks), 
the largest (9.2 mm) of the three specimens of lot USNM 
175018, Ireland, 54°19' N,ir50' W, 1180 fms [2158 m] 
(Porcupine 1869 stn 16); other paralectohpes: USNM 
175017, Off West coast of Ireland, (Porcupine 1869 stn 
19a), 2 ,specimens; USNM 175019, Greenland, 56°11'N 
37°41'W, 1450 fms [2646 m]. Valorous stn 12, 1 speci- 
men; USNM 175020, 55°4()'N 12°46'W, 1476 fms [2694 
m]. Porcupine 1869 stn 21, 3 specimens; off Azores, 
Challenger E.xpedition stn 78, 37°26' N 25"13' W, 
Azores, 1000 fathoms [1825 m], 7 specimens and 3 frag- 
ments, BMXH 1887.2.9.36-40. Two other paralecto- 
tvpes from Porcupine expedition are deposited at 
BMNH under code 1885.11.5.1393-4, with a mention 
"further synt)pes in USNM 175017-020". 

The fact of the species has been partially desciibed on 
the base of a young specimen (see Remarks) and that the 
specimen illustrated was not found in the collections re- 
vised, lead us to designate the lectot\pe based on an 
adult shell. 

Measurements of Lectotvpe (mm): Length 9.20, 
oral aperture 1.00, apex 0.12, arc 0.42 at 1.9 from apex. 

Tyije Locality: West of Ireland, 54°19'N, ir50'W, 
1489 m. Porcupine 1869, stn 16 (here selected). 

Other Material Examined: Off Gabon (Africa), 2°32' 
S, 8°18' E, stn DS20. 2514 m (1 dd); Gulf of Guinea, 
4°40'N, 5°39'2E); stn DS3(). 3109 m (8 dd) (bodi W.^lda 
E.xpedition, CNEXO, Nickles, 1979), all MNHN. 

Figures 1-14. Heteroschisinokles species. 1-6. H. subteifissus (JeClreys, 1977). 1. Lectotype USNM 175018, 9.2 mm. 2. Para- 
leclotvpe. .3. Detail of tjie apical fissure. 4. Specimens showing embiyonic sliell. 5. Detail of longitudinal ribs. 6. Detail of 
microseulpture on the outer surface of intercostals space. 7-10. H. nichdioiialis new species. 7. Holot\pe, MNHN, 14 nun. 8. 
Paratype, detail of apex. 9. Detail of sculpture. 10. l^etail of microseulpture on the outer surface of intercostals space. 11-14. 
Hcleroschismoides nnlipodes new species. 11. jioiotjpe, MNHN, 9.95 nini. 12. Detail o( upc\. 1.3. Detail of sculpture. 14. Detail of 
microseulpture on the outer surface of intercostal space. 

y. Scarabino and C. H, S. Caetano, 2()()(S 

Page 173 

Page 174 

THE NAUTILUS, Vol. 122, No. .3 

Figures 15-17. Radulae of Heteroschismoicles species. 15. H. subteifissus, rachidian and lateral teeth, internal view. 16. H. 
antipodes new species, external vdew of lateral tooth (left), I'achidian, lateral, and marginal teeth (right), in internal view. 17. H. 
meiidionalis new species: rachidian and lateral teeth, internal view. 

New Records: Central Atlantic Ocean: Azores Islands: 
BugoRES 1971: stn DS 54, 38°12' N, 2S°15' W, LSIO ni, 
2 dd, stn DS 165, 37°33' N, 25°58' W, 2085-2050 ni, 1 
dd, DS 173, 37°57' N, 26°08' W, 3225 m, 1 dd; Norat- 
LANTE 1969: stn 02, 53°55' N, 17°52' W, 2456 - 2420, 3 
Iv; stn 84, 36°21' N, 08°43' W, 2871-2875 m, 3 Iv, 3 dd; 
stn 85, 36°25' N, 08°48', 2573-2580 m, 2 Iv, 3 dd; 
Thalassa 1971, Stn X 336, 44°11' N,05°10' W, 1850- 
2050 m, 1 Iv, 1 dd; Eastern Atlantic Ocean: Incal 1976: 
stn. DS 01, 57°59' N, 10°40' W', 2091 ni, 56 Iv, 7 dd; stn. 
DS 02, 57°59' N, 10°49' W, 2081 m, 51 Iv, 3 dd; stn. DS 
05, 56°28' N, iri2' W, 2503 m, 24 Iv, 7 dd; stn. DS 07, 
55°01' N, 12°31' W, 2884 m, 4 Iv, 1 dd; stn. DS 07, 
56°27' N, iril' W, 2884 ni, 5 Iv, 1 dd; stn. DS 08, 
55°02' N, 12°35' W, 2891-2884 m, 3 Iv, 2 dd; stn. DS 09, 
55°08' N, 12°53' W, 2897 m, 2 Iv, 2 dd; stn. CP 01, 
57°58' N, 10°55' W, 2040-2068 m, 12 Iv; stn. CP 02, 
57°58' N, 10°42' W, 2091 ni, 2 dd; stn. CP 03, 56°38' N, 

Figure IS. Discriminant function analysis of shell iiiorplici- 
irietric parameters of HetenKrhis}iii>idi.'s species. 

O H.merldlonalis new speaes 

O H. ant/podssnewspedes 



• H suOfarflssus ( Jetlreys, 1877) 






ir06' W, 2466 m, 3 Iv. 1 dd; stn CP 04, 56°33' N, 11°11' 
W, 2438-2513 m, 7 h: 7 dd; stn CP 05, 56°00' N, 12°29' 
W, 2884 m, 1 dd; stn CP 06, 55°02.3' N, 12°40' W, 
2888-2893 ni, 3 Iv, 1 dd; stn CP 07, 55°03' N, 12°46' W, 
2895-2897 m, 1 dd; stn CP 09, 50°15' N, 13°16' W, 
2659-2691 m, 2 Iv; stn WS 01, 50°19' N, 13°08' W, 
255(V2539 ni, 5 K-; stn WS 02, 50°19' N, 12°55' W, 
2498-2505 m, 1 Iv; stn WS 03, 4S°19' N, 15°23' W, 4829 
m, 2 dd; stn WS 09, 47°28.8 N-47°27.9' N, 09°34' W, 
4277 m, 1 dd,; stn OS 01, 54°14' N, 13°ir W, 2634 m, 
1 Iv; BiOG.AS II: stn DS 32, 47°32' N, 08°05' W, 2138 m, 
3 Iv, 3 dd; BiOGAS III: ,stn DS 37, 47°:32' N, 08°35' W, 
2110 m, 2 dd; BIOGA.S IV: stn DS 52, 44°06' N, 04°22' W, 
2006 m, 2 dd; stn DS 51, 44°11' N, 04°15' W, 2430 m, 
1 Iv; stn DS 58, 47°34' N, 09°08' W, 2775 ni, 3 dd; stn 
DS 62, 47°33' N, 08°40' W, 2175m, 1 dd; ,stn DS 64, 
47°29' N, 08°35' W, 2156m, 4 Iv, 1 dd; BIOGAS V: stn 
CP 07, 44°09.8' N, 04°16.4' W, 2170 m, 5 dd; Biogas VI: 
stn DS 61, 47°34.7' N, 08°38.8' W, 2250 m, 1 Iv, 1 dd; 
stn DS 62, 47°33' N, 08°40' W, 2175 m, 1 dd; stn DS 63, 
47°33' N, ()8°35' W, 2126 m, 1 Iv; stn CP OS, 47°03' N, 
08°39' W, 2177 m, 2 K , 1 dd; stn CP 09, 47°33' N, 08°44' 
W, 2171 m, 1 Iv; stn CP 24, 44°08' N, 04°16' W, 1995 m, 
3 dd; stn DS 71, 47°34' N, 08°34' W, 2194 m, 3 Iv, 5 dd; 
stn DS 86, 44°04.8' N, 04°1S.7' W, 1950 m, 1 dd; stn DS 
87, 44°05' N, 04°19' W, 1913 m, 3 Iv; Biogas VII: stn CP 
26, 47°33' N, 08°34' W, 2115 m, 1 Iv; Biogas VIII: stn 
KG 145, 47°33' N, 08°39' W, 2170 m, 1 dd; stn KG 147, 
47°33.40' N, 0S°40.7()' W, 2190 m, Iv, dd; stn KG 149, 
47°33' N, 08°39' W, 2165 m. 1 M: stn KG 151, 47°34' 
N, 08°39' W, 2205 m. 1 \v: Biogas IX: stn CP 34, 47°32' 
N, 08°25' W, 1970 in, I dd; Bi< .gas XI: stn CP 37, 47°34' 
N, 08°41' W, 2175 m, 2 Iv. 

Distribiilion: Eastern Atlantic Ocean: Greenland, off 
West hihiiid. Gulf ol C^ascony, oil Spain and Portugal to 
Gulf III (iiiinea. West Africa (|effreys, 1877; Watson, 
1879, IS8(i, Locard, 1898; Nickles, 1979). Alive in 940- 

\\ Scarabino and C. H. S. Caetano, 2008 

Paee 175 

29S7 111, shells downi to 4S29 m (present paper). Richest 
depth for h\e material: 2040-2503 m (present paper). 

Remarks: |etlre\s paitialK' described the species based 
on a jmenile specimen, since he mentioned ". . . posterior 
end bulbous" and illustrated this moiphologv in plate 49, 
fig. 3. Ho\ve\er, no juvenile specimen was detected 
among the t^pe material. In addition, he wrote that the 
slit is in the "under or ventral side", which in tact corre- 
sponds to the dorsal side in normal position of the ani- 
m;il. Jeffre\s also mentioned the presence of 12 to 16 
ribs, but we did not find an\' specimens with that number 
of ribs. Howexer, it is possible that secondaiy ribs and 
intercost;il striae might have been, in part, counted in- 
appropriateh' for the original description. To clarify, in 
the description \\'e enumerated only the priman' ribs 
because the number of secondaiy ribs varies wdth age 
and among specimens but the number primaiy ribs is 
constant among specimens and through the life of the 
specimen. Specimens with embiyonic shell are often ob- 

Hetcroschismoides .siibtcifissus was also mentioned in 
the Challenger expedition from "stn 120 - 8°37' S 
34°28' W, Pemambuco, South America 675 fathoms 
[1232 111] red mud" (Watson, 1886). The lot in which this 
reference was based was not located in B.VINH collec- 
tions bv the senior author. In the distribution paragraph, 
V^'atson (1886) wTote "Habitat.- Davis Strait. 1785 fath- 
oms. North Atlantic; various Stations off the West Coast 
of Ireland. 1180 to 1476 fathoms (Jeff.)." He did not 
mention Pemambuco. 

The material from Pemambuco iiia\ liekjiig to the new 
species described below or, as suggested by Henderson 
(1920; 58), could also correspond to specimens of Per- 
tiisiconcha caUitluix (Dall, 1889), since young specimens 
of diis species, specially those with a broken apex, can 
easily be misidentified as species of Hetcroschismoides 
(Scarabino, pers. obseiv). 

Hetcroschismoides meiidionaUs new species 
i Figures 7-10, 17) 

Description: Shell 14 mm length, slightlv cui"ved, 
regularlv tapering, opaque white. Longitudinal sculpture 
composed of nine rounded-edge primary ribs, all reach- 
ing but fading towards the oral aperture. No secondan* 
ribs. Intercostal spaces concave in posterior three quar- 
ters and straight to convex at tlie anterior fifth, present- 
ing 12 verv' faint, fine striae throughout. Apex with a 2 
mm long, wide irregular fissure on dorsal side. Cross- 
section polygonal, less angular at apex and at the slightly 
laterally compressed anterior aperture. 

Measurements of the Holotvpe (mm): Length 14.0, 
oral aperture 1.0. apex O.I. arc 0.8 at 6.7 from apex. 

T\pe Material: Holot\pe (Iv) VINHN 20902 and 14 
paratvpes (dd), 13 MNHN 2090.3-20906, 1 MNRJ 

T\'pe Locality: Off Espirito Santo, Brazil, 1S°59.1' S, 
37=47.8' W. 1540-1550 m [MD 55 stn DC 70]. 

Material Examined: Southwestern Atlantic Ocean; 
MD 55 stn CP 68, 18°55.6' S, 37°49.1' W, 1200-1500 m, 
1 dd (paratyix- MNHN 20903); stn DC 70, 18°59.1' S, 
37°47'8 W, 1540-1.550 m, 1 Iv (liolot)pe MNHN 20902), 
26 dd (7 paratvpes MNHN 20904,' 1 parat)pe MNRJ 
12707); stn DC'72, 19°00.4' S, 37°48.8' W, 950-10,50 m, 
4 dd (paratypes MNHN 20905); stn CB 77, 19°40.6' S, 
37°48.1' W, 790-940 m, 9 dd (2 paratypes MNHN 
20906); stn. CB 79, 19°01.8' S, 37°47.8' W, 1500-1575 
m, 10 dd; stn CB 106, 23°54.2' S, 42°10.5' W, 830 m, 15 
dd; stn. CB 107, 24°00.3' S, 42°14.4' W, 1020 m, 1 1 dd. 

EtAinology: MeridionaUs: Southern. Name refers to 
tlie taxon's distribution in the Southem Hemisphere. 

Distribution: Brazil; Espirito Santo and Sao Paulo. 

Alive in 1540 m, shells in 790-1540 m depth. 

Hcteroscliismoidcs antipodes new species 
(Figures 11-14. 16) 

Description: Shell to 9.95 mm lengdi, slightly cui-ved 
at the apex, regularly tapering. Opaque white. Longitu- 
dinal sculpture of ten primaiy ribs, all reaching the oral 
aperture. Intercostal spaces concave and smooth. Apex 
with a wide irregular fissure 1.7 mm long on dorsal side. 
Cross section polygonal, less obvious at apex, slightlv lat- 
erally compressed at the anterior aperture. 

Measurements of Holotype (mm): Length 9.95, oral 
aperture 0.9, apex, 0.1, arc 0.4 at 2.76 from apex. 

Type Material: HoloHpe (dd) MNHN 20907 and 10 
paratypes (7 Iv, 3 dd) MNHN 20908-20910 

Type Locality: Solomon Islands, 07°49.3' S, 
157°41.2' E, 104.5-1118 in [Salomon 2 Stn. CP2217]. 

Material Examined: Solomon Islands. SALOMON 2 
stn CP2182, 08°47.0' S, 157°37.9' E, 762-1060 m, 16 dd: 
stn CP2197. 08°24.4' S, 159°22.5' E, 897-1057 m, 2 Iv. 
2 dd (3 paratvpes MNHN 20909); stn CP2217, 07°49.3' 
S, 157°41.2'' E, 1045-1118 m, 1 Iv, 3 dd (holot^pe 
MNHN 20907 and 3 paratypes MNHN 20908), 'stn 
CP2218, 07°56.3' S, 157°34.6' E, 582-S(-i4 m, 4 Iv 
(paratypes MNHN 20910) 14 dd; stn CP2253, 7°26.5' S, 
156°15.0' E, 1200-1218 111, 10 dd. 

Etymology: From Greek antipodes, meaning those 
living on dianietrallv opposed places on Earth. 

Distribution: Solomon Islands, alive in 1200 ni; shells 
762-1200 ni depth. 


Table 1 lists the mean, standard deviation, minimum, 
and maximum values for each nioiphometiic parameter 
for the three species. Significant differences in all mor- 
phometric parameters were obsei-ved between the spe- 
cies (Kmskal-Wallis test; p<0.01), except for die maxi- 
mum curvature (Arc). This univariate comparison 
showed that H. meridionaUs new species and H. siibter- 

Page 176 

THE NAUTILUS. Vol. 122, No. 

Table 1. Shell moiphometric parameters with minimum (min), maximum (max), mean values and standard deviation (+SD) for the 
three new species of Heteroschismokles. L: length; Max: maximum diameter {= anterior aperture diameter); Arc: maximum cun'ature; 
Larc: distance of point of maxiniinu cunature from the apex: Apd, apical aperture diameter; Fiss: fissure length. KW: Kruskal-WaUis 
test): ns: not significant; ''p<0.(){)l. The horizontal lines at Dunn's test represents absence of statistical differences. 

H }iu-ri(li(>ii(ilis (n = ]2) 
incan(±SD) niin-max 

//- antipodes (n=S) 

H. subteifissus (n=23) 

mean (±SD) 

niin-max mean (±SD) 



Dunn's test 




0..5 (0.4) 
0.1 (0.05) 

2.1 (0.5) 



8.9 (0.8) 
1.0 (0.07) 
3.5 (0.6) 
0.2 (0.00) 

1.4 (0.2) 







5.4 (2.5) 
0.2 (0.05) 




14.98 *■ 


st sb an 

I 1 H 

St an sb 

I 1 l-l 

st sb an 

I 1 

st sb an 

I 1 l-l 

st an sb 

'— ^1 
st sb an 
I 1 l-l 

fissus are more similar, differing significanlK' only in 
maximum diameter (Max) and apical aperture diameter 
(Apd) (Table 1). Hctcroficliisinoidc.s iiiciidioiialis new 
species is also less shiny and the intercostal spaces are 
more densely striated: 12-15 striae (observed on stained 
or coated specimens due to the difficulty to be obsei"ved 
without any treatment) against 8-10 in H. subfeifissiis 
(well observed under lens) (these data was olitained from 
counting 10 specimens of each species). 

Lidixiduals of Hctcroschisiuoidcs rintipodcs new spe- 
cies are significantly smaller than individuals of the other 
two species for both shell length and fissure extension. 
The distance of point of maximum cun'atvu-e from apex 
(Larc) in //. antipodes new species specimens is located 
nearer to apex than in specimens of both H. iiiciidiniKiIis 
new species and H. siibteijissus (Table 1). \n ackhtion, 
H. antipodes specimens are opaque, due to their coarsest 
surface microsculpture. Specimens of this species have 
smooth intercostal spaces without secondaiv ribs and 
striae, and the primary ribs are more apparent. 

The multivariate discriminant function analysis was 
able to distinguish three groups among the specimens we 
examined that correspond to three species (Wilks" 
Lambda = 0.12; Fe-g = 24.27; p < 0.0000). This analysis 
classifies about 97% of the cases correctly (only 1 out of 
43 specimens were incorrectly classified) (Figure 3). The 
model was constructed with 3 parameters (Max, L : Max, 
Larc). The variable L. which was highly con-elated with 
Larc (r = 0.80) and Fiss (r = 0.74), was excluded. The 
Discriminant Functions (DF) based on the raw coeffi- 
cients of canonical variables are shcjwn below: 

DFl = 2.5439Max + 1,0378L : Max - 0.7366Lurc 

DF2 = 0.7249Max + 2.1404L : Max - 0.8.328Larc 

The radulae of tlie three species, here illustiated, are 
quite similar and fmther studies would be necessary to 
test the existence ol interspecific differences, as it is tlie 

case in most Scaphopotla. Finally, the findings of Het- 
eroschisnioides in the Solomon Islands sho\\' the genus to 
have worldwde distributetl instead of being confined to 
the eastern Atlantic Ocean. However, better sampling of 
areas of the Indian Ocean (one of the less known tropical 
areas for scaphopod diversity) could eventually confirm 
whether the genus is also represented in that ocean. 


The authors want acknowledge the support and help re- 
cei\ed from Pliilippe Bouchet, Virginie lieros, and Phil- 
ippe Maestrati (MNHN), Ellen Strong (USNM), Kathie 
Way (BMNH), and Anders Waren (Swedish Museinn of 
Natural Histow, Stockholm). Ronald Shimek is also 
g]'atefnlly acknowledged for his constructix'e connnents 
on the manuscript. 


ChistikuN % S. D. 1982 The modern Entahnidue iScapliopoda, 
Gadilida), 1. Subfamily Heterosehismoidinae - 1 [in Rus- 
sian]. Zoologicheskii Zhurnal 61 (5): 671-682. 

Dall, W. H. 1889. Reports on the results of dredging, under the 
supenision ol Alexander Agassiz, in the Gull ol Mexico 
(1877-78) and in the CarilDbean Sea (1879-80), b\- the 
U.S. Coast Suivey Steamer "Hlake", Lieut. -Commander 
CD. Sigsbee, U.S.N.. and Commander J.R. Bartlett, 
U.S.N., commanding. XXIX. Report on the Mollusca. Part 
2, Gastropoda and Scaphopoda. Bulletin of die Museum 
of Comparati\-e Zoolog)' at Hai^ard College IS: 1—192, pis. 

Jeffreys, ]. G. 1877. New and peculiar Mollusca ol the order 
Solenoconcliia procined in the "Valorous" expedition. An- 
nals and Magazine of Natural Ilistoiy (4) 19: 153-15S. 

Jeffreys, J. G. 1883. On the VIollusca procured during the 
"Lighting" and rorciipiiic" expeditions, 1868-70. Pro- 
ceedings ol llic Zoological Society oi London 1882: 656- 
687, pf. 49-50. 

\'. Scarabino and C. H. S. Caetano, 2008 

Page r 

1 k'uderson, ]. B. 1920, A nionograpli of the East Anierican 
scaphopocl rnolhisks. United States National Mnsenni Bnl- 
letin 111: 1-177. 20 pis. 

Locard, A. 1.S9.S. Mollusijues Testaces, In: Expeditions Scien- 
titiqnes du Tnivdillciir et du Talisiiuni pendent les annees 
1SS0-1S.S3. Masson, Paris, 515 pp., 1,S pis. 

Niekles, M. 1979. Scaphopodes de rOuest-Africain (MoUusca, 
Scaphopoda). Bulletin du Museum National d'Histoire 
Naturelle, Section A. Zoologie, Biologie et ecologie Ani- 
males (series 4) 1(1): 41-77. 1 pi, 

Pilsbiy, H. A. and B. Sliaip. 1S97-1S9S, Scaphopoda, 1-280, in: 
Manual of Concholog\ . 17: 1-280. Conehological Section, 
.Acadeni\- of Natural Sciences. Philadi'lphia. [1897: 1-144, 
pis. 1-26; 1898: i-xx.\ii + 145-280, pis. 27-37], 

Roniesburg. H. C, I9S4. Cluster .Analysis for Researchers. Life- 
time Learning Publications, Belmont, 334 pp. 

Scarabino, V. 1995. Scaphopoda of the tropical Pacific and 
Indian Oceans, with descriptions of 3 new genera and 42 
new species. In: Bouchet, P. (ed.), Resultats des Cam- 
pagnes MUSORSTOM, \ol.l4. Memoires du Museum na- 
tional d'Histoire naturelle 167: 189-379. 

Shimek, R. L. 1989. Shells moiphometrics and svstematics: a 
re\iew of the slender, shallow-water Cadulus of tlie Noitli- 
easteni Pacific (Scaphopoda: Gadilida). The Veliger, 32, 

Steiner, G. 1997. Scaphopoda from tlie Spanish coasts. Iberus 
15: 95-111. 

Steiner, G. 1998. Ph\"logen\- of Scaphopoda (Mollusca) in the 
light of new anatomical data on the GadiUnidae and some 

problematica, and :i icpl\- to Revniolds. Zoologica Scripta 
27: 7.3-82. 

Stehier, G. 1999. A new genus and species of the family An- 
ulidentaliidae (Scaphopoda: Dentaliida) and its systematic 
implications, |ournal ol Molluscan Studies 65: 151-161. 

Steiner, G. and K. Linse. 2000, Ssstematics and distribution of 
the Scaphopoda (Mollusca) in the Beagle Channel (Chile). 
Mitteilnngen aus dem Hamburgischen Zoologischen Mu- 
seum und Institut 97: 13-30. 

Steiner, G. and H. Dreyer. 2003. Molecular phylogeny of 
Scaphopoda (Mollusca) inferred from 18S rDNA se- 
quences: support for a Scaphopoda-Cephalopoda clade. 
Zoologica Scripta 32: 343-356. 

Steiner, G. and A. R, Kabat. 2001. Catalogue of supraspecific 
taxa of Scaphopoda (Mollusca). Zoosx-stema 23: 433—460 

Steiner, G. and A. R. Kabat. 2004. Catalogue ot species-group 
names of Recent and lossil Scaphopoda (Mollusca). Zoo- 
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Jeffreys, widi the location of the t\pe material. Coneho- 
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50 pis. 

THE NAUTILUS 122(3):178-181, 200S 

Page 178 

Parasitism of Monogamus minibiilla (Olsson and McGinty, 1958 
(Gastropoda: Eulimidae) on the red sea-urchin Echinometra 
hicimter (Linnaeus, 1758) (Echinodermata: Echinometridae) on 
the Caribbean coast of Mexico 

Norma Emilia Gonzalez-Vallejo 

Depto. Ecologi'a Acuatiea 

El Colegio de la Frontera Sur 

Apdo. Postal 424, 77014 

Chetumal, Quintana Roo, MEXICO 


Eulimids are gastropod mollusks parasites of echinoderms. In- 
tertidal red sea-urchins, Echinometra liiciinter, collected along 
die Mexican Caribbean shores were examined for tlie presence 
of those ectoparasitic gastropods. The analysis of 206 sea- 
urchins resulted in 56 having eulimids on them (27% preva- 
lence), mostly living in pairs, with about 7 eulimid pairs per 
sea-urchin. The eulimid gastropods found are Monogamus 
minibuUa (Olsson and MacGinty, 1958), described from 
Panama. Its host was unknown. This article consists of the first 
record of this host-parasite relationship. A short description of 
the shell stiiicture and some details of the biological relation- 
ship are given. The females are much larger than the males, 
and the latter are usually attached to die female bodies. Most 
female eulimids live inside the hypertrophied foot, which this 
allows for partial enveloping of the entire body. A few females, 
however, were found living fixed directly on the echinoderm 
integument. The position of the gastropod on the sea-urchin is 
fixed via boring through the skeleton. Anchoring is achieved via 
die proboscis. Some females had egg capsules (0.9-1.5 mm) 
fixed to their bodies; each is a spherical structure attached to 
the body by a short stalk. Egg capsules contained 14-15 em- 
bryos in different stages of development. The only previously 
published account of this type of relationship involves a similar 
species living on Echinometra mathaei (Blaimalle, 1825) from 
the Red Sea. 

Additional Keijwords: Rosenia, Ecliinometra mathaei, Monoga- 
mus interspinea 


The red sea-urchin Echinometra lucunter (Linnaeus, 
1758) is frequently found on shallow water rocky- 
bottoms along the Mexican Caribbean shores, and occa- 
sionally in sea-grass beds. In that region, some specimens 
living in the reef lagoons host one or more tiny whitish 
eulimid gastropods firmly attached to their bodies. 
Ecological associations including eulimids gastropods 

have been recognized (Waren, 1983). The original de- 
scription of the eulimid Rosenia minibuUa Olsson and 
McGinty, 1958, was made from an empty shell found on 
the beach of Isla Bocas, Panama; however at that time, its 
host was unknown. 

Liitzen (1976) studied the sea urchins Echinometra 
mathaei (Blainville, 1825) from the Red Sea and the 
Western Pacific Ocean and found eulimid parasites on 
them. He named a new genus as Monooanms because he 
found these eulimids always living in pairs; he included 
two different species, one (M. entopocha Liitzen, 1976) 
living inside the hypertrophied tube feet of E. matthaei 
from the Red Sea, and another one (M. interspinea 
Liitzen, 1976) living on the integument of E. matthaei 
from several localities in the Western Pacific Ocean. 

Waren (1983) established Ro.senia Schepman, 1914, as 
a junior synonym of Pelseneeria Koehler and Vaney, 
1908. However, the transfer oi Rosenia minibuUa to Mo- 
nogamus was informally proposed by Waren and 
Moolenbeek (1989), perhaps because both share internal 
growth lines in the body whorl. In addition, the former 
author had found individuals of the species living on E. 

The faunal associations of the red sea-urchin were 
studied in the southern Caribbean Sea by Schoppe and 
Werding (1996), but these authors did not find eulimids 
associated with the red sea-urchin in the study localities. 
In this contribution, my objectives are to report the para- 
sitism of die eulimid Monogamus minibuUa (Olsson and 
McGinty, 1958), on the red sea-urchin Echinometra hi- 
cunier (Linnaeus, 1758), found along the Caribbean 
coast of Mexico, and to describe some aspects of the 
relationship between the two species. 


The material was collected along several places in the 
Caribbean coast of Quintana Roo, Mexico: Isla Contoy 

N. E. GonzalezA'allejo, 2008 

Page 179 

(2r30'S.4" N, S6°47'45.3" W. 2S/II/01); Plava Paraiso. 
Cozumel (20°35'21.5" N, 86°43'44" W, 24/111/01); Playa 
Ana and Jose (19°54'22.4" N, 87°26'14.5" W, 2 km S 
from the junction to Boca Paila, 18/11/01); Majagual 
(1S°43'2S"'N, 87°42'05" W. 22/111/00); Punta Herradura 
(1S°32'23" N, 87°44'32" W, 27/VI/OO); and Buenavista 
(1S°30'42" N, 87°45'30" W, 15AV99). 

Sea-urchins were counted and those uath euhmids 
fixed, sorted and kept indixidualK', and identified accord- 
ing to Hendler et al. (1995). The number ol euhmids per 
sea-urchin was counted to determine the prevalence: the 
relationship heh\een the number of infested hosts (sea- 
urchinsl and the total number of hosts; and the intensity: 
number of parasites (eulimids) present in infected host 
i,Margohs et al. 1982). Voucher specimens were depos- 
ited in the Swedish Museum of Natural History, Stock- 
holm. Sweden, SMNH 27858, and in the Reference Col- 
lection of El Colegio de la Frontera Sur (ECOSUR), 
Chetumal, Mexico). 


Monooanuis iniiiihiilla (Olsson and McGintv, 1958) 

Description: Mean shell length: 1.85 mm (range 1.5- 
2.5), uitlth 1.15 mm (range 1.0-1.5). Females are easily 
distinguished due to their bodv-size, being twice as large 
as males. The shell is tiny, spherical, fragile, and trans- 
parent. The spire is low, the nuclear apex is long and 
erect like a stump (mucro). The shell color is white with 
gloss\- texture, with a very fine suture line. There are two 
post-nuclear whorls; the first one is small and convex, 
with a small shoulder close to the suture, which is better 
seen on the aperture side. The bodv whorl is large, and 
inflated, polished, with some delicate growth lines. The 
aperture is wide, the outer lip thin, expanded along its 
median part. There is no operculum. When the animal is 
alive, the mantle and visceral mass are deep-red. There 
are two distinct black eves. Tentacles were not obsei"ved. 

Parasitism: From a total ot 206 sea-iu'chins, 56 were 
infested with eulimids. The prevalence was of 0.27 (pres- 
ence of the parasite in 27% of the specimens). The in- 
tensity w^as of 7.03: there was an average of 7.03 eulimid 
pairs per sea-urchin (including only those that were in- 
fested by at least one eulimid [Table 1]). 

Attachment Locations of Parasites on Hosts: Many 
parasites ha\'e a preference for the oral side of the host. 

although some were located on the dorsal side 
(periproct); in only two cases parasites were found on the 
peristomial membrane (buccal tube foot). Most eulimids 
invade the tube feet, becoming inserted in its integu- 
ment. This invasion can produce a hypertrophy of that 
tube foot. Often, the deformation is such that when the 
eulimid grows into adulthood, the tube feet are modified 
beyond recognition. The eidimids, in particular juveniles 
with a single teleoconch whorl, use the skin of the tube 
as a protective hood; later, as the animal grows, its owm 
mantle (the pseudopallium) covers the fragile shell, ex- 
cept for the apical part, which is left uncovered (Fig- 
ure 1). 

Another attachement site is the interambulacral zone. 
Eulimids that are not covered by the pseudopallium are 
attached by the snout (plus phaiyngeal anchor). Few oth- 
ers may he found adhering to the integument at the base 
of spines, where according to stage of shell growth, it 
may be collapsed, broken, and later disappear, leaving an 
open space where the "euhmid-scar" becomes evident. 

Eulimids may bore the sea-urchin test: this mav be 
seen clearly when detached specimens were placed on a 
host. They may also enlarge the poi-e associated with the 
foot tube: a typical feeding pattern of ectoparasitic eu- 
limids involves insertion of the proboscis through the 
tube pore followed by suction of coelomic contents of the 

The gastropods were found mosth in pairs, but, less 
frequently, two or three males were found associated 
with a single larger female. These pairs look like irregu- 
larly shaped protrusions of the body wall, each with one 
or two circular openings through which the eulimid shell 
tips may project. Egg capsules were obsei-ved packed in 
globular sacks (3-5), attached for a short stalk to the body 
of the female, and partiallv or completelv covered by the 
pseudopallium. Each capsule (size 0.09-1.5 mm in 
length) contained 14-15 embryos. Egg capsules con- 
tained all stages from early egg stage to small jiueniles 
ready to hatch. 


Olsson and McGintv (1958) described Monogamus 
minilniUa based (jn an empty shell. In the present study 
details of the shell morphology are presented based on 
several alive specimens and those fixed on the sea- 
urchins. Waren in Waren and Vloolenbeek (1989) briefly 
pointed out the presence of AL ininihuUa associated to 

Table 1. Intensit\' and prevalence oi Monogamns iniitihiilln on Eiliiiutinctni Imiiittcr on the Carililiean coast of Mexico. 


St'u in"cliiiis 





limid females 



Isla ContoN 






P. Paraiso 






P. Ana V Jose 












P. Herradura 












Page 180 

THE NAUTILUS, \'ol. 122. \o. 3 

Figures 1-4. Monogamus mlmbuUa. 1. On Echiiioinetra liiciintcr (host test = 16.54 mm diameter x S.72 nun height); 2. 7/! sHii. 
3. Male (left. 1.5 mm) and female (right, 2.5 mm). 4. Female, anterior and lateral Wews (1.8 mm height ■ 1.4 unn width.) 

Echinometra hiciiiitrr. However, they only recorded the 
dimensions of the shell ( 1.5 mm of longitude and a lai-val 
shell of 1.0 mm), because their contrihution was focused 
mainlv in the relationship ol the eulimid Trochostilifcr 
eiicidaricola (W'aren and Moolcnheek, 1989) associated 
with the pencil sea-urchin Euckhins trihiilnidcs (La- 
marck, 1816). This stud\' constitutes the first report of 
the relationship ol Moiiogaiiius miiiihiilhi with Eclii- 
nometra lucunter for the Me.xiean Caribbean region. 

Liitzen (1976) reported the presence of Monogainus 
entopodia in the tube feet of the Red Sea sea-urchin. In 
the Mexican Caribbean, M. mhi'dndla infests the tube 
feet, but was also obsei'ved attached to the epidermis 
host, using its snout and tissues of the host but without 
being covered or protected with the pseudopallium. The 
two different patterns were rarely obsewed on the same 
sea-urchin. Liitzen (1976) had (omid the same patterns 
but on sea urchins collected in dillcrent localities and 
regarded them as belonging to two different species. 
Further, Ecliinininlr/i Iroiii llu' Indian and western Pa- 

cific oceans have been found to belong to three different 
moi"phospecies that might be reproductiveh' isolated 
from each otlu'i- as well (Arakaki and Uehara, 1999). 
Thus, it seems that C'aribbean sea-urchins ha\e not di- 
verged as much as those present in the Indo-Pacifie 

Few studies liave detailed the prevalence and intensity 
of eulimids parasites, Salazar and Reyes (1998) studied 
the relationship of TlujCd rallista Beriy, 1959, and the 
starfish Phatnri/i iiiiifascialis (Gray, 1840), finding that 
the association was more frequent in two sites of the Gulf 
of California where hiunan influence was more accentu- 
ated. In this case, only one of tlic localities (Playa Ana 
and Jose) is liea\il\ inniicnced li\ drxilopment, and is a 
protected, semi-enclosed embaymeut, with many resi- 
dences and tourists. That locality presented two sea- 
urcliiTis that had unusualK' high infestations of about 100 
eulimids each. Most ol the sea-urcliiiis were adjusted 
close to the average ol alicml 7 ciiliniid paiis per sea- 

N. E. Gonziilez-\'iillejo. 200S 

Page 181 

\\'aren (19S3) pointed out that mam' tropical eulimids 
tliat attach their egg capsules to their shells to or to their 
hosts, ha\e almost inxariablv been found with spawai, an 
indication that spa\\iiing might be continuous. On the 
other hand, Liitzen (1976) suggested for M. entopodin 
that the presence of different de\'elopmental stages 
found simultaneousK' in samples of egg capsules indi- 
cates continual production of eggs, which is similar to 
what happens in A/. minibuUa. 

A thorough studv of the association is requnetl to elu- 
cidate the mechanisms of feeding, including histology. 
There is need to clarif\- \\'hether the proboscis sucks of 
part of the hquid fraction of the hosts coelom or, as 
Liitzen (1976) pointed out for M. cntopodia. the eulimid 
feeds exclusi\el\- of the area adjacent to attachment on 
die sea-urchins. 

To determine the specificity of this association it is 
necessan" to do experiments and obsenations in the col- 
lection place, to determine whether, as Waren (1983) 
pointed out, diere is lack of specificity of the parasite on 
a single species guest. Although I had opportunity' to 
examine some specimens of Echinonwtra liiiclis Agassiz, 
1863, and I did not observe any attached eulimids, it 
could be said that if some preference exists in this case, 
but also diis \\'ould ha\'e to corroborate it. Further, in 
2001 1 had examined about 50 specimens belonging to E. 
htcuntcr from Guana Island, hut none had anv associated 
eulimid parasite. 


Manv dianks to Anders Waren (Swedish Museum of 
Natural Histor\'), who always answered my questions 
about euhmids and is aluws ready to help. To Sergio 
Salazar-X'aUejo and Maria Ana Tovar, who reviewed pre- 
\ious drafts of the manuscript and to colleagues at 
ECOSUR for collecting the sea urchins for the present 
stud\'. The careful reviews by Anders Waren and an 

anonymous referee enhanced the clarit\' of tliis contri- 
bution. Humberto Bahena helped impro\e the qualit)' of 
the photograpiis. 


Arakaki, Y. and T. Uehara. 1999. Moipliological comparison of 
black Echi no metre individuals among those in tJie Indo- 
West Pacific. Zoological Science 16: 5.51-.558. 

Hendler. G., J. E. Miller, D. L. Pawson, and P. M. Kier. 199.5. 
Sea stars. Sea-urchins and Allies: Echinoderms of Florida 
and the Caribbean. Smithsonian Institution Press, Wash- 
ington, 390 pp. 

Liitzen, J. 1976. On a new genus and two new species of proso- 
branchia (Mollusca) parasitic on the tropical sea urcliin 
Echinometra mathaei. Israel Journal of Zoolog\' 25: 38-51. 

Margolis, L., G. W. Esch, J. C. Holmes, A. M. Kuris, and A, 
Schad. 1982. The use of ecological terms in parasitology 
(report of an ad hoc Committee of the American Society of 
Parasitoiogists). [ournal Parasitology 68: 131-133. 

Olsson, A. A. and T. L. McGinty. 1958. Recent marine moliusks 
from die Caribbean coast of Panama with tlie description 
of some new genera and species. Bulletins of American 
Paleontolog)- 39(177): 1-58. 

Salazar, A. and H. Reyes. 1998. Parasitismo de Thijca callista 
(Gastropoda: Capulidae) sobre Phataria unifascialis (As- 
teroidea: Ophidiasteridae) en el Golfo de Cahfornia, 
Mexico. Revista de Biologia Tropical 46(3): 1^. 

Schoppe, A. and B. Werding. 1996. The boreholes of the sea 
urchin genus Echinometra (Echinodermata: Echinoidea: 
Ecliinometridae) as a microhabitat in tropical South 
America. Publicazioni della Stazione Zoologica di Napoli 1, 
Marine Ecolog)' 17: 181-186. 

Waren, A. 1983. A generic revision of the famii)' Enlimidae 
(Gastropoda:Prosobranchia) The Journal of MoUuscan 
Studies. Supplement 13, 96 pp. 

Waren, A. and R. Moolenbeek. 1989. A new eulimid gastropod, 
Trochostilifer eucidaricola, parasitic on die pencil urchin 
Eiicidaris trihuloides from the southern Caribbean. Pro- 
ceedings of the Biological Society of Washington 102: 

THE NAUTILUS 122(3): 182, 2008 Pare 182 


Due to an editorial lapse, the article by Simone and Cunlia (2008) in the most recent issue of The Nautilus incorrectly listed "FMNH" 
instead of the correct form "UF" as the collection acronym for the Florida Museum of Natural History. 


Simone, L. R. L. and C. M. Cunha. 2(K)S. Revision of the genus Spinosipella (Bivalvia: Verticordiidae), with descriptions of two new 
species from Brazil. The Nautilus 122: 57-78. 

THE NAUTILUS 122(3):183-1S4, 2()()8 

Page 1S3 


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Volume 122, Number 4 
December 22, 2008 
ISSN 0028-1344 

A quarterly devoted 
to mahicologij. 


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Vahimc 122, Number 4 

December 22, 2008 

CONTENTS ^SSW 0028-1344 

Daniel L. Geiger New species of scissurellids from the Austral Islands, French Pohaiesia, and the 

Indo-Malayan Archipelago (Gastropoda: Vetigastropoda: Scissurellidae, 
Anatoiriidae, Larocheidae) 185 

Sven N. Nielsen EarK' Pleistocene niolliisks ot the Tuhnl Formation, South-Central Chile 201 

Claudio \'alclovinos 

Juliana M. Harding Radula moi-phology in veined rapa whelks, Rapana vcnosa (\'alenciennes, 1S46) 

Stefanie M. Gera (Gastropoda: Mnricidac) from Chesapeake Bay, USA 217 

Roger Mann 

Maria Cristina Dreher Mansur .\ new species ot Sphacriuiii Scopoli, 1777, Ironi southern Brazil (Bi\al\ia: 

Glaus Meier-Brook Spliacriidae) 228 

Cristdan Ituarte 

Maria Eugenia Segade Some remarks on tlie gross anatomy o{ Adelomelon fenissacn 

Guido Pastorino (Donovan, 1824) (Gastropoda: Volutidae) from the coast of Patagonia, 

Argentina 236 

-Alexandre Dias Pinienta A new species and a ii('\\ record ol Muricidae (Gastropoda) ircjm Brazil: genera 

Diogo Ribeiro do Couto Ptenjnotus and Leptotrophon 244 

Paulo Marcio Santos Gosta 

Luiz Ricardo L. Simone A new species oi Chlaunjdocomha Dall, 1SS4. horn southeastern Brazil (Bixalvia: 

Chlamvdoconchidae) 252 

Research Note 

Jeremy S. Tiemann Sinistral Campehma dccmim (Sav, 1817) (Gastropoda: X'ixipaiidae) from the 
Kerin S. Cuniniings Fo.x River, Illinois 259 

Book Reviews 2fil, 263 

Errata 264 

Notice 264 

Author Index 265 

MBLWHOI Library 

JAN 7 2008 


Massachusetts 02543 


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THE NAUTILUS 122(4): 185-200, 2008 

Page 185 

New species of scissurellids from the Austral Islands, French 
PoKiiesia, and the Indo-Malayan Archipelago (Gastropoda: 
\ etigastropoda: Scissurellidae, Anatomidae, Larocheidae) 

Daniel L. Geiger 

Santa Barbara Museum of Natural Histon' 
In\ertebrate ZoologN' 
2559 Puesta del Sol'Road 
Santa Barbara, CA 93105 USA 


Four new species of Scissurellidae, .Vnatomidae. and Larochei- 
dae are described from the Austral Archipelago, French 
PoKnesia and the Indo-M;ila)'an Archipelago: Sinezoiiii 
diinielclreicri new species, S. iviley new species (both Scissur- 
ellidae), Anatoma rnpaensis new species (Anatomidae) and 
Trogloconclw lozoueti new species (Larocheidae). One ol 
tire species is currently only known from the Austral Islands 
[T. lozmic'ti). while the others seem to show a Imiad 
Indo-Mala)an Archipelago to western Pacific distribution. 
Shells of all species and the radulae of S. danieldreieri, 
T. lozoueti, and A.rnpacnsis are illustrated with scanning elec- 
tron micrographs. 

Additional Keywords: MoUusca, Rapa Island, radula 

three genes ( I listone 3, COI, 18S) recovered in some of 
his analyses a clade rniiting all three of these lineages 
(LcpctiHlrihis. Siiiczona, Anatoma). Scissurellidae and 
Anatomidae are characterized by a slit or foramen in 
the shell (lacking in Larocheidae), a rhipidoglossate rad- 
ula, and a lack oi nacre. Approximately 140 species are 
cin"reiitly described with an additional 90 remaining to 
Ik' formalK' recognized (Geiger, 200.3; 2008). To date, no 
species in these families have ever been recorded from 
the Austral Islands. The species described herein are all 
new to science, one witli its range restricted to the 
Austral Islands, and three others known from several 
Ifjcalities in the broad Indo-Pacific. 


The present contribution describes some microgas- 
tropods in the families Scissurellidae, Anatomidae, and 
Larocheidae known from tlie Austral Islands and be- 
yond. The impetus stems from a Frencfi expedition to 
the southeastern-most Austral island, Rapa (see Lozouet 
et al. 2004; 2005 for details); for tlie species found 
at Rapa, additional material from other localities located 
in \arious institutional collections has also been inclu- 
ded. This article is part of a series describing the bio- 
diversit)^ of tliis remote archipelago (e.g., Schwabe and 
Lozouet, 2006). 

Scissurellidae sensu lato comprises lineages of small 
basal marine snails in Vetigastropoda (Geiger et al., 
2008). The phylogenetic position of Anatomidae is cur- 
rently unsettled. Whereas Geiger and Thacker (2005; 
unpubhshed dataj resolved a Lepetodrihis (Lepetodrili- 
dae) + Scismrella, Sinezona, Sukashitrochiis (Scissurel- 
hdae sensu stricto) clade with more distantly related 
Anatoma (Anatomidae), Kano (2008) using the same 


Standard methods for scanning electron microscopy 
(SEM) were employed as detailed in Geiger (2006a, 
b, c) and Geiger et al. (2007). Terminology for shell 
morphology and details on method of whorl count have 
been given elsewhere (Geiger, 2003; Geiger and Sasaki, 
2008). Specimens cited are diy lots; ": complete" indi- 
cates wet-preserved lots with animals. All depth indica- 
tions refer to bottom depth. 

Institutional abbreviations used in the te.vt are: BRC: 
Bret Raines Collection, Kansas City, USA; DLG: Daniel 
L. Geiger Collection, Los Angeles, USA; JTC: Jean 
Trondle Collection, Paris, France; LACM: Los Angeles 
County Museum of Natural Histoiy, Los Angeles, USA; 
MNHN: Museum national d'Histoire naturelle, Paris, 
France; SBMNII: Santa Barbara Mu.seiuii of Natural 
History, Santa Barbara, USA; USNM: United States Na- 
tional Museum of Natural Histoiy, Washington (DC), 
USA; ZMA: Zoological Museum, Amsterdam, The Neth- 

Other abbreviations used in text are: M: Monotyjiy; 
OD: Original designation; SEM: Scanning electron 

Page ISfi 

THE NAUTILUS, Vol. 122, No. 4 


Scissurellidae Gray, 1847 
Sinezona Finlay, 1926 

Type species: Sc]usinoj>e brevis Hedley, 1904 (OD). 

Remarks: The genus has recently been treated by 
Marshall (2002) and Geiger (2003). 

Sinezona clanicldrcieri new species 
(Figures 1-9) 

Description: Shell small (to 0.77 mm), trochilorm. Pro- 
toconch of 1 whorl, with strong axial sculpture not 
reaching apical suture, apertnral varLx not connected to 
smootli embnonic cap, apertural margin straight. Tele- 
oconch I of 0.875 whorls, suture at peripheiy, approxi- 
mately 9-15 strong, raised axial cords; first spii^als after 
0.5 whorls. Teleoconch II of 0.6 whorls, approximately 
four fine spiral cords at onset of selenizone between 
selenizone and deep suture below peripheiy, descending 
noticeably on last 0.25 whorls; shoulder slightly concave, 
approximateh' 1.3-20 axial cords, raised near suture, as 

high as wide towards selenizone; approximately 10 spir- 
als, distinct cords near suture, diminishing to veiy fine 
cords near selenizone; interstices with fine irregular 
growth lamellae. Base with constriction below seleni- 
zone, approximately 15 elevated spiral cords, crossed 
and run over by approximately 16 fine spiral lines. Um- 
bilicus cone-shaped, moderately wide, wall smooth, at 
distinct angle to base. Aperture D-shaped, roof overhan- 
ging. Selenizone above peripheiy, keels quite strong, 
moderately elevated, distinct lunules, elongated foramen 
closed anteriorly. 


Thin, llexible, multispiral, with central 

K,\i)ULA (Figures 8-9): Rachidian tooth triangular cusp 
with approximately live equal sized denticles. Lateral 
teeth 1-3 similar; outer edge of cusp with 4-5 denticles. 
Lateral tootli 4 reduced in size, hook-shaped. Lateral 
tooth 5 enlarged bv broadening, approximately six, large 
denticles on inner edge, 1-2 small ones on outer edge. 
Central denticle ol inner marginal teeth large, appro.xi- 
niatcK' three denticles on inner edge, four on outer 

Figure 1. HoJotype oi Sinczoiui danieldreieri new species. (SBMNFI 83540). Luniu Kect, Kliiilic Ba\, New Britain, Papiiu New 
Guinea, 5.283° S, 150.131° E, 0-5 m. Scale bar, shell = 500 |iin. Scale bar, protoconch = 100 |.nn. 

D. L. Geiwr. 2008 

Paw ] 87 

edge; outer marginal teeth spooii-sliaped, hilaterallx 
S'sinmetricd with approxiiiiateK' a dozen line dentieles 
on eaeli side. Radnlar interloek moderate. 

T\pe Material: Holot\pe: SBMNll 83540: Figure 1. 24 
Parat\pe.s: .3— t m. Little Santa Cruz Island. Mindanao, 
6.884" N, 122.04'^' E (USNM 8124.53. 1: Figure 2). 
1.5 m, Okina\\-a, Okii, Japan, 26.847° N, 128.287° 
E (LACM 77-61, 3: one illustrated Figure 3). 20 m. Off 

Bohol Is, I'annlaean Island, 9.5° N, 123.917° E (AMS 
406342, 1: Figure 4). 

Type Locality: 0-5 m, Lunm Keel, Kinihe Bav, New 
Bdtain, Papua New Guinea, 5.283° S, 150.131° E. Col. 
D. L. Geiger Oet. 2005. 

El)inolog)': Tlie name honors Daniel Dreier, son of 
Douglas and Ilaima Dreier of Santa Barbara, who have 

Figures 2—4. ParatN-pes of Sinezona danicldreieri new species. 2. Little Santa Cruz Island, Mindanao, Philippines, 6.884° N, 
122.04= E, .3-4 m (USNM 8124.53, 1). 3. Okinawa, Oku, Japan, 26.847° N, 128.287° E, 1.5 ni (LACM 77-61, 3). 4. OIT Bohol 
Island, Pamilacan Island, Philippines. 9.5' N, 123.917° E, 20 m (AMS 406342, 1). Scale hars, shell = 50(1 \\m. Scale bars, protoconch 
= 100 nm. 

Page 188 

THE NAUTILUS, Vol. 122, No. 4 

made generous contributions 
Museum of Natural Iliston'. 

to the Santa Barbara 

Distribution: Indo-Malayan Archipelago to southern 
Polynesia; shallow reef en\'ironments. 

Other Material E.xamined: French Polynesi.\: m, 
Rapa Iti Island, Rapa, Austral Islands, 27.622° S, 
144.302° W (MNHN, 1). m, Kotuaie Point, Tupuaki 
Bay, Rapa, Austral Islands, 27.577° S, 144.343° W 
(MNHN, 1). 2 m, Nordi of Puldtarava, Rapa, Austral 

Islands, 27.597° S, 144.227° W (MNHN, 4: two illu- 
strated Figures 5-6). 2-4 m, N of Aturapa Island, Rapa, 
Austral Islands, 27.572° S, 144.350° W (MNHN, 4). 2 m, 
N of Rapa Iti Island, Rapa, Austral Islands, 27.620° S, 
144.303° W (MNHN, 1). 2 m, N of Anatakuri Bay, Rapa, 
Austral Islands, 27.623° S, 144.307° W (MNHN, 15). 
3-24 m, Hiri Bay, Rapa, Austral Islands, 27.622° S, 
144.370° W (MNHN, 4). 5 m, Anarua Bay, Rapa, Austral 
Islands, 27.605° S, 144.378° W (MNHN, 7: one 
illustrated Figures 7, 8-9 [radula]). 6 m, Mei Point, 

Figures 5-7. Sinezona danieldreieri new species. 5-6. Nortli ol I'ukilaiaxa, llapa. Austral Islands, French Polynesia 
U422T W, 2 m (MNHN). 7. Anarua Bay, Rapa, Austral Islands, French Polynesia, 27.605° S, 144.378° VV, 5 in (MNHN 
specimen is shown in Figures 8-9. Scale bar, shell = 500 |im. .Scale bar, protoconch = 100 |.nii. 

, 27.597° S, 
), Hadula of 

D. L. Geiser. 200S 

Page 189 

Figures 8-9. Radula ot Sinczoiui danicldrcicri new species, from specimen shown in Fi<;;nie 7. Anaiua Bav, Rapa, Anstral Islands, 
27.605" S, 144.378° W, 5 m. 8. Lateral teedi 4 (L4) and 5 (L5) plus marginal teeth. 9. Hal " " ' 

bars =10 \\.m. 

'idth of radula with central field. Scale 

Rapa, Austral Islands, 27.637° S, 144.303° W (MNHN, 1; 
MXHX, 1). 6 m, SW of Gotenaonao Point, Rapa, 
.\ustral Islands, 27.645° S, 144.320° VV (MNHN, 3). 
15-20 m, \^avai, Rapa, Austral Islands, 27.590° S, 
144.381° W (MNHN, 1; MNHN, 1). 16-20 m, Mei 
Point, Rapa. Austral Islands, 27.637° S, 144.303° W 
(MNHN, 2). 18 m, Rarapai Island, Rapa, Austral 
Islands, 27.572° S, 144.368° W (MNHN. 1). 27 m, 
Kauira Point, Rapa, Austral Islands, 27.592° S, 144.347° 
W^ (MNHN, 1). 30 m, NW of Tauna Island, Rapa, 
Austral Islands, 27.605° S, 144.303° W (MNHN, 17). 
36 m, around Rukuaga Point, Rapa, Austral Islands, 
27.568° S, 144.368° w"(MNHN, 8). 

Philippines: Canipo Island, Cuyo Islands, Palawan 
Province, 11° N, 120.948° E (USNM 808195, 1). 

Giam: 0.7 m, Ajavan Bav, S. Guam, 13.245° N, 
144.717° E (LACM 77-19, 2).' 

Remarks: The protoconch sculpture shows some in- 
traspecific variability in die material examined. More 
specifically, the strong axial cords are sometimes discon- 
tinuous. This character is not restricted to the present 
species, hence, it does not seem to be of any systematic 
\alue. Those axial cords are also somewhat variable in 
widtli, die tliickenings occasionally seeming to form 
some faint spiral elements. Given the intra- and inter- 
specific variability of protoconch scidpture at this level 
of detail, the presence/absence of faint spiral elements is 
considered irrelevant for systematic purposes. Discrete 
differences can be observed between strong and weak 
axial cords as utilized previouslv in a svstematic context 
(Geiger 2003). 

Sinczona plicata (Hedley, 1899) from the Indo-Pacific 
has an overall wider and larger shell (2.3 mm: Geiger 
and Jansen, 2004b), more pronounced but fewer raised 
axial cords, and a protoconch with spiral lines. Sinezona 
ferriezi (Crosse, 1867) (? = S. hoeniesi Semper, 1865: 
whereabouts of Semper types unknowai. original illu- 
stration lacking detail) with broad Indo-Malayan 

Archipelago distribution, is about twice as large, has a 
smooth protoconch with a distinct apertural varix, and 
>1 teleoconch I whorls. Sinczona globosa Geiger, 2006, 
from New Caledonia and Wallis Island is more rounded 
in overall appearance, lacks the elevated spiral cords, is 
larger (3.1 mm; Geiger, 2006a), and has spiral sculpture 
on the protoconch. Sinezona niacleani Geiger, 2006, 
with broad Indo-Malayan distribution is more globular 
in overall outline, larger (2.3 mm; Geiger, 2006a), has a 
protoconch with flocculent sculpture, lacks the raised 
;L\ial lines and has a wider umbilicus. All the above 
species grow to much larger size and would be imma- 
ture with open slit at the maximum size of S. chiniel- 

Sinezona wilci/i new species 
(Figures 10-16) 

Description: Shell medium size (up to 1.08 mm) tro- 
chiform turreted. Protoconch slightly sunken in, with 1 
whorl, fine irregular axials, no apertural varix, apertural 
margin straight. Teleoconch I with 0.6-0.7 whorls, su- 
tiu-e at periphery, usually with approximately 14-22 faint 
to distinct fine axial cords, usually without spiral lines 
(occasionally faintest spiral threads only visible by SEM 
[Figure 11]), interstices with irregular thickenings. Tele- 
oconch II witli up to 1.25 whorls, suture below periph- 
ei")'. Shoulder with approximately 32-44 barely 
perceptible to distinct fine commarginal axial cords, 
stronger near sutiu'e becoming less distinct towards sele- 
nizone; approximatelv eight barely perceptible to very 
indistinct spiral cords, concentrated on middle of shoul- 
der; occasional specimens with predominant spiral 
sculpture (Figure 11). Base distinctly constricted below 
selenizone, axial cords of same strength and density as on 
shoulder starting in lower portion of constriction below 
selenizone; approximately 17 fine, irregularly spaced spi- 
ral cords, starting in lower portion of constriction widi 
onset of axial lines. Umbilicus wide, walls straight. 

Page 190 

THE NAUTILUS, Vol. 122, No. 4 

Figure 10. Holotvije of Sinezaiia icilci/i new specie.s. Lumu Reef, Kimbe Bav, New Britain, Papua New Guinea, 5.283° S, 
150.1.31° E, 0-5 m, (SBMNH 8.3541). Scale bar, shell = 500 |,im. Scale bar, protoconch = 100 |.nn. 

smooth, peripheiy at clfstinct angle to base, bordered by 
carina. Aperture subquadratic, D-shaped, roof overhan- 
ging. Selenizone above peripheiy, keels moderately 
strong, quite elevated, lunules distinct at regular inten'al; 
foramen narrow, elongated, margins con\'erging and 
touching, without raphe. Animal unknown. 

Type Material: Holotype: SBMNH 83541 (Figure 10). 
35 Paratypes: 0-5 m, Lumu Reef, Kimbe Bay, New Brit- 
ain, Papua New Guinea, 5.283° S, 150.131° E (DLG 
639, 19: one illustrated in Figure 15). Laisse de plage 
e.xterieure, Mururoa, Tuamotu, French Polynesia, 
21.842° S, 138.895° W (JTC, 2: one illustrated in Figure 
12). 20 m, Herald Pass, W of Ndravuni Lsland, NW 
end of Great Astrolabe Reef, Fiji, 18.767° S, 178.467° E 
(LACM 85-135, 2: one illustrated in Figure 14). 20 m. 
Herald Pass, W of Ndravuni Island, NW end of Great 
Astrolabe Reef, Fiji, 18.767° S, 178.467° E (LACM 85- 
135, 2). 10-20 m. Off Francis Island in Beqa Lagoon, 

just off of Beqa Island (south of \'iti LeMi), 18.300° S, 
178.067° E (DLG 279, 10: one illustrated in Figure 16). 

Tjipe Locality: 0-5 m, Lumu Reef, Kimbe Bav, New 
Britain, Papua New Guinea, 5.283° S, 150.131° E. 

EtMiioIogy: The name honors the collector of the first 
known specimens of the species Tony Wiley of River- 
side, California, USA. 

Di.slriliiilioii: Tropical Indo-Malayan Archipelago; 
shallow shelf 

Other Material Examined: Fkench Polynesia: 36 m. 
Around Rukuaga Bnv, Rapa, Austral Islands, 27.568° S, 
144.368° W (MNHN, 20: Fi,gin-es II, 13). 2-4 m, N of 
Aturapa Island, Rapa. Austral Islands, 27.572° S, 144.350° 
W (MNHN, 1). 5 m, Anama Bay, Rapa, Austral Islands, 
27.605° S, 144.378° W (MNHN, 4). 45 m, Haurei Bay, 
Rapa, Austral Islands, 27.613° S, 144.305° W (MNHN, 1). 

D. L. Geieer. 2008 

Page 191 

Figures 11-13. Sincznna wilei/i new species. 11-12. Around Rukuaga Baw Rapa, Austral Islands. French PolvTiesia. 27.568° S, 
144.368= W. 36 rn (MNHN, 20). 1.3. Laisse de plage exterieure, Muraroa.' Tnamotu. French Polynesia, 21.842° S. 138.895° W 
:Parat\pe: JTC, 2). Scale bars, shell = 500 ^m. Scale bars, protoconeh = 100 |jm. 

New C-;ledoni.^: 25-30 m, Santal Bay. Lifon, 20.822° 
S, 167.173" E (MNHX, 1). .55-57 m, Koumac Sector. 
20.672° S, 164.195° E (MNH\. 1). 5-25 m, Toiiho Sec- 
tor. 20.878= S. 165..325= E (MNHX. 1). 8 m, Tonho 
Sector. 20.742° S, 165.265° E (MNHN, 1). 

Fiji: 10-20 m. Off Francis Island in Beqa Lagoon, just 
off of Beqa Island (south of \'iti Levu), 18.300° S, 
178.067= E (DLG 279, 10). 

Papua New Guinea: 70 ni, off South Enia Reef Kimbe 
Bay, New Britain, 5.202° S, 150.152° E (DLG 768, 3). 

Remarks: The organization of tlie foramen in this 
species seems to be intermediaiy bet\veen the tjpical 
open slit in Scissurella and tlie closed foramen in Sine- 
zona. Altliough the margins of tlie slit are converging 
and fused anteriorl); a raphe is not evident. The facts 

Paee 192 

THE NAUTILUS. \'oI. 122, No, 4 

that there is a marked downward deflection oi the last 
quarter whorl and tliat the roof of the peristome 
attaches belo^^' the peripheiy of the pi'eNdous whorl dem- 
onstrate that tliose specimens are fully grown and sliare 
more similarities witli Sinezona tlian with SciisiircUa 
with an open slit. Accordingly .S. ici!ci/i is placed in 

The species seems to have variable strengths of sculp- 
ture. Most specimens have predominant axial sculpture 
(Figures 10, 12, 14-15), while in occasional specimens 
(Figure 11) the axial sculpture is barely perceptible and 
the spiral structures, particularly on the adumbilical part 
of base, become most prominent. As there are interme- 
diate specimens (Figure 13) with distinct adumbilical 

Figures 14- K>. Siuciuki u iliij, i\r\\ spicirs, paiuUpr.s, 14. Heiakl, \\ (il Ndravuni Island, NWend ul liical A.stidlalH' Hcrl, 
Fiji, 18.767° S, 178.467° E, 20 m (LACM 85-135, 2). 15. Lumu Reef, Kimlie Bay, New [Britain. Papua New Guinea, 5.283° S, 
150.131° E, 0-5 m (DLG 639, 19). 16. Off Francis Island in Beqa Lagoon, jnst olTof Beqa Island (sonth ofViti Levn). Fiji, 18.3° S, 
178.067" E, 10-20 m (DLG 279, 10). Scale bars, shell = 500 ^ini. Scale bars, protoconch = 100 |.ini. 

D. L. Geiger. 2008 

Page 193 

spiral lines and i"elati\'e!\- weak ;L\ial lines, coniliiiu'd 
widi identic;il condition of the protoconc-h and \ei\ 
small size, tiiese scnlptm'al differences arc coiisidcrcd 
to constitute intraspecitic \ariabilit\. 

There are no similar species in the indo-Pat'llic. Siiic- 
zona feniczi (Crosse, 1867) with hroail Indo-Malayan 
Archipelago distribution is most similar, but is about 
h\ice the size of S. wileiji. has a smooth protoconch \wth 
distinct apertin'al \arix, a teleoconch 1 with more than 
whorl, is more rounded o\'er;rll, and a distinct raphe 
anterior to the closed foramen. Siiiczona garciai Geiger, 
2006, from the Caribbean, shares the protoconch sculp- 
ture, the absence of an apertural varix on the pioto- 
conch, the relati\el\' short teleoconch I with 0.6 whorls, 
and die narrow umbilicus. However, in addition to its 
occurrence in a separate ocean, S. garciai has a more 
inflated shell with an oval aperture, which connects to 
the previous whorl bareK' below the periphen', has onlv 
about 0.6 teleoconch II whorls, and bears a distinct 
raphe anterior to the ioramen. 

Anatoniidae McLean, 1989 
Aiiafoina Woodward, 18.59 

T>pe Species: .SV'/.s.svnv//*/ ciispiitd Fleming, 1828 (M). 

Remarks: The genus has recently been treated by 
.\hirshall (2002), Geiger (2003), and'Ceiger and Jansen 


Anatoina rapacitsis new species 
(Figures 17-23) 

Description: .Slu'll ol medium size (to 1.82 mm), tro- 
cliilonii biconical. Protoconch with 0.75 whorls, flocculent 
scnjptui'e, no apertiu'al varbc, apeiiural margin slightly si- 
nusoid. Teleoconch I with 0.3-0.4 whorls, approximate- 
ly 9-12 tLxial cords, interstices with line flocculent 
sculpture, occasionall)' (holot)pe) flocculent sculpture 
somewhat concentrated in position of selenizone. Teleo- 
conch II with up to 2.25 whorls, sutm-e immediately below 
selenizone in early growdi, offset by width of selenizone 

Figure 17. Holot\pe of Anatoina rapaensis new species. St. 43, Haurei Bay. Rapa, Austral Islands, French PoKmesia. 27.61.3° S, 
144.30.5^ \\". 4.5 ni. ('MNHX 20S23j. Scale bar shell = 1 mm. Scale bar protoconch = 100 |.un. 

Page 194 

THE NAUTILUS. Vol. 122. No. 4 

Figures 18-20. Aiiatoiiui rapaeiisis new specie.s, parahpes (MNHN). 18-19. St. 4.3. Haiiiei Baw Rapa. .\ustral Lslands. French 
Polviiesia. 27.613" S, 144.305" W, 45 m (MNHN 20S24). 20. St. 48. around Kukuasja Point, Rapa. Austral Lslands. French PoKiiesia. 
27.568" .S. 144.368" W, 36 m (MNHN, 30). Scale bars, shell = 1 mm. Scale bars, protoconch = 100 ^im. 

in fully grown specimens. Shoulder convex, approximately 
5.'3-66 (n = 4) axial cords on first teleoconch II whorl, same 
density on remaining whorls; first fine spiral cords after 
0.125 whorls, approximately 7-10 after one teleoconch II 
whorl, appro.ximately 15-22 at apertural margin (jf fully 
grown specimen, becoming less distinct and more un- 
evenly spaced towards apertural margin; intersections of 

a\i;J and spiral cords loiining minute points. Base contin- 
uously sloping with narrow umbilicus; axial cords of same 
density and strength as on shoulder, approximately 19 
spiral cords, fine spiral cords below selenizone tuniing 
into low steps from mid-base onwards; intersection of 
spiral and a,xial cords with line points. .Aperture o\oid 
D-sh:iped, lool oNcrJKiiigiug, l);is;il ;idunil)ilK';Ll portion 

D. L. Ceiger, 2008 

Pas^e 195 

ilared. Selenizone at periplien*. keel iiKKlerateK' strong, 
iiKxlerateK' elexated, slit open, margins converging 
towarils apeitural margin. 

Operculum (Figure 23): As large as apeilure, thin, I'or- 
neons, nucleus central, ninltispiral. 

Radul.\ (Figures 21-22): Rachidiaii tootli trajKvoid. 
central denticle with 3—1 denticles on each side. Lati'ral 
teeth 1-3 similar, dexelopment ol cnsp reduced periph- 
eralK. terminal denticle largest, 4-2 denticles on outer 
edge. Lateral tooth 4 reduced in size, hook-shaped, with 
one minute point on each side. Lateral tooth 5 enlarged 
wdth four strong denticles on inner margin. Inner mar- 
gin;J teeth elongated, terminal denticle largest, 3-4 
snurller denticles on inner margin, three larger denticles 
on outer margin. Outer marginal teetli with cup-shaped 
cusp with mam' small denticles. 

Type Locality: 45 m, RAPA St. 43. Haurei Bay, Rapa, 

Austral Islands, French PoKi 

27.613'' S, 144.305° W. 

T\pe Material: 

60 Parat\pcs: 45 
-\ustral Islands, F 
^^" (MNHN 20S24 
36 m. RAPA St. 
.\ustral Islands, 2' 
one illustrated in 
St. CP 1159, Eiao 
1: Figures 21-23 I 

Holot\pe: MNHN 20823 (Figin-e 17). 
m, RAPA St. 43, Haurei Bay, Rapa, 
rench PoKiiesia, 27.613'' S, 144.305" 
, 29; t\vo illustrated in Figures 18-19). 
48, Around Rukuaga Point, Rapa, 
568° S, 144.368° W (MNHN, 30: 
Figure 20). 145 m, MUSORSTOM 9 
Ishnid, 7.972° S, 140.728° E (MNHN, 
radula, operculum]). 

Et>iii()log}': The species is named alter its t)pe locali- 
f\, Rap;i Island, ["rench PoKiicsia. 

Di.stribution: Tropical Indo-Malayan Archipelago and 
W'csteiii Pacilic; sliallow- shelf to upper slope. 

Other Material Examined: Fkengh Polynesm: 10-15 m. 
Pake Ba\-, Rapa, Austr;xl Islands, 27.617° S, 144.310° 
W (MNHN, 2). 30 m, NWof Tauna Island, Rapa, Austral 
Islands, 27.605° S, 144.303° W (MNHN, 50). 33 m, 
Haurei Bay, Rapa, Austral Islands, 27.612° S, 144.318° W 
(MNHN, '10). 52-57 m, SE of Tauna Island, Rapa, 
Austral Islands, 27.608° S, 144.295° W (MNHN, 50; 
MNHN, 20). 100 m. North of R;iivavae, Austral Islands, 
23.828° S, 147.693° W (MNHN, 2). 

PHiLtPPiNEs: 92-97 m, 12.517° N, 120.650° E 
(MNHN, 8). 

New Caledonia: 105-110 m, Poindimie Sector, 
20.817° S, 165.317° E (MNHN, 9). 250-350 m, south- 
ern New Caledonia, 22.500° S, 166.400° E (MNHN, 1). 
250-350 m, southern New Caledonia, 22.500° S, 
166.400° E (MNHN, 1). 495 m, southern New Caledo- 
nia, 22.367° S, 166.233° E (MNHN, 3). 600-616 m, 
northern New Caledonia, 18.817° S, 163.250° E 
(MNHN, 3). 

Figures 21-23. Kadiila and operculum oi Anatoma rapaensis new species. Eiao Island, Marquesas Islands, 7.972° S, 140.728° E, 
145 ni I MXnX, 1 '. 21-22. Radula. 23. riperculum. Scale bar, radula Figure 21 = 50 \xm. Scale bar, radula Figure 22 = 20 mm. Scale 
bar, opeculum = 500 |im. 

Page 196 

THE NAUTILUS, Vol. 122, No. 4 

M.\RQUESAS LsLWD.s: 145 m, Eiao Island, 7.972° S, 
140.728° E (MNHN, 1: paratype). 200-220 m, Ua Huka 
Island, 8.900° S, 139.633° W (MNHN, 6). 352-358 m, 
Hiva Oa Island, 9.850° S, 139.150° W (MNHN, 1). 

Fiji: 149-168 m, S of Viti Levu, 18.207° S, 178.552° E 
(MNHN, 1). 260-305 m, S of Viti Levu, 18.308° S, 
178.097° E (MNHN, 6). 275-430 m, S. of Viti Le\ai, 
18.297° S, 177.907° E (MNHN, 1). 441-143 m, S. of Viti 
Levu, 18.320° S, 177.862° E (MNHN. 1). 

Remarks. Anatoiiui rapacusis i.s characterized In tlie 
rather flattened o\erall shape, the short teleoconch I 
with less than 0.5 wliorls, and the line reticular sculpture 
that is axial-dominated on the iirst half teleoccMich II 
whorl, subsequently becoming spiral-dominated. The 
most similar species is Anatoma (sensu lato) cxqiiisita 
Schepman, 1908, from die Indo-Malayan Archipelago; 
the comparison is based on SEM imaging of the holo- 
type (ZMA 3.08.101; Figure 24). The" shell is slightly 
taller, the spiral and axial lines are denser, forming more 
prominent points at their intersection, and the a.xial lines 
are slightly stronger than the spiral lines even in larger 

specimens (specimens 1.8-^3.5 nun examined); protoconch 
and teleoconch I are eroded beyond recognition. Anatoma 
paucispiralia Bandel, 1998, from Satonda, Indonesia, has a 
smooth protoconch, a shorter teleoconch I (0.125 vs. 0.3- 
0.4 whorls), and a sHghtly undulating shoulder profile. AH 
odier Indo- Pacific species are eidier more turreted or have 
noticeably different sculpture on shoulder and base. The 
only other docimiented case of a species that shows a 
change of sculpture on the teleoconch II is Anatoma jane- 
tae Geiger, 2006, knowai from 2,500 m off the west coast of 
North -America (Geiger, 2006c). 

Larocheidae Fleming, 1927 
Trogloconcha Kase and Kano, 2002 

Type Species: Trogloconcha ohasliii Kase and Kano, 
2002 (OD). 

Remarks: The genus was recently treated by Geiger 


Trogloconcha lozoneti new species 
(Figures 25-30) 

Figure 24. lliildlype oi' Aiialoiiia cMiidsita .Scliepmiin. 1908 (ZMA 3.08.101). Silioga Station 95, 5 43.5' N, lt9°40' E, 522 m 
[between Sal);ili, .Malaysia, and southeastern Tawitawi Group, Pliilippines]. Scale bar, shell = 1 mm. Scale bar, protoconch = 100 |im. 

D. L. Geieer. 2()()S 

Page 197 

Figures 25-27. Tw^loconclia lozoueti new species. 25. HoloUpe. Vavai. Rapa, Austral IslaiiLls. I'niiul] IwUniMd, 27.590° S, 
144..3Sr \\". 1.5-20 m (MXHX 20825). 26-27. Radula from specimen .shovvii in Figure .30. Hiri Bay, Rapa, Austral Islands, French 
PoK-nesia. 27.622' S. 144.37" W. .'3-24 m (MNHN). Scale bar, shell = 1 mm. Scale bar, protoconch = 100 |jni. Scale bar, radiilae = 10 |im. 

Page 198 

THE NAUTILUS, Vol. 122. No. 4 

Figures 28-30. Trogloconcha lozoueti iif\\ s[)rri(s. l^aial\[>r,s, 2iS-29. \a\ai, Hapa, Austral IslaiaLs. trciuli PoKiR'.sia, i i .590'' S, 
144.:381° \V, 15-20 m (MNHN 20826). 30. Hiri Bay, Rapa, Austral Islands, French Polynesia, 27.622° S, 144.37° W, 3-24 m 
(MNHN. 20). Scale bar. shell 28 = 1 nun. Scale bars, shell 29-30 = 200 |.ini Scale bars, pnituconch = 100 jiiii 

Description: Shell medium size (to 1.17 mm), trochi- 
lorm globular, with rapidly increasing whorls. Proto- 
conch with 0.75 whorls, flocculent sculpture somewhat 
spirally ai"ranged, weak apertural \'ari\, apertural margin 
convex. Teleoconch with up to two wliorls, approximate- 
ly ]5 axial cords on First 0.3-0.5 whorls with no spiral 
sculpture, interstices with fine flocculent sculpture; on- 
set ol spiral cords alter 0.5 wliorls; approximately 

31 spiral cords, Irom suture to uiid-base as fine spiral 
cords, transitioning to low spiral steps in 5-6 adimibilical 
spirals; approximately 70 fine axial cords on last whorl, 
forming distinct points at intersections: o\erall appear- 
ance of shell fine reticulate aud spiky. Suture bordered 
by strong irregularly lamellalc tiiiekening on shoulder. 
Base anomphalus with weak c'alius in umbilical n^gion. 
Aperture oval, roof ovei-hanging. 

D. L. Geis;er, 2()0S 

Page 199 

Operculum: Corneous, round, multispiral, with central 
nucleus, covering onl\' approxiniateK' i i ol api'rture. 

Radula (Figures 26-27): Kachidian tooth triangular, 
cusp with central denticle largest, three denticles on 
each side, arranged in convex cin-\'e. Lateral teeth 1-4 
similar, L-shaped, cusp with apical denticle largest, 3—4 
denticles on outer margin, 1-3 denticles on inner mar- 
gin; later;il tooth 5 enlarged bv half, apical denticle lai'g- 
est, 3-4 denticles on each side. Inner marginal teeth 
witli elongated shaft central denticle largest, 3—4 denti- 
cles on inner margin, 5-6 denticles on outei- margin; 
outer marginal teeth with cup shape cusp, with man\- 
fine denticles on each side. Radular interlock ot central 
field strong. 

T>'pe Material: Holot\-pe: MNHN 20825. 26 Para- 
t\^es: 15-20 m, RAPA' St. 32, Vavai, Rapa, Austral 
Islands, 27.590° S, 144.381° W (MNHN 20826, 6: two 
illustrated in Figure 28-29). 3-24 m, RAPA St. 9, Hiri 
Ba\-, Rapa, Austral Islands, 27.622° S, 144.370° W 
(MNHN, 20: one illustrated in Figure 30). 

T%pe Locality: 1.5-20 m, RAPA St. 32, Vavai, Rapa, 
AustnJ Iskmds, French PoKnesia, 27.590° S, 144.381° W. 

Et\TnoIog\': Named in honor of Pierre Lozouet 
(MNHN) for his accomplishments paiticularly in the field 
of fossil mollusks including ScissurelHdae sensn lato, and 
his work in connection with the Rapa expedition. 

Other Material Examined: French Polynesia: 36 m. 
Around Rukuaga Point, Rapa, Austral Islands, 27.568° S, 
144.368° W (MNHN, 100). 33 m, Hanrei Bay, Rapa, 
Austral Islands, 27.612° S, 144.318° W (MNHN, 6). 52- 
57 m, SE of Tauna Island, Rapa, Austral Islands, 27.608° 
S. 144.295° \V (MNHN, 20). 30 m, NW of Tauna Island, 
Rapa, Austral Islands, 27.605° S, 144.303° W (MNHN, 
3). 8 m. S of Tarakoi Iskmd, Rapa, Austral Islands, 
27.093° S, 144.308° W (MNHN, 1). 52-57 m, SE of 
Tauna Island. Rapa, Austral Islands, 27.608° S, 144.295° 
W'MNHN, 9). 

Remarks: Trogloconclia ohashii from southern Japan 
has fewer spiral and lodal elements that also form elevated 
points and a wide umbilicus. Trogloconcha tesschta Kase 
and Kano, 2002, from the Indo-Malayan Archipelago has 
a smooth protoconch and lacks the fine points at the 
intersection of axials and spirals. Trogloconcha chiistiiidc 
Geiger, 2003, from W'estem Ausb'alia lacks spiral sculp- 
ture and has flattened-flocculent protoconch sculpture. 

The new species is currently only known from the 
Austral Islands. The radula represents the general veti- 
gastropod pattern (rhipidoglossate) and shows the tvpi- 
cal larocheid arrangement ot similar lateral teeth 1—4, 
without reduced, hook-shaped lateral tooth 4; the radula 
confirms the placement of the species in Larochaeidae. 
The lack of a brood pouch seen in Larochaea and 
Larocheopsis places the species in Trogloconcha. 
Juveniles have a proportionally wider appearance 
with the suture approximately at the periplieiv of the 

previous whorls. Fully growai specimens ha\'e a more 
clexated appearance with the suture connecting well 
below the peripheiy of the previous wliorl. 


The new species all belong to recognized genera. They 
are diagnosed by particular character combinations not 
known from any described species. The two Sinezona 
species are among the smaller species in tlie genus, 
while the Anatoma and Trogloconcha species are witliin 
the usual size range for their respective genera. All spe- 
cies show characters of shell and radular morphology 
that are known from other species; no new character 
states were found. The combination of particular fea- 
tures and their paiticnlar strength of development, how- 
ever, are unique for each of the nev\' species. 

The distribution of three species reaches beyond die 
Austral Islands towards the equator {Shwzona wilei/i, S. 
(lanicldrcicri, and Anatoma rapacnsis) . while one spe- 
cies (Trogloconcha lozoucti) is currently onlv known 
from Rapa Island. This pattern suggests a closer faunal 
aflinit)' of tlie Austral archipelago widi the broad Indo- 
Malavan Archipelago, rather than with the temperate 
Southern Ocean. The two more widelv distributed spe- 
cies S. chniicldrieri and S. wilciji are also those that have 
been found in deeper water, confirming the idea that 
deep-water species in general show a wider distribution 
than those restricted to shallow water 


I thank on behalf of P. Lozouet all colleagues who 
participated in "RAPA 2002" and especially the moUusk 
group: R. von Cosel, V. Heros, A. Le Goff , P. Maestrati, 
|.-L., Menou, S. Schiaparelli and J. Trondle. Many 
thanks also to Claude Payri (Universitv- of Papeete, 
Tahiti), who organized the logistics with the French 
Navy and the mayor and local council ot Rapa. Pierre 
Lozouet provided helpful criticism on a draft version of 
the present contribution. Diego Zelaya and an anon- 
ymous reviewer improved tlie manuscript with helpful 
pointers. Total Foundation funded the ex|iedition within 
its "Coral Reef Biodiversity Programme." The SEM 
investigation was supported by NSF MRl-0420706 
grant to Henn' Chaney Michael Caterino, and Daniel 
L. Geiger Visiting curatorships from AMS and MNHN 
to DLG are kindly acknowledged. 


Geiger, D. L. 200.3. Phylogenetic assessment of characters 
proposed for the generic classification of Recent Scissur- 
ellidae (Gastropoda: Vetigastropoda) with a description of 
one new genus and six new species fiom Easter Island 
and Australia. Molluscan Research 2.3: 21-83. 

Geiger, D. L. 2006a. Eight new species of Scissurellidae and 
Anatomidae (Mollusca: Gastropoda: Vetigastropoda) from 

Paee 200 

THE NAUTILUS, Vol. 122, No. 4 

around the world, wath discussion of hvo new senior syno- 
nyins. Zootaxa 1128: 1-33. 

Geiger, D. L. 2006b. Sasakiconcha elegantissima new genus 
and species (Gastropoda: Vetigastropoda: Anatomidae?) 
with disjointly coiled base. The Nautihis 120: 45-51. 

Geiger, D. L. 2006c. A new blind Anatoma species irom the 
bathyal of the northeastern Pacific (Vetigastropoda: Ana- 
tomidae). Molluscan Research 26: 108-122. 

Geiger, D. L. 2008. Monographing micromolluscs: A case 
study on Scissurellidae s.I. (Vetigastropoda). Zoosymposia 
1: 133-145. 

Geiger, D. L. and P. Jansen 2004a. Revision of the Australian 
species of Anatomidae (MoUusca: Gastropoda: Vetigastro- 
poda). Zootaxa 414: 1-35. 

Geiger, D. L. and P. Jansen. 2004b. New species of Australian 
Scissurellidae (Molhisca: Gastropoda: Vetigastropoda) 
widi remarks on Australian and Indo-Malayan species. 
Zootaxa 714: 1-72. 

Geiger, D. L., B. A. Marshall, W. F. Ponder, T. Sasaki, and 
A. Waren. 2007. Techniques ibr collecting, handling. 

and preparing small 

molluscan s 

pecimens. Molluscan 

Research 27: 1-50. 

Geiger, D. L., A. Niitzel, and T Sasaki. 2008. Vetigastropoda. 
In; Phylogeny and Evolution of the Mollusca. W. F. Pon- 
der and D. R, Lindberg (eds). Universit)' o( California 
Press, Berkeley, pp. 297-330. 

Geiger, D. L. and T Sasaki. 2008. Four new species of Anato- 
midae (Mollusca: Vetigastropoda) from the Indian Ocean 

(Reunion, Mayotte) and Australia, witli notes on a novel 
radular type for the family. Zoosymposia 1: 247-264. 

Geiger, D. L. and C. E. Thacker. 200.5. Molecular phylogeny of 
Vetigastropoda reveals non-monophyletic Scissurellidae, 
Trochoidea, and Fissurelloidea. Molluscan Research 25: 

Kano, Y. 2(.)08. Vetigastropod phylogeny and a new concept of 
Seguenzioidea: independent evolution of copulatoiy 
organs in the deep-sea habitats. Zoologica Scripta 37: 

Lozouet, P, R. von Cosel, V. Heros, A. Le Goff, P. Maestrati, 
J.-L. Menou, S. Schiaparelli, and J. Trondle. 2004. 
L'Atelier Rapa 2002 (Polynesie franfaise). Xenophora 
107: 17-28. 

Lozouet, P., R.von Cosel, V. Heros, A. Le Goff, P. Maestrati, 
J.-L. Menou, S. Schiaparelli, and J. Trondle. 2005. Biodi- 
versity gradient in the Pacific: first results of RAPA 2002 
(French Polynesia). In: G. Richard (ed.) Les moUusques 
dans la recherche actuelle. Actes du Illeme Congres 
International des Societes Europeennes de Malacologie 
(La Rochelle 24-27 juin 2003): 93-99. 

Marshall, B. A. 2002. Some Recent scissurellids form die New 
Zealand region, and remarks on some scissm'ellid genus 
group names (Mollusca: Gastropoda). Molluscan Re- 
search 22: 165-181. 

Schwabe, E. and P. Lozouet. 2006. Chitons (Mollusca, Poly- 
placophora) from Rapa, the soutliernmost island of 
Polynesia. Zoosystema 28: 617-633. 

THE NAUTILUS 122^^:201-216, 20(),S 

Pa^e 201 

Early Pleistocene mollusks of the Tubul Formation, 
South-Central Chile 

Sven N. Nielsen^ 

GeoForsehnngsZeiitiTim Potsdam 

Sektion 3.1 


14473 Potsdam GERMANY 


Claudio Valdovinos 

Center ui Enviionniental Sciences, EULA-Cliile 

Universidad de Concepcion' 

Casilla 160-C 

Concepcion, CHILE 


Patagonian Ecosystems Research Center (CIEP) 

Covhai()ue, CHILE 


Earh' Pleistocene mollusks of the Tubul Formation are re- 
xlewed and 23 species are recognized. These are twelve 
bi\al\es, ten gastropods, and one scaphopod. With two excep- 
tions, all of them are known from the Recent. The fauna is 
compared witli Miocene, Pliocene-Pleistocene, and Recent 
faunas ot die Chilean coast. The composition of the Tubul 
fauna resembles the Recent one of the Magellan Region, sug- 
gesting cooler water temperatures for south-central Chile dur- 
ing the earlv Pleistocene than at present. 

Additional Kfijwords: MoUusca, eastern Pacific, climate 


The marine late Pliocene to early Pleistocene of Chile 
comprises localities distributed along almost the whole 
lengtli of die coast of Chile (Figure 1). The northern 
locahties attributed to the Pliocene and Pleistocene are 
e.\clusi\elv nearshore emironment outcrops between 
Peninsula Mejillones and Coquimbo that have been de- 
scribed in detail by Herm (1969) and Le Rou.x et al. 
(2004, 2005, 2006)' Herm (1969) described the faunas 
from these locahties and also commented on the fauna 
of die La Cueva Formation, which overlies the Miocene 
Navidad Formation of central Chile. Pliocene faunas 
from soudrem Chile have been described from the 
islands Guafo (Frassinetti, 1997, 2000) and Guamblin 
iFrassinetti and Covacevich, 1995). Another fauna of 
late Miocene or early Phocene age from central Chile is 
knowTi from Lo Abarca near San Antonio (Covacevich 
and Frassinetti, 1990; DeVries, 2003). 

Current address: Institut fiir Geowissenschaften, Christian- 
Albrechts-Universitat zu Kiel, Ludewig-Me\Ti-Str 10, 24118 

The Tubul Formation (Figure 2) was first described by 
Ferugho (1949), wth its ty^ie area located on the south- 
ern side of the Gulf of Arauco (37° 14' S, 73°26' W). 
Onshore outcrops are limited to the east bv the Cor- 
dillera de Naliuelbuta and to tiie west by the Pacific 
Ocean. Feruglio (1949) described the sediments of the 
Tidiul Formation as light-gray soft tuffaceous sandy 
nmdstones with fossil remains, which can reach a 
thickness of up to 100 m (Garcia, 1968). Using macro- 
fossils, Briiggen (1950) showed that these sediments 
can be correlated vi.'ith those of Coquimbo. From their 
study ot microfossils, Martinez and Osorio (1968) con- 
cluded that the basal part of the Tubul Formation 
corresponds to the middle Pliocene. However, later 
studies by Martinez (1976) in Caleta Hueton led to 
the conclusion that the base of the Tubul Formation 
corresponds to the late Pliocene. Suhseqiientlv, Biro- 
Bagoczky (1979) recognized a lower and an upper 
member: the lower member has a thickness ot approx- 
imately 50 m and is formed by veiy fine, slightly 
calcareous, dark-gray sandstones, including abundant 
fossil marine fauna; the upper member also has a 
thickness of approximately 50 m and is formed by 
light-gray to olive fine sandstones and mudstones with 
rare fossils. 

According to Pineda (1983, 1986), deposition of the 
sediments ot the Tubul Formation was rapid, in a quiet 
zone close to the coast, maybe lagoonal. These sedi- 
ments should correspond to the Plio-Pleistocene-bound- 
aiy (Pineda, 1983, 1986) and occur at the following 
localities: Punta Pichicui, Estero Chupalla, Villa Alegre, 
Los Alamos, Quebrada Raquilco, El Tique, Estero 
Licauquen, and Minas Trihueco. According to Pineda, 
the Tubul Formation represents a transgressive se- 
quence. The presence of quartz-sand and a conglomer- 
ate at the base of the formation has been inteqireted as 
a former coastal line. Above this level, veiT fine clayey 
sandstones have been deposited, which should represent 

Paee 202 

THE NAUTILUS, Vol. 122, No. 4 



13 5 7! 

10 12 14 17 19 
13 18 

^ •Santiago 
w Lo Abarca 

La Cueva 


Tubul Formation 

, Valdivia 

Puerto Montt 


Elevation (m) 


200 400 



Figure 1. Extant ruiiges oi bivaK-t- and gastropod species oceiirniig at Las I'enas. 1. Einiiiciilii ^riiiji. 2. Tiiulciriopsis sulculiila. 3. 
Malletia chilensis. 4. Zijgochlamys patagonica. 5. Ctjclocardia vehitinus. 6. Macoma iiionuita. 7. Darina solenoides. 8. Ensis iiiticha. 
9. Retrotapes exalhidus. 10. Pandora cistula. 11. Epitoniu'm magellanicus . 12. Fiisitiiton iiKigilhniinis. I'l Tro/i/ioo geversianus. 14. 
"Xijmenopsis" dispar. 15. Chorus gigantens. 16. Nasaarius taeniolatus. 17. Adclniiiflon niicilld. IS, licln jiiic\hii. 19. Scaphander 
intcrntptus. Stars indicate Plio-Pleistocene fossil localities. 

S.N. Nielsen and C. Valdo\inos, 2008 

Page 203 

5880 km 


^\- ""V ]&»d^ 

637 km 639 . . 

Figure 2. Area ol tlie Tubul Formation and sample site at Las Pefias. Height of coa.stal liluli aliout 18 m. 

a shallow-marine emironment. Radic et al. (2005) cited 
a diickness of 100 to 500 m for die Tubul Formation 
based on well data and seismic lines. 

The molluscan fauna decribed in diis paper comes 
from the coastal bluffs near Las Peiias (Figure 2), which 
represent the uppermost part of the Tubul Formation. 


The mollusks described in this study belong to the col- 
lection of the late Professor Lajos Biro and are housed in 
the Departamento de Ciencias de la Tierra, Universidad 
de Concepcion, Chile. The material comes from near 
Punta Las Peiias (37"14'32" S. 73°26'02" W, Figure 2), 
south of the \iLlage Tubul and is designated hv the let- 
ter T' in die collections. The number of specimens 
inspected for each species is indicated bv a number in 
parendieses if die number is greater than one. Species 
are not fomialh' described because all except two are 
well known. Modem geographic ranges of species are 
mochfted from Valdovinos (1999). 


Class Bi\al\ia Linnaeus, 1758 
Family Nucuhdae Gray, 1824 
Genus Ennucula Iredale, 1931 

Ennucula gi'aiji id'Orbigm", 1846) 
(Figure 3) 

Xucida obliqua Sowerby, 1833: 5, pi. 16, fig. 21. 
Xiicula graiji d'Orbigny, 1846: vol. 5, 625 (nomen novum 

for A", obliqua Sowerbv, non Lamarck, 1819). 
Ennucula graiji (dOrbignvj.-Soot-Ryen, 1959: 13, pi. 1, 

fig. 8; Villarroel and St^iardo, 1998: 133, figs. 8, 70, 71, 

73, 107-109. 
Ennucula araucana (Philippi).-Vi]larroel and Stuardo, 

1998: 165 (partim), figs. 142-143. 

Ennucula valcliviana (PhiHppi). -Villarroel and Stuardo, 

1998: 165 (paiiim). figs. 1.34-137. 
Ennucula lehuensis (Philippi). -Villarroel and Stuardo, 

1998: 165 ipartun), figs. 138-139. 

Material E.xamined: 47 specimens: T/4 (2), T/5 (6), 
T/7 (5), T/11, T/17, T/204, T/237, T/386, T/1072, T/1276 
(3), T/1313, T/1394-1397, T/1537, T/l 749-1754, T/1968 
(7), T/1969 (2), T/1989-1990, T/204S, T/2090. 



Width 19.2 mm, lenetli 16.' 

Occ-urrence: Recent: Coquimbo to Magellan Region; 
Plio-Pleistocene: -'\rauco to ?Isla Guafo. 

Remarks: Nuculti barrosi Philippi, 1887 (=A'. arau- 
cana Philippi, 1887), A', lebuensis Philippi, 1887, and 
N. valcliviana Philippi, 1887, are Miocene species from 
the Navidad, Ranquil, and Sto. Domingo formadons of 
central and southern Chile. Howexer, their generic 
placement is not known and therefore possible relation- 
ships to E. graiji remain unclear. The specimens men- 
tioned and figured by Villarroel and Stuardo (1998) 
lielong in E. graiji and not in one of these Miocene 
species. The taxonomic positions of these older species 
must await a rexasion of the Chilean Miocene nuculoids. 

Genus Tindariopsis Verrill and Bush, 1897 

Tindaiiopsis sulculata (Gould, 1852) 
(Figure 4) 

Nucula striata King and Broderip, 1832: 337 (non 

Lamarck, 1805). 
Nucula sulculata Gould, 1852: 12, 434, pi. 37. figs. 539 

a-e (Couthouy MS). 
Nucula elcgans Hupe, 1854: 305, Conquiliologfa pi. 5, 

fig. 7; Philippi, 1887: 189, pi. 31, fig. 6. 
Tindariopsis sulculata (Gould).-Dell, 1964: 149; \'illar- 

roel and Stuardo, 1998: 144-145, figs. 39-41, 80-82, 


Page 204 

THE NAUTILUS, Vol. 122, No. 4 


;:^i>>i-- N 


9 --^^.^ ^^ 









.^ ^%. 



'4^ ft ^^ . ^ 

&> O - — (^ 






'* M 




^ h(>^-^ 

S. N. Nielsen and C. X'aldovinos, 200S 

Page 205 

Xiiciilana siilciilata (Gould). -Dell, 1971: 167. pi. I. Pigs. 

1-i, pi. 2, fig. 9. 
Niiculaiia elc^iani: iHupe).-Frassinetti and Co\ace\ifh, 

1995: 51, text-fig. 3a, pi. 1, figs. 1-5; Fra.ssinetti, 

1997: 60, pi. I, fig.s, .5-7. 
Tiiulariii sulcidaia (Gould).-Griltiii and Niel.sen, 2008: 

16. pi. 5, figs. 3-5. 

Material Examined: ca. 1S50 specimens: T/1 (92), T/S 
(24), T/9 (89), T/12 (17), T/13 (36), T/14 (17). T/15 (4S), 
T/16 (9), T/IS (198), T/19 (252), T/32 (10), T/53-54, 
T/14S (4), T/149 (9), T/164 (2), T/168 (8), T/206, T/207 
(4), T/236 (4), T/296 (3), T/542, T/974 (2), T/982-984 (ca. 
150), T/1083 (7), T/1 182-1 187 (ca. 90), T/1235 (6), 
T/1270 (293), T/130S (ca. 30), T/1379 (92), T/1518 (11), 
T/1536 (21), T/1624, T/1681 (ca.45), T/16S2 (6), T/1802- 
1804 (ca. 90). T/1911 (23), T/1912 (ca. 90), T/1913, 
T/1970 (15), T/2001 (5), T/2009 (23), T/2042 (7). 

Measureinent.s: \\'idth up to 16 mm (T/1682). 

Occurrence: Recent: Talcahuano to Strait of Magel- 
lan: Plio-Pleistocene: Arauco to Isla Guamblni. 

Remarks: As for man\' odier species, there are different 
names for Recent and fossil records of die same species. 
\'iUm'roel and Stuardo ( 1998, p. 145), in their review of the 
Recent and fossil species of Chilean Protohranchia, noted 
tliat "The ornamentation of die shell of 7^ .sidciilata is veiA' 
similai- to diat of tlie fossil species T. elegan.s (...). It differs 
from that b\- its shoitlier length and by the tmncation of its 
extreme posterior." However, we do not consider these 
differences as sufficient for specific separation and regard 
tliem as intraspecific \ariation instead. 

FamiK' Malletiidae H. and A. Adams, 1858 
Genus Malletia Moulins, 1832 

'SlaUctia chilctms Moulins, 1832 
(Figure 5) 

Malletia chilcnsis Mouhns. 1832: 85, pi. 1, figs. 1, S. 

Malletia incquaJis Dall, 1908: 219, 383. 

?MaHetia sp. Frassinetti and Co\ace\ich, 1995: 50. 

Material Examined: Nine specimens: T/544(2), 
T/1091-1093, T/1917, T/2176-2178. 

Measurements: Figiu'ed specimen width 26 mm 

Occurrence: Recent: Coquinibo to Magellan Region; 
Plio-Pleistocene: .Arauco to Isla Guamblm. 

Remarks: -\ccording to Villarroel and .Stuardo (1998), 
there are four or fi\e species of Malletia known troni 
Cliile. all of wliich are radier similar to each other. Apart 
from M chileusis, thev recorded M. pataopuica Mabille 

and Rochebrnne, 1889, M. ineiiualis Dall, 1908, and 
M. inafiellaiiica (Smith, 1875). Malletia hijadesi Mabille 
and Rochebrune, 1889 was listed as a proliablc .sviionym 
oi M. patagotuca. Malletia volchnanni (Philippi. 1887) 
most probably is of Miocene age and has not been 
rexised and its placement should be regarded with caution. 

Family M)tilidae Kalinesque, 1815 
Genus ?Miitilus Linnaeus, 1758 

Mi/tiliis sp. 
(Figure 6) 

Material E.xamined: One specimen: T/106. 

Measurements: Length 58 mm (T/106). 

Remarks: According to Valdo\inos (1999) there are 
nine species of Mvtilidae known from Chile. Based on 
the main characters of the our shell fragment, such as 
the pointed morphology of the umbo and fine external 
growtli striae, our specimen may correspond either to 
the genus Mijtilus or Choroinijtihts. In southern Chile 
the species Choromiitilus chonis (Molina, 1782) and 
Mijtilus cdulis clulcnsis Hupe, 1854, coexist, but it is 
not possible to identify our specimen to species level. 
M\-tilidae of uncertain generic and specific status are 
kniown from Miocene dirough Recent and are in need 
ol a detailed rexision. 

Family Pectinidae Rafinesque, 1815 
Genus Zi/gochlanii/s Ihering, 1907 

Zi/gochlami/f; patagonica (King and Broderip, 1832) 
(Figure 7) 

Pecteii patagoniciis King and Broderip, 1832, 5: 337. 

Pecten tenuicostatus Hupe, 1854: 291. pi. 5, fig. 4; Phi- 
lippi 1887: 203, pi. 47, fig. 1; Moricke, 1896: 580. 
pi. 12, figs. 13-16. 

Chlamifs patagonica patagonica (King and Broderip). 
-Beu, 1985: 1-11, pi. 1. figs. 1-4. '~ 

Zi/gflehlai)ujs patagonica (King and Broderip). -Waller. 
1991: 28-30, pi. 2, figs. 13, 14; Griffin and Nielsen, 
2008: 37, pi. 16, figs.^5, 6. 

Chlanii/s tenuicostatus (Hupe).-Frassinetti and Covace- 
\-ich. 1995: 52, text-fig. 3b, pi. 1, figs. 6-9. 

Material E.xamined: 305 specimens: T/63, T/79, 
T/1.35, T/147. T/166. T/175. T/180. T/222-223. T/243, 
T/255. T/256. T/293. T/305. T/307-309, T/311-312. 
T/329-334. T/361-380, T/387-399. T/432^34, 
T/491-504, T/577-578, T/725-726, T/728-772, T/775- 
777, T/806-820, T/1006-1017, T/1039, T/1074-1077, 
T/1080-10S2, T/1156-1157. T/1160-1166, T/1269. 

Figures 3-13. Bi\aKia. 3. Eitiuiciila grayi (d'Orbigny, 1846) [length 19.; 

4. Tindm-iopsis sulcuhita (Gould, 18.52) [length 

16.0 mm]. 5. Malle'tia chilensis .Mouhns, 1832 [length 16.0 mm]. 6. Mijtilus sp. [length .58.0 mm]. 7. Zijgochlmmjs patagonica (King 
and Broderip, 1832) [lengdi 75 mm]. 8. Cijclocardia vehitinus (E. A. Smith, L8S1) [length 18.5 mm]. 9. Macotna inornata (Hanlev 
1844) [lengdi 16.0 mm]. 10. Darina soJenoides (King and Broderip, 1.832) [lengdi 44.0 mm]. 11. Ensis macha (Molina, 17.82) [length 
97.2 mm]. 12. Retrotapes exalbidus (Dillw\Ti, 1817) [length 62.0 mm]. 1.3. Pandora cistula Gould, 1.S50 [length 23.0 mm]. 

Pase 206 

THE NAUTILUS, Vol. 122, No. 4 

T/1280-1281, T/1284, T/1309-1310, T/1358, T/1371- 
1378, T/1380-1386, T/1389-1392, T/1415-1424, 
T/1514-1516, T/1553-1559, T/1579-1586, T/1632, 
T/1677-1680, T/1714-172S, T/1827, T/1863-1875, 
T/1964-1967, T/2003-20()8, T/2030-2031, T/2033, 
T/2037-2038, T/2049, T/2052, T/2055-2056, T/2059- 
2065, T/2154-2167, T/2181-2192, T/2225-2229. 

Measurements: Height 75 mm, width 73 mm (T/223). 

Occurrence: Recent: Island of Chiloe to Strait of 
Magellan: Plio-Pleistocene: Arauco to Isla Guamblin. 

Remarks: Because revision of Chilean pectinids is not 
within die scope of this work, relationships of Z. patago- 
nicus to tlie northern species described by Herm (1969) 
are not discussed here. Zi/gochlamys pafagonicus is 
the type species of Psijchrochlamys Jonkers, 2003. 
However, like Griffin and Nielsen (2008) and Dijkstra 
and Marshall (2008), we consider Psychrochlamijs a 
synonym oiZijgochlamys. 

Family Carditidae Fleming, 1828 
Genus Ci/clocardia Conrad, 1867 

Ci/clocardia vehitiniis (E.A. Smith, 1881) 
(Figure 8) 

Carchta {Actinobohts) vehitiniis E.A. Smith, 1881: 42, 

pi. 5, fig. 8. 
Cardita volckrnanni Philippi, 1887: 167, pi. 37, Hg. 4. 
Cardiocardita volckrnanni (PhiHppi).-Frassinetti and 

Covace\4ch, 1995: 53, text-fig. 3c, pi. 1, figs. 10-17; 

Frassinetti, 1997: 65, pi. 1, figs. 17-20. 

Material Examined: 19 specimens: T/507-51(), T/ 
639, T/786-787, T/1546-1552, T/1628-1630, T/1756, 

Measurements: Width 18.5 mm, length 17 mm 

Occurrence: Recent: Arica to Beagle Canal; PHo- 
Pleistocene: Arauco to Isla Guainblin. 

Remarks: A number of different Cyclocardia species has 
been reported from soutliem Chile (Forcelli, 2000). 

Family Tellinidae Blainville, 1814 
Genus Macoma Leach, 1819 

Maconia inornata (Hanley, 1844) 
(Figure 9) 

Tellina inornata Hanley, 1844, 1847: 315, pi. 59, fig. 127. 
Tellina tubulensis Phihppi, 1887: 134, pi. 32, fig. 7. 
Macoma (Psaniinacoina) inornata (Hanley). -Soot-Ryen, 

1959: 62, pi. 4, figs. 34-36. 
Macoma tuhulensis (Philippi). -Frassinetti, 1997: 72, 

pi. 2, fig. 6. 

Material Examined: Eight specimens: T/64, T/721- 
724, T/789, T/1236, T/2067. 


26 mm. 

Width figured specimen (T/64) 

Occurrence: Recent: Atacama to Gulf of Ancud; Plio- 
Pleistocene: Arauco to Isla Guamblin. 

Remarks: Herm (1969, p. 119) described a specimen 
of Ardeamya sp. as the only tellinid from the Pliocene 
soutli of San Antonio. Since Herm knew the Philippi 
collection, it must be assumed that it is not conspecific 
with the species reported here. 

Family Mactridae Lamarck, 1809 
Genus Darina Gray, 1853 

Darina solcnoides (King and Broderip, 1832) 
(Figure 10) 

Erycina solenoides King and Broderip, 1832: 335. 
Daiina solenoides (King and Broderip). -Smith, 1905: 

Mactra sp. -Frassinetti and Covacevich, 1995: 54. 

Material Examined: One specimen: T/1238. 

Measurements: Width figured specimen (T/1238) 
44 mm. 

Occurrence: Recent: Strait of Magellan and Tierra 
del Fuego: Plio-Pleistocene: Arauco to Isla Guamblin. 

Remarks: Herm (1969) discussed the problems of 
related forms appearing in the Pliocene of Coquimbo 
to San Antonio which he placed in the genus Mulinia 
Gray, 1837. 

Family Pharidae H. and A. Adams, 1858 
Genus Ensis Schumacher, 1817 

£(i.s(.s' madia (Molina, 1782) 
(Figure 11) 

Solen macha Molina, 1782: 203; Hupe, 1854: 369, pi. 8, 

fig. 6. 
Solen gladiolus Sowerby, 1839: 153, pi. 43, fig. 4. 
Solen gladiolus? Sowerby. -Philippi, 1887: 169, pi. 34, 

fig: 9. 

Ensis macha (Molina).-Carcelles, 1944: 292. 
Ensis sp. -Frassinetti, 1997: 67. pi. 2, figs. 4-5. 

Material Examined: 38 specimens: T/192, T/579-580, 
T/793-797, T/1084, T/1 101-1102, T/1175-1176, T/1285- 
1290, T/1368-1370, T/1412-1414, T/1694, T/1707-1712, 
T/1 923, T/1972, T/2000, T/2068, T/2226-2227. 

Measurements: Up to 97.2 mm (T/1707). 

Occurrence: Recent: Caldera to Magellan Region; 
Plio-Pleistocene: Coquimbo to Isla Guafo. 

Remarks: Ensis macha is one of the few species which 
has a veiy wde geographical range in Pliocene and Re- 
cent faunas of Chile. During the Pleistocene it also 
reached Mejillones (Herm, 1969; own obsen'ations). 

Family Veneridae Rafinesque, 1815 
Genus Rcfrotapcs del Rio, 1997 

Retroiapes exalhidns (Dillwyn, 1817) 
(Figure 12) 

S, N, Nielsen ami C \'aklo\inos, 20()S 

Paee 207 

Venus cxalbida Chemnitz, 1795: 225. pi. 202. fiu;. 2 {iioii 

binoin.): Dillw)ii, 1S17: 170. 
Venus aerea Hupe, 1S54: 338. 
Venus subalbicfins Hnpe, 1854: 339. 
Venus araucana Philippi, 1887: 117, pi. 17. fig. fi; Ta\era 

and \eyl, 1958: 170, pi. 4, fig. 13c. 
Samarangia cxalbida (Dillw-Mil.-Carcelles, 1944: 287, 

pi. 12, figs. 93, 94. 
Eurbomalea araneana (Philippi). -Frassinetti, 1974: 47, 

figs. 1-2; Frassinetti and Co\'ace\ich, 1995: 54, 

text-fig. 3c, pi. 1, fig. 18; Frassinetti, 1997: 74, pi. 2, 

fig. 6." 
Retrotapcs cxalbida (Dillwni).-del Ri'o, 1997: 80-82, 

figs. 22, 23, 41. 
Retrotapes exalbidus (DilhvMij.-Reid and Osorio, 2000: 

139, fig. 5[; Griffin and Nielsen, 2008: 7. 35, pi. 1, 

figs. 2^, pi. 16. figs. 1-3. 

Material Examined: 669 specimens: T/21-25, T/44- 
49, T/56, T/58-62. T/65-73, T/75, T/78, T/84-l()9, 
T/136-140, T/151, T/153-163, T/173-176, T/18 1-191, 
T/198-203. T/208-213, T/216-221, T/226-235, T/241- 
242. T/249-253, T/257-263, T/266-272, T/277-282, 
T/319-32S, T/400-421, T/449-490, T/516-539, T/545- 
557, T/607-614. T/653-671, T/681-703, T/710-719, 
T/802, T/829-881, T/901-910, T/919-968, T/978, 
T/987-989, T/1027-1037, T/1 109-1 114, T/1 128-1 134, 
T/1142-1154, T/115S-1159, T/1194-1197, T/1212- 
1217. T/1228-1229, T/1237-1268, T/1298-1306, 
T/1316-1324, T/1333-1356, T/1425-1428, T/1511- 
1513. T/1534. T/1538-1544. T/164S-1676, T/1781- 
1796, T/1805-1809, T/1817-1826, T/1835-1862, 
T/1949-1963, T/201 1-2014, T/2035-2036, T/2039- 
2041. T/2046, T/205S, T/2081, T/2093-2097, T/2140- 
2147, T/2230-2249. 

Measurements: Up to width 62 mm, length 54.5 mm 

Occurrence: Recent: Chiloe to Strait of Magellan; 
Plio-Pleistocene: Arauco to Isla Guambli'n. 

Remarks: A number of different Eurbomalea species, 
most probabK- now belonging in Retrotapes. has been 
reported from nortliern Chile (Herm, 1969). 

FamOv Pandoridae Rafinesque. 1815 
Genus Pandora Bniguiere, 1797 

Pandora cistula Gould. 1850 
(Figure 13) 

Pandora cistula Gould, 1850: 217. 
Kennerhjia patagonica Dall, 1915: 450. 
Pandora sp.-Frassinetti and Co\ace\ich, 1995: 55, pi. 1. 
fig. 19. 

Material Examined: T/141-146, T/345, T/353-354, 
T/615-634. T/781. T/783, T/784 (3), T/972-973, T/986 
(2), T/1273 T/1291-1292, T/1311-1312, T/1405-1411, 
T/1535 (8), T/1755. T/2175, T/2225. 

Measurements: Width 23 mm (T/2225). 

Occurrence: Recent: Valparafso to Magellan Region; 
I'liu-Plcistocene: Arauco to Isla Guambli'n. 

Remarks: This species is the onlv ]i\ing species of 
Pandora iccordcd in Chile (see Forcelli. 2000). 

Class Gastropoda Cux-ier, 1795 
Family Epitoniidae Beny, 1910 
Genus Epitonium Roding, 1798 
Subgenus Nitidiseala de Bonn-, 1909 

Ej)ifonium (Nitidiseala) inagellanicus (Philippi, 1845) 
(Figure 14) 

Sealarid nitigelliinica Philippi, 1845; 46. 

Sealaria (Opalia) niagellaniea Strebel, 1905; 656, pi. 23, 

i'igs. 44a-f. 
Cirsotrenia (Coroniseala) magelliinicuiii (Philippi). - 

VVenz, 1938: 798, fig. 2322'. 
Epitonium (Nitidiseala ) )H0g(?//fl»iir(7(Philippi). -Ramirez, 

1987: 70, fig. 82. 
Epitonium biroi Frassinetti and Covacevich, 1995: 56, 

pi. 2, figs. 1-2; Frassinetti, 1997: 135, pi. 1, figs. 4-5. 

Material Examined: One specimen: T/2071. 

Measurements: Height 20.5 mm. 

Occurrence: Recent: Strait of Magellan; Plio-Pleisto- 
cene: Arauco to Plsla Guamblfn. 

Remarks: A nuniber of different epitoniid species has 
been reported Injm soiitliern Chile (Forcelli, 2000). The 
t)'pe material of Epitonium biroi Frassinetti and Cova- 
ce\ich, 1995 from Guambli'n was not revised, so it 
remains unclear if it represents a different species or a 

Family Naticidae Guilding. 1834 
Genus Euspira Agassiz, 1838 

Euspira guaniblinensis Frassinetti and Co\-ace\ich, 1995 
(Figure 15) 

Polinices (Euspira) guainblincnsis Frassinetti and Cova- 
cevich. 1995: 57, text-fig. 4a-e, pi. 2, figs. 3-5; Fras- 
sinetti, 1997: 135, pi. 1, figs. 6-9. 

Material Examined: 37 specimens: T/30, T/152, 
T/215, T/335-337, T/778, T/779, T/976, T/1177-11S1, T/ 
1272, T/1279, T/1627, T/1684, T/1696, T/1704, T/1705, 
T/1777, T/1899, T/1901, T/1902, T/1909, T/1940-1942, 
T/1991, T/2053, T/2068, T/2151, T/2210-2212, T/2255. 

Measurements: Height up to 22.5 mm, figured speci- 
men (T/1909) height 14 mm. 

Occurrence: Recent: Exfinct; Plio-Pleistocene: Ara- 
uco to Isla (.iuamblin. 

Remarks: None of tlie extant naticid species rev- 
iewed In- Pastorino (2005a) matches Euspira guanibli- 
nensis in its callus characters and we regard it as a valid 

Family Ranellidae Gra\, 1854 
Genus Fusitriton Cossmann, 1903 

Page 208 

THE NAUTILUS, Vol. 122, No. 4 




J 14 


,,\ V \- 



16 -^ 







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V 5 



^ 1^- 

i^ 21 





^ 24 \ 


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S. N. Nielsen ami C. \aldr 


Page 209 

Fusitritou magcllaniciis (Rodina;, 1798) 
(Figure 16) 

Mnrex inagcllaniciis Cheninitz. 17S.S: 275, pi. 164, 

fig. 1570 {non binoin.). 
Neptunea mogcUanica Roding, 179S: 116. 
Triton cancellatiim Lamarck, 1816: 4. 
Pricnc (Fusitriton) canccllatiis (Lamarck). -Cos.smann, 

1903: 109, fig. 8. 
Argobucciuiiin (Fusitritou) ciiucclldtus (Lamarck).- 

"" ^^'enz, 1938: 1058, fig. 3022. 
Argobucciuum (Argobuccinum) magelhiuicum (Rod- 

'" ing).-Carcelles. 1944: 247, pi. 2, fig. 23. 
Argobucciuum (Fusitritou) luagcllauicum (Roding). - 

Carcelles and Williamson, 1951: 286. 
Fusitritou cauccllatus ( Lamarck). -Smidi, 1970: 475, pi. 

42, figs. 4-10. 
Fusitritou magcllauicus (R6ding).-Ceniohorsl<y, 1977: 

107, fig. 3'. 

Material Examined: 20 specimens: T/150, T/171, 
T/178, T/214. T/315, T/340, T/511, T/512, T/572, T/971, 
T/llOO, T/12S2, T/1360, T/1575, T/1731, T/2026, 
T/2039, T/2169, T/2214, T/2252. 

Measurements: Figured specimen (T/2026) height 
73 mm. 

Occurrence: Recent: Los \'ilos to Magellan Region 
and Juan Fernandez Archipelago; Plio-Pleistocene: 

Remarks: This is a well known species that clear]\' 
does not present anv taxonomic problems. 

Genus Sa.ssia Bellardi, 1S72 

Sa-ssia leucostomoides (Sowerby, 1846) 
(Figure 17) 

Triton leucostomoides Sowerby, 1846: 240, pi. 4, fig. 64. 
Sassia leucostomoides ( Sowerby ).-Frassinetti, 1997: 136, 

pi. 1, figs. 11-14: Griffin and Nielsen, 2008: 49, 

pi. 21, figs. 9-14. 

Material Examined: SLx specimens: T/310, T/574; 
T/2032. T/2153. T/2196, T/2215. 

Measurements: Figured specimen (T/574) height 
44.4 mm. 

Occurrence: Recent: Extinct; Plio-Pleistocene: 
.\rauco to Isla Guafo. 

Remarks: This species was originally described from 
Guafo (Sowerby, 1846) and this record is the first from 
anotlier locafitw 

FamiK- Muricidae RaFinesqne, 1815 
t^'nus Tropliou Montlbrt, 1810 

Tropliou gcvcrsitiuus (Pallas, 1774) 
(Figure 18) 

Bucciuum gcversiauus Pallas, 1774: .33, pi. 3, figs. 1, 2. 
Murcx magcllauicus Gmelin, 1791: 3548. 
Fusus geversiauus ( Pallas). -Hupe, 1854: 167. 
Tropliou gcversiauus (Pallas). -Gould, 1852: 227. pi. 6, 
fig. 227. 

Material E.xamined: Nine specimens: T/513, T/569, 
T/575, T/640, T/798, T/1361, T/1574, T/2029, T/2253. 

Measurements: Height 62 mm (T/640), figured spec- 
imen (T/1361) 39 mm.' 

Occurrence: Recent: Ghiloe to Magellan Region; 
Plio-Pleistocene: Arauco. 

Remarks: The Chilean species of tlie genus Trophon 
piesent a high intraspecific variability, with many species 
described as synonyms of T. gcvcrsianus. A detailed re- 
vision of the genus Tropliou, including T gevcrsianus, 
was recently provided by Pastorino (2005b). 

Genus "Xipneuopsis" Powell, 1951 

"XipueiHipsis" ct. "-V " dispar (Rochebnnie and Mabille, 


(Figure 19) 

Fusus dispar Rochebrune and Mabille, 1889: H57, pi. 2, 

fig. 3. 
Trophon dispar (Rochebrune and Mabille). -Carcelles 

and Williamson, 1951:289. 
Xi/menopsis dispar (Rochebrune and Mabille). -Powell, 

1951: 159. 

Material Examined: 16 specimens: T/352, T/586 (5), 
T/588 (7), T/1357, T/2219, T/2245. 

Measurements: Figured specimen (T/.352) 22 mm. 

Occurrence: Recent: Tiei'ra del Fuego: Plio- 
Pleistocene: Arauco. 

Remarks: Pastorino and Harasewych (2000) stated 
that this species does not belong in Xijmeuopsis but did 
not pro\ide a more appropriate generic-level placement 
in the Muricidae. 

Genus Choiiis Gray, 1847 

Chorus gigauteus (Lesson, 1830) 
(Figure 2()) 

Monoceros gigauteus Lesson, 1830: 405, N° 165, pi. 11, 
fig. 4. - 

Figures 14-23. Gastropoda and Scaphopoda. 14. Epitonium (Nitidiscah) magellanicus (PhiUppi, 1845) [height 25.0 mm]. 15. Euspirn 
guamhlineiisis Frassinetti and Covacevich. 1995 [height 22.5 mm]. 16. Fusitiiton magellanicus (Roding, 1798) [height 73.0 mm]. 17. 
Sassia leucostomoides (Sowerby, 1S46) [height 44.4 mm], 18. Trophon geversianus (Pallas, 1774) [iieight 62.0 mm]. 19. "Xijmeuopsis" cf. 
"X" dispar (Rochebrune and Mabille, 1889) [height 22.0 mm], 20. Chonis gigauteus (Lesson, 1830) [height 95.0 mm]. 21. Nassariiis 
taeniolatus (Philippi, 1845) [height 11.0 mm], 22. Adehmelon ancilla (Ligtiifoot, 1786) [height 142.0 mm], 23. Bela paeslcii Strebel, 
1905 [height 16.0 mm], 24. Scaphander intemiptus Dall, 1889 [height 18.5 mm], 25. Dentalium sp. [lengtii 72.0 mm]. 

Page 210 

THE NAUTILUS, Vol. 122, No. 4 

Monoceros giganteus? Lesson.-Moricke, 1896: 564. 
Rapana (Chonis) gigantea ( Lesson ).-Wenz, 1941: 1038, 

fig. 3080. 
Chonis giganteus (Lesson). -Carcelles, 1954: 271, pi. 5, 

figs. 1-11; Herni, 1969: 135, pi. 15, figs. 4a, 4b; 

DeVries, 1997: 132, pi. 1, figs. 1-4, pi. 4, fig. 4. 
Chonis grandis {Philippi).-Herni, 1969: pi. 15. figs. 

3a, 3b. 

Material E.\amined: 17 specimens: T/576, T/985, 
T/1174, T/1193, T/1904-1905, T/1907-1908, T/1910, 
T/1984, T/2170-2172, T/2208, T/2213, T/2272, one 
specimen without number. 

Measurements: Height up to 95 mm (T/985), figured 
specimi'ii (T/iy()5) 80.5 mm. 

Occurrence: Recent: Papudo to Calbuco; Plio-Pleis- 
tocene: southern Peru to Arauco. 

Remarks: The species of Chonis have been reviewed 
by Herm (1969) and DeVries (1997). DeVries (1997) 
also gave new diagnoses and descriptions. Regarding 
the stratigraphic range of Chonis giganteus cited by 
DeVries (1997), its presence in the Tubul Formation 
implies a maximum age ot veiy late Pliocene to early 
Pleistocene for these deposits. 

Family Nassariidae Iredale, 1916 
Genus Nassaiiiis Dumeril, 1806 

Nassariiis tacniolatiis (Pinlippi, 1845) 
(Figure 21) 

Buccinum taeniohitns Pinlippi, 1845: 69; Hupe, 1854: 

207, pi. 4, fig. 9. 
Nassaiius taciiiolatns (Philippi).-Keen, 1971: 609, fig. 


Material Examined: Tlu'ee specimens: T/586 (2), 

Measurements: 11 nnn (T/1687). 

Occurrence: Recent: Papudo to Chonos Archipelago; 
Plio-Pleistocenc: Arauco. 

Remarks: This species is the only representative of 
Nassarius in southern Chile, while two more species are 
found li\dng off central and northern Chile (N. gaiji 
(Kiener, 1835) and N. dentifer (Powys, 1835); see Mar- 
incovich, 1973). 

Family Volutidae Rafinesque, 1815 
Genus Adelomclon Dall, 1906 

Adelomelon ancilla (Ligthfoot, 1786) 
(Figure 22) 

Vnhila anciUa Lightfoot, 1786: 84. 

Vohila graciUs Wood, 1828: 59, pi. 3, hg. 2. 

Cimhiola ancilla (Lightfoot).-Pace, 1902: 28, pi. 7, 

figs. 1-16. 
Adelomelon (Adeloniclon) ancilla (Liglitf()Ot).~Wenz, 

1938: 1347, tig. 3816. 
Adelomelon ancilla (Lightfoot). -Dall, 1906: 143. 

Material Examined: One specimen: T/1227. 

Measureinents: Figured specimen (T/1227) 142 mm. 

Occurrence: Recent: Chiloe to Magellan Region; 
Plio-Pleistocene: Arauco to ?Isla Guafo. 

Remarks: It is nuclear ii A. recondifus Frassinetti, 
1997, is a diiferent species or a juvenile A. ancilla. A 
large specimen ol presumably A. rcconditus was found 
on Guafo (coll. Nielsen), but presei"vation does not allow 
assignment to a species. The Chilean fossil species of 
Adelomelon were recently re\ased by Nielsen and Fras- 
sinetti (2007a). 

Family Turridae H. and A. Adams, 1853 
Genus Bela Leach in Gray, 1847 

Bela paesleri Strebel, 1905 
(Figure 23) 

Bela paessleri Strebel, 1905: 588, pi. 22, figs. 35, 35a-b. 
Mangelia {Bela) paessleri (Strebel). -Carcelles, 1950: 67. 

Material Examined: 14 specimens: T/5S9 (6, in part), 
T/59() (7), T/1626. 

Measurements: Height 16.7 nun (T/1626). 

Occurrence: Recent: Valdivia to Magellan Region; 
Pho-Pleistocene: Arauco. 

Remarks: This species is in need of revision, but be- 
cause the protoconch is not presened in these speci- 
mens, it is not possible to verify its generic position. 

Familv Scaphandridae Montfort, 1810 
Genus Seaplnnider Montfort, 1810 

Scaphander (i)/('rr;(/)//(.s D;ill, 1889 
(Figure 24) 

Scaphander iuterruptus Dall, 1889, 12: 297, pi. 12, 
fig. 12. 

Material Examined: Six specimens: T/244, T/583, 
T/584 (2). T/975, one specimen without number. 

Measurements: Figured specimen without inunber, 
18.5 nun. 

Occurrence: Recent: Panama to Magellan Region; 
Plio-Plcistocene: Arauco. 

Remarks: The Recent and fossil Chilean Cephalaspi- 
dea aic in need of revision. Scaphander cosmophilus 
(Sowerby, 1846) is known from the Pliocene of Isla 
Guafo (Frassinetti, 2000) and possibly from Isla Guam- 
blin (Frassinetti and Covacevich, 1995). Scaphander cos- 
mophilus was placed in the Genus K/:iitoa by Griffin and 
Nielsen (2008). It is not clear if the two species are 
closely related. 

Class Scaphnpoda l-ironn, 1862 
Familv Dcntaliichic (iiax. 1S47 
Genus Denfaliuni Linnaeus, 1758 

Dentalium sp. 
(Figure 25) 

S. N. Nielsen and C. \''aldo\inos, 2()()S 

Page 211 

Material Examined: T\\() specimens: T/1937-193S. 
Measurements: Fignred specimen (T/1937) length 

Remarks: Dentaliidae are well represented in Chile at 
least since the Miocene (see e.g., Philippi, 1887; Herm, 
1969). A rexiew of the group has never been under- 


Sexenil t;L\a outside the mollusks are knox\n from the 
Tubul Formation. The Biro collection contains about 
225 specimens ot barnacles, remains ot two decapod 
crabs, bnozoans, about 260 specimens of the brachio- 
pod Mogcllania venosa Solander, 1786, two different 
echiuoids and some shark teeth. 

Some of die fauna represented in the collection tloes 
not come from the Tubul Formation but from die Mio- 
cene Ranijuil Formation, most probablv from the nearby 
loc;dit\^ Punta El Fraile {see Nielsen et al, 2004; Finger 
et al., 2007). Miocene representatives in the Tubul col- 
lection include the nautiloid cephalopod Atiiria cuhacn- 
sis (Lea. 1841) and the olivid gastropod Ldiitprocloiitina 
dimidiata (Sowerbw 1846). 


UnderKing the Tubul Formation is the late Miocene to 
earK" Phocene Ranquil Formation (Garcia, 1968; Pineda, 
1986; Finger et al., 2007), the fauna of which is similar 
to that oi die Navidad Formation south of Valparaiso 
(see Philippi, 1887; Groxes and Nielsen, 2003; Nielsen, 
2004, 2005; Nielsen and Frassinetti, 2003; Nielsen et al., 
2004). This fauna has been reworked and is of earK- to 
middle Miocene age (DeXnes and Frassinetti, 2003; 
Finger et al., 2007) and contains manv tropical to sub- 
tropical gastropod genera, such as Neiita, Strombus, 
Xe)wp]wra. Distorsio. Echinophoiia, Ficus. Terebra, 
and Architectonica (Philippi. 1887; Covace\ich and 
Frassinetti, 1980; Nielsen, 2005; Nielsen and DeVries, 
2002; Nielsen and Frassinetti, 2007b), diat disappeared 
from Chile during the late Miocene climate cooling. 
A number of die gastropod genera described herein 
{Epitoniiim. Trophon. Chonis, Kasmrius) were already 
present in Chile during the Vliocene, but were repre- 
sented h\ different species. If and how these are related 
to the li\ing taxa has vet to be investigated. 

The Pliocene to Pleistocene faunas from noitlieni Chile 
have a different composition than diat of the Tubul For- 
mation, containing abundant Trochoidea, FissureUa, Tiir- 
ritella, Cnicibulum, Trochita, Crepidula, different species 
of Naticidae (including Sinttm cijmba). Ar^obuccinum, 
Xassaiius. Acanthina. Chonis, and Ohvci peniviana 
(Herm, 1969: De\nes, 1997. 2003; own data), represent- 
ing toda\s coastal fauna of noithem to central Cliile. 
Howe\'er. these are nearshore faunas witii many taxa from 

rock"\' coasts, while that of Tubul is a soft-bottom fauna 
Ironi slightly deeper water, missing typical rocky-shore 
elements such as FissureUa. Acanthina, and Concholepas, 
which are present in the Plio-Pleistocene deposits of 
noi-theni Chile and in the modern iauna along the whole 
Chilean coast (McLean, 1984; DeVries, 1995, 2003). 

The Tubul fauna has prexaously been compared to 
faunas coming from Guambh'n and Guafo islands farther 
south (Frassinetti, 1997, 2000; Frassinetti and Covace- 
\ich, 1995; Table 1). The fauna from Guaniblin is close 
to the Tubul fauna and Frassinetti and Covacevich 
(1995) and Frassinetti (2000), comparing both faunas 
directk', cited new Tubul species (Epituniuni biroi, Tro- 
jihon covaccvichi. and Hindsiclava ignorata\, and sug- 
gested tliat Epitoniiim biroi may be conspecific widi 
E. (Nitidiscala) magcUaniciis, Trophon covacevichi may 
be conspecific with Trophon gcversianiis, and Hindsi- 
clava ignorata may be conspecific with Bela paesleri. 
However, to confirm these synon\niies more material 
should be directly compared. 

The fauna described by Frassinetti (1997, 2000) from 
Guafo contains a niLxture of species also present at 
Guambh'n island and Tubul and species known from 
older deposits, e.g., Panopca chihwnsis, IncatcUa cliilen- 
sis. Echinophoria sp. (cited as Semicassis sp.), Chonis 
(hiUaris. Fcninn spp., and Gcmmula siibacquaUs. It seems 
likely that these taxa represent a fauna intermediate 
in age betxveen the older faunas and the Tubul-Guam- 
bli'n fauna but it is possible that specimens of tiie 
(jlder fauna ha\'e been reworked from older beds and 
incoqDorated into \ounger sediments as has been 
demonstrated for the Navidad, Ranquil and Lacui for- 
mations of .Mio-Pliocene age (Finger et al, 2007). The 
presence of Chonis doliaris and Incatella chilensis 
suggests a latest Miocene age for the Guafo fauna 
(D^eVries, 1997, 2007). 

The first appeareance of Chonis giganteus in soudi- 
ern Peru during the latest Phocene (DeVries, 1997) 
makes this a likely maximum age for the Tubul fauna. 
Indeed, this agrees well with strontium isotope stratigra- 
phy data from calcitic shells of Zijgocldamijs patagonica 
coming from the same localih', which confirm an early 
Pleistocene age (unpublished data). The two apparently 
extinct species present, Euspira guanibUnensis and Sas- 
sia leucostonioides , may well be found living off Chile in 
the future. Comparison with living ranges of the species 
lound in the Tubul Formation (Figure 1) permits one to 
identify two regions with 16 species in common. The 
fauna of the Tubul Formation resembles most closely 
the present-day faunas of northern Cliiloe island and die 
-Magellan Region, whereas northern faunas of Pliocene 
to Pleistocene age have a composition similar to that of 
uuxlern northern and central Chile. Water temperatures 
for the Arauco area are accordingly interpreted to have 
been colder than today. It is presently not known if the 
more northern species (Figure 1, numbers 6, 15, 16) 
previously tolerated cooler waters or if the southern spe- 
cies (Figure 1, numbers 7, 11, 14) tolerated warmer 
waters. However, it is possible that an additional factor 

Page 212 

THE NAUTILUS, Vol. 122, No. 4 

Table 1. Comparison of iiKjUusk species found at Tubul. 
Guafo and Guanibli'n. 




? (sp.) 


Ennucula graiji x ? (ban-osi) 

Tindariopsis x x 

Malletia chilensis x 
Mijtibis sp. X 

Zijgochlaim/s x ? (cf huj^cnnus) < {tciinirostatus) 

Lucinomn sp. x 

Cijclocarcha x x (volchnanni) x (volckmanni) 

Macomu inoniata x x {tuljulcnsis) 

Dahlia solenoides x x (Mactm sp.) 

Ensis macha x x (sp.) 

Rctrotapes x x (araucana) x (iiniucaiia) 

Pandora cistula x x (sp.) 

Panopea x 



Epitoniiim x ? (biroi) ? (biroi) 

Turritella x 

Euspira x x x 

Echinophoria sp. x (Semicassis) 

Fiisitriton x 

Sassia x x 

Cijmatium x 

Trophon x ? (covacevichi) ? (sp.) 

Trophon parens x 

"Xymenopsis" cf. x ? (hiiilliche) 

"X." dispar 
Xymenopsis hero x 

Chorus giganteus x 

Chorus doliaris 





Nassarius gaiji 


Phos chilensis 


Pcnion spp. 




? (reconditus) 


Bela paesleri 













? icosmophilns) 

? (cosmophilus) 



Deiilalinm sp. 


? (.sp.) 

independent of water temperature is responsible 
for these widely separated species triplets today in con- 
tiast to tlieir common occurrence during die early 


First of all we express our gratitude to Profesor Lajos 
Biro-Bagoczky (1929-1993) who, together wnth others, 
assembled this collection during more than 15 years. 
Arturo Quinzio (Departamento Ciencias de la Tierra, 
Universidad de Concepcion, Chile) made this work 
possible by gi^ing access to this collection, Klaus Bandel 
(Geologisch-Paliiontologisches Institut, Universitiit 
Hamburg, Germany) is tiianked for discussion on the 
Tubul Formation and fauna. Daniel Melnick (GFZ 
Potsdam) helped with the DEM map for die distribution 
figure. Comments and suggestions by Tom DeVries 
(Burton, USA) improved die manuscript. Work of SNN 
was supported financially by Deutsche Forschungsge- 
meinscliaft erant Ni 699/4-1 and 4-2. 


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THE NAUTILUS 122(4):2 17-227, 2()()S 

Fat^e 21' 

Radula moiphology in veined rapa whelks, Rapana venosa 
(\^alenciennes, 1846) (Gastropoda: Muricidae) from 
Chesapeake Bay, USA 

Juliana M. Harding' 
Stefanic M. Gera 
Roger Mann 

Department of Fisheries Science 
\'irs;inia Institute of Marine Science 
Gloucester Point. Mrginia 23(162 USA 
jharding@\ims . edu 


Radula length, widdi, niuiiber of transverse rows of teeth, and 
rachidian tooth dimensions (central cusp height, central cusp 
b;ise width, and rachidian tooth base width) were examined in 
relation to leined rapa whelk shell length. Radula length and 
widdi increase hnearK' with whelk shell length. The number of 
ti'ansverse rows of radular teetli increase with whelk shell 
lengtli. \\'ithin an individual, central cusp height of the rachi- 
dian tooth increases with increasing distance from the anterior 
of die radula. Central cusp height of the rachidian tooth, an 
indicator of tootii wear or use. was least for teeth in rows 1 and 
11. Teeth in radular row 21 appear to be in a transition zone 
from high to low wear or use. Within a radula and within a size 
class, die ratio of central cusp base widdi to rachidian tootli 
base widdi does not change. Central cusp base width, central 
cusp height and rachidian toodi base widdi were significantly 
smaller in females than in males indicating sexual chmoqDliism 
in rachidian tootii shape for rapa whelks. Patterns of wear as 
indicated h\ central cusp base widtii to central cusp height ratio 
values w'ere not significantK- different between sexes and mav 
sen'e as an indication that feeding strategies and/or prey may 
be similar behveen animals of different sex lint similar size. 

Addi tion al Kcijico rd.s: 

Neogastropoda, rac'hidian teedi, allometiy 


The radula is a chitinous ribbon-like series of nearlv 
colorless transverse tooth rows resting atop the radula 
membrane (\Vu, 1965; Radwin and Wells, 1968). Muri- 
cid gastropods use die anterior teeth when drilling holes 
in bixahe prev (Carriker, 1961, 1981; Fujioka, 1985). 
As anterior teeth are worn down, tliey are replaced by 
xounger teedi diat are formed in the radular sac and 
"radualK' mox'ed fonvard along; the radula (Isarankura 

.■Vutlior for correspondence 

and Runhani, 1968; Carriker, 1981). Muricid radulae 
have between 100 and 500 transverse rows of teeth (e.g., 
Carriker, 1961; Radwin and Wells, 1968; Fujioka, 1985). 
Each transverse row of teetli consists of a central rachi- 
dian (R) tootii and two slender marginal teeth (M) in die 
tooth formation M + R + M (Carriker, 1969). The cen- 
tral rachidian tootii in each transverse row is responsible 
for most of the rasping and physical shell removal during 
drilling while the marginal teeth synchronously tear flesh 
from prey (Carriker, 1969; Carriker et al., 1974; Krutak, 
1977). Thus, the rachidian teeth show more wear, or 
reduction in size with use, than marginal teeth found in 
the same tran.sxerse rows (Carriker et al., 1974). This 
trend is particularly evident at the anterior end of the 
radula where the rachidian cusps in the most anterior 
row(s) may be completelv remo\'ed by use (Carriker, 
1969, 1974; Fujioka, 19S5)'. 

Veined rapa whelks {Rapana venosa, \'alenciennes 
1846, Muricidae) are predatoiT marine gastropods that, 
while originally native to Japanese and Korean waters 
(Tsi et al., 1983), have successfully invaded marine and 
estuarine habitats in the Black, Adriatic, Aegean, Medi- 
terranean (Mann et al, 2004), and North Seas (Vink 
et al., 2005) as well as the Rio de la Plata (Pastorino 
et al., 2000) and Chesapeake Bav, USA (Harding and 
Mann, 1999). At the present time, the Chesapeake Bay 
rapa whelk population is die only kiiowai population of 
rapa whelks in North America. 

Rapa whelks provide an unusual opportunity' to 
investigate allonietric changes in radula morphology 
across a wide size range of individuals because diey 
reach terminal shell lengths in excess of 170 mm (Wu. 
1988; Harding and Mann, 2005). Like otiier muricids 
(Paine, 1966), rapa whelks experience ontogenetic 
shifts in diet (Harding and Mann, 2001) as well as 
predation strategy and resulting predation signatures 
in prey valves (Harding et al., 2007). Small (<35 mm 
shell length) rapa whelks drill their prey (Harding and 
Mann, 2001; Hardina; et al.. 2007) incluchng barnacles 

Paee 218 

THE NAUTILUS, Vol. 122, No. 4 

(Balanus sp., ClitJwmahis sp.), mussels {Mtjtihis sp., 
Gcukensia demissa), soft shell clams [Mija arenaria), and 
ON'sters (Crassostrea virginica). At shell lengths above 
35 mm, rapa whelks eat larger bixalves (Harding and 
Mann, 2001) including oysters and northern quahogs 
{Merceuaiia mercenaria) and t)''pically either edge bore 
their prev or leave no signatures (Morton, 1994; Harding 

We quantitati\ely describe radula and rachitlian tooth 
moiphology for a size range of rapa whelks irom Chesa- 
peake Bay, USA. Rachidian teeth in Rapaiia have a large 
central cusp flanked bv two smallei^ cusps (Arakawa, 
1964; Wu, 1965). Shell length, the maximum dimension 
from the tip of tlie spire to the Iiottom of the siphonal 
canal, is used as the metric ot \vhelk size. Shell length 
does not fluctuate with season or other lactors. Relation- 
ships between shell length and radula dimensions are 
quantitativeh' described ior male and female rapa 
whelks. Within each radula, rachidian tooth moq3holog\' 
is described along the length oi the I'adula bv measuring 
ratios of rachidian ttjoth central cusp base width to cen- 
tral cusp height and central cusp base width to rachidian 
tooth base width. The resulting ratios are compared 
bet\veen teeth along the length of an individual radula 
and across radulae from male antl female whelks as well 
as from a size range of Chesapeake Bay rapa whelks. 


Rapa whelks with shell lengths (SL) in excess of 70 mm 
were obtained from the lower Chesapeake Bay, USA as 
donations to the Virsinia Institute of Marine Science 
(VIMS) rapa whelk bounts' program. Rapa whelks less 
than 70 mm SL were cultured at VIMS, Gloucester 
Point, \irginia to supplement the lower SL range of rapa 
whelks because individuals less than 70 mm SL were not 
available through the boimty program. 

At the time of whelk collection, SL was measured in 
mm and whelks were assigned to shell length classes 
diat were established to evenly categorize the potential 
SL range (1-180 mm SL). Petite, small, medium, and 
large classifications corresponded to whelk SL ranges 
of less than 45 nmi, 45.1-90 nun, 90.1-1.'35 mm, and 
135.1-180 mm, respectively. 

Whelks were frozen after collection and thawed to 
facilitate dissection and removal of radulae. Whelks 
wei-e sexed during dissection and distinguished as male 
or female on tlie basis of penis length and gonad color 
after Mann et al. (2006). For the purposes of discussion 
herein, true females (penis length = mm, bright vellow 
gonad) and imposex females (penis length <2() mm, 
bright yellow gonad) are grouped together per Mann 
et al. (2006). Tvpically, radulae were dissected out of 
the whelk proboscis. However, two conti'ol radulae were 
removed bv soaking the proboscis for 24 hours in 10% 
sodiiun hvdroxidc to ensure that dissection removed the 
entire radula intact. Onlv intact radulae were used in 
this studv. 

After removal from whelks, images of the complete 
radula were taken using a digital camera mounted on a 
dissecting microscope for measurement of total radnla 
length, total anterior to posterior distance (mm) and rad- 
ula width, dre maximum lateral distance across the bases 
of the marginal and rachidian teeth at the first transverse 
row of teeth (Figure 1). The odontophore was removed 
and then the rachidian teeth were systematically 
removed from eveiy 10th transverse tooth row (e.g.. 
Row 1, 11, 21 etc.) along die radulae moving from ante- 
rior to posterior (Figures 2 and 3). Tooth rows were 
removed with a size 10 scalpel blade for larger indi- 
viduals, and with shaipened needles for culttired 
individuals less than 66 mm SL. 

Digital images were taken of each individual rachidian 
tooth after removal with the tooth positioned convex 
side down. T)'pical magnification of intlividual teeth 
used for digital images ranged from 50 x for whelks with 
SL greater than 147 mm to 90 x for whelks less than 
45-50 mm SL. Measurements (mm. Figure 4) of the 
rachidian tooth central cusp base width (LI), maximum 
central cusp height (L2) measured from the tip of die 
central cusp to the midpoint of LI, and the maximum 
rachidian tooth base width (L3) were made on the 
resulting images. The terniinologv' used to describe 
tooth moipholog)' follows that of Kool (1993). 

Data Analyses: Significance levels for all statistical 
tests were set at alpha = 0.05 a j)nori. Fisher's multiple 
comparison tests were used for j)ost hoc comparisons 
when appropriate. 

Radula Allometn and Gross Moi-jihologj': Linear 
and power regression models wxTt' used to describe 
relationships within sexes between rapa whelk shell 
lengdi and radula length, radula width, and number of 
rows of transverse teetfi per radula. The relationship 
between radula lengdi and both radula vviddi and die 
number of transverse rows of teeth within radulae from 
males and females were also examined widi both linear 
and power regressions. The slopes of all moiphological 
regressions were compared between sexes with t tests 
(per Zar, 1996) on raw data or on logarithm transformed 
data if the power model was deemed more appropriate 
than the linear model. 

Rachidian Tooth Dimensions: The relationsliip bet- 
ween central cusp base width (LI, Figure 4) and size 
class of the whelk from which it came was evaluated widi 
a tliree-vvay ANONA. (whelk size class x tooth row x 
sex) with tlie response being the maximum base width of 
the rachidian tooth. These data satisfied assumptions 
of homogeneity of variance aftiT the logarithm trans- 
formation was applied but not normalitv. 

Tlie relationship between rachidian tooth central cusp 
height (L2, Figure 4), the size class of the v\helk from 
which it came, and the tooth row was evaluated with a 
thr(H' factor ANOVA (whelk size class x tooth row x 
sex). These data satisfied neither the assumption of 
normality nor homogeneity of variance regardless of die 

J. M. Harding et al., 2008 

Page 219 

Figures 1-3. Radula of \eined rapa whelk. 1. Radula fiT)iii an 84. 2 mm shell lengdi (SL) whelk showing general moiplinlogical 
features and orientation. 2. A rachichan tootii from row 1 of tlie radnla from a 134 mm SL whelk showing wean 3. An unworn 
rachidian toodi from row 41 from a 122 mm SL whelk. Scale bar = 1 mm. Abbreviations: r = I'ow number; R = rachidian; Od = 
odontophore; A = anterior; P = posterior 

m, .sqr 

transformation (logarithm, natural logarith 
root, arcsine). 

The relationship between rachidian tooth base width 
(L3, Figure 4), row number, sex, and size class of the 
whelk from which it came was evaluated witli a three- 
wa\" AN OVA (whelk size class x tooth row x sex). These 
data satisfied the assumption of homogeneity of variance 
without transformation but did not satisfy the assump- 
tion of normalitv regardless of the transformation (loga- 
rithm, natural logarithm, square root, arcsine) and were 
analyzed without transformation. 

The ratio of rachidian tootli central cusp base width 
(LI) to central cusp tooth height (L2) was calculated for 

each rachidian tooth. Using a ratio that compares base 
width to tooth height is appropriate in a structure where 
both the base width and tooth height change along the 
length ot the structure with ontogeny. Not only does the 
ratio allow for scaling when comparing individual teeth 
along the radula length, but it can also be used as an 
index of wear because width does not change with use. 
In this case a central cusp base vwdth: central cusp tooth 
lieight ratio >1 is indicative of wear The hrst 81 rows 
were chosen for analysis because eveiy radula dissected 
had at least 81 rows. These data satisfied neither the 
assumption of homogeneity of variance nor normality- 
regardless of the transformation (logarithm, natural 

Page 220 

THE NAUTILUS, Vol, 122, No. 4 

Figure 4. Veined rapa whelk lachidiaii tootli with the mea- 
surements made in diis stiidv identified: raeliidian tooth central 
cusp base width (LI), rachidian tooth central cusp height {L2), 
and rachidiau tooth base width (L3). Scale bar = 0.25 mm. 

logarithm, square root, arcsine). A three-way ANOVA 
(whelk size class x tootli row number x sex) was used 
to evaluate tlie ratio of central cusp width to central cusp 
height (L1:L2). 

The ratio of central cusp base width (LI) to rachidian 
tooth base width (L3) ior each rachidian tooth was eval- 
uated with a 3 way ANOVA (size class x row x sex) to 
describe potential changes in tooth shape with ontogeny. 
Data satisfied assumptions of homogeneity of variance 
without transformation but not normality (either with or 
without transformation, e.g., logarithm, natural loga- 
rithm, square root, arcsine). Changes in the L1:L3 ratio 
across whelk size classes reflect ontogenetic changes in 
tooth moiphology that may be related to sexual dimor- 
phism (Fujioka, 1982, 1984) and which may act to dis- 
perse relatively greater strike force during feeding in 
larger whelks. 


Only radulae from whelks collected when water tem- 
peratures were above 11-12°C and feeding were used 

(Harding, unpublished data). All radulae examined were 
intact. Descriptive morphological data were collected 
from 39 rapa whelk radulae. These radulae were from 
rapa whelks with shell lengths between 20.2 mm and 
174 mm (Table 1, Figure 5). Rapa whelk radula lengths 
ranged from 4.33 to 51.05 mm with corresponding 
radula widths of 0.23 to 2.67 mm and total number of 
transverse rows of teeth of 89 to 210, respectively. 
Radula length was an average of 21.4% (standard error 
= 0.61%) of shell length. 

Radula Allometry and Gross Moi-phology: Regres- 
sion coefficients for the fitted linear and power regres- 
sion models used to describe relationships between rapa 
whelk shell length and radula morphology and between 
rapa whelk radida measurements are given in Table 2A. 

The linear model is suggested as a better descriptor of 
the relationship between rapa whelk shell lengdi (SL) 
and radula length (RL; Figure 6A) for both sexes since 
the coefficients of determination from both models are 
identical (Table 2A) and the linear model provides the 
simplest description of the data. The slope for the SL- 
RL relationship in males is significantly higher than that 
for females (t-test. Table 2B)." 

The power Tuotlel more accurately described the rela- 
tionships between rapa whelk SL and radula width (RW; 
Figure 6B) by predicting a radula witlth e(|ual to at a 
shell lengdi ecjual to 0. The coefficient of determination 
for the linear model describing the relationship between 
shell length and the number of rows of radular teeth was 
higher (females = 0.63, males = 0.76, Table 2A) than that 
of the corresponding power model (females = 0.58, 
males = 0.69, Table 2A) for both sexes and the linear 
model predicted a positive number of I'ows of teeth at 
shell lengths of mm (Figiu'e 6C). 

The relationship between radula length (RL) and rad- 
ula width (RW) was described with a power model for 
both sexes (Table 2A, Figure 6D) which predicted a 
radula width of at a radula length of and had a higher 
coefficient of determination than the corresponding lin- 
ear model (Table 2A). The linear model describing die 
relationship between radula lengtli (RL) and number 
of transverse rows of teeth has a higher coefficient of 
dctcniiiiiation tlian the corresponding power model 

Table 1. Description ol rapa whelks usetl m tliis study with basic statistics on ratlulae. Abbre\iations usetl below arc as follows: 
F = female, M = male, Avg = average for female and male whelks combined, SL = shell length, mm, SEM = standard eri'or of the 
mean in parentheses. RL = radvila length, mm. RW = radula wiflth, mm; NRT = number of rows of teeth. 

# of Whelks 

Avg SL 

A\g RL 

Avg RW 

Avg NRT 

.'\\g RIVSL % 

Whelk size class 







Petite (<45 mm SL) 

V = 5 

25.64 (2.64) 

5.28 (0..56) 

0.48 (0.18) 

106 (6.07) 

20.59 (0.58) 

M = 5 

34.72 (3.07) 

7.62 (0.95) 

0.71 (0.31) 

122.20 (4.78) 


Small (45.L-90mm SL) 

F = 5 

70.38 (5.47) 


0.97 (0.15) 

133.80 (5.05) 


M = 6 

69.13 (6.21) 

17.63 (1.92) 


126.67 (3.86) 

25.56 (1.79) 

Medium (9().l-]:35 mm SL) 

F = 4 

104.75 (5.07) 

22..32 (2.52) 

1.47 (0.03) 


21 .37 (2.53) 

M =5 





25 (1.51) 

Large (L35. 1-180 mm SL) 

F = 2 

153 (10.0) 

35.7 (3.2) 



23.3 (0.57) 

M = 7 

153.71 (4.47) 

41.12 (2.23) 


182.29 (6.38) 


M. Hardine et al, 2008 

Page 221 

■ Females 
□ Males 

11 IMill rr I III! B 

n = 10 

n = 11 

11-^ ' ,r\ .i-^ it^ ' irt ' I 

n = 9 

n = 9 

. 1 I I M I I I I I 

Midpoint of rapa whelk shell length size class (mm) 

Figure 3. Shell length (mm) trequenc\ distfibutiun for the 
39 veined rapa whelks whose radulae were examined in this 

(females = 0.78 vs. 0.71, males = 0.81 vs. 0.72, Table 2) 
and predicts a positi\e number of tooth rows at radula 
lengths of mm (Figure 6E). 

Rachidian Tooth Dimensions: Raehidian tooth cen- 
tral cusp base width (LI, mm; Figure 4) increased 
siguiticantK' uith increasing whelk size class (Table 3, 
Figures 7A and 7D). There were no significant differ- 
ences in central cusp base width obseived between rows 
witliin a size class for die 81 rows of teeth that were 
e.xamined. Central cusp base widdi was significantly 
larger in males than in females (Fisher's test. Table 3). 
Differences between male and female central cusp base 
width were particularK- evident in the medium and large 
size classes (Figure 7A and 7D). 

The height of die rachidian toodi central cusp (L2, 
mm) varied significanth' with size class, row number, 
and sex (ANO\'A, Table 3, Figures 7B and 7E). In gen- 
eral, larger whelks have larger central cusp heights in 
rows 31 through 81 than whelks of odier size classes. 
Within all size classes and bodi sexes, central cusp 
heights from rows 1 and 1 1 are significantly less than in 
rows 31 dirough 81 (ANOXA., Table 3, Figures 7B 
and 7E). Central cusp heights from rows 1, 11 and 21 in 
large whelks are significantlv different from central cusp 
heights in petite whelks but similar to cusp heights ob- 
ser\"ed in rows 1 and 21 for medium whelks and row 11 
for small whelks (ANOVA, Table 3, Figures 7B and 7E). 
Female whefks have significantly lower L2 values than 
male whelks (Fisher's test. Table 3, Figures 7B and 7E). 

Rachidian tooth base width (L3, mm) increases signi- 
ticantK' with increasing whelk size class (ANOX'A, 

































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Page 222 

THE NAUTILUS, Vol. 122, No. 4 



—»— Females 

- n- Males 



a ,1 

° .'-"■' / 



dj' y 



^^' y^ 







•■• /• ° 

















1 ■ ■ ■ ■ 1 ■ ■ ■ ■ 1 

3 ,. 

50 100 150 

Shell length (mm) 














A °X 



y c 





° p ^ y^ 
D y *X* 



1 J 

1 Jf 





. . . 1 . 

■ ■ ■ 1 ■ ■ 

1 1 1 1 . 1 1 1 



J ^^ 



.o /-a 




a -^ -/ 






1 5 

D A 

















1 1 


.. 1 .... 1 .... 1 .... 1 











50 100 150 

Shell length (mm) 




°% y' 

u _■ / 

D yy 

^' y 

n . ' ']y° 

■ yy* 

» .'V^ 

. y° 

o . a ,'':;>^ 


a ' V^ ° ° 

■ '^>^ °« 


^y • 

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" ■ ■ ■ ■ 1 ■ ■ ■ ■ 1 ■ ■ ■ ■ 1 ■ ■ ■ ■ 1 


10 20 30 40 50 

Radula length (mm) 


50 100 150 

Shell length (mm) 


10 20 30 40 50 

Radula length (mm) 

J. M. Harding et al.. 2008 

Table 2B. Suminan' iif t-tests comparing regression I'tpia- 
tions lor female and male wlielks gi\en in Table 2A tliat are 
recommended for descriptions of these relationships. T-tests 
were performed tor the power model nsing logarithm 
transformed data. Abbreviations are the same as those nsed 
in Table 2B abcne. 







Female \ s. 




SL vs. 


Femiile vs. 






Female vs. 




RL vs. 


Female \s. 






Female \s. 




Table 3. Figures 7C and 7F). However, no differences 
were obsened in rachidian tootJi base widths bet^veen 
radnlar tooth row's within a whelk size class and within a 
sex (ANO\'A, Table 3, Figures 7C and 7F). Rachidian 
tooth base width was simificantK' lars;er in male whelks 
than in female whelks and this trend is particularly evi- 
dent in die medium and large size classes (Table 3, 
Figures 7C and 7F). 

Large whelks had significantlv higher ratios of rachi- 
dian central cusp base widtli (LI) to central cusp height 
(L2) tlian all other whelk size classes (ANOVA, Table" 3, 
Figure 8). The first row of teeth in the radulae had 
significantK' higher LLL2 ratios tlian all other rows 
(ANOVA, Table 3, Figure 8). The eleventh row of teeth 
also had an L1:L2 ratio tliat was significantly higher than 
that observed in rows 21-81 (ANOVA, Table 3, Figure 8). 
Since central cusp base to height (L1:L2) ratios >1 are 
indicative of toodi wear, die anterior 1-11 rows of teeth 
are more worn dian newer teeth occurring in rows 21 
and higher. W'idiin each size class and sex, a wide range 
of L1:L2 \'alues was obsen'ed for row 1 and/or row 11 
(Figure 8). This variabilitv was the result of one or two 
individual whelks per size class having ven' low central 
cusp heights (extreme wear) in row 1 or row 11, the rows 
of teedi that are activek' used in feeding. Patterns of wear 
as indicated bv L1:L2 ratio values were not significantly 
different betxveen sexes (Table 3, Figure 8). 

The ratio of central cusp base width to rachidian base 
width (L1:L3) was significantly affected by size class and 
sex (ANOVA, Table 3, Figure 9). Within a size class and 
widiin a sex, the ratio of central cusp base widtli to 
rachidian toodi width did not change significantly with 
row number. Male whelks had greater L1:L3 ratios than 
female whelks (Fisher's test. Table 3, Figure 9). 


Radula lengdi. radula width, and number of rows of 
teeth in the radula increase with increasing rapa whelk 

shell length. Ontogenetic increases in radula length and 
the number of I'ows of teeth with sliell length have 
also been documented for other muricid species (e.g., 
Stramonita floridrnui. Radwin and Wells, 1968 (as Thais 
floridand): CrouUi indfiuiiilii-itla and Monila imisiva, 
Fujioka, 1984; Titdis liroimi and T. clavigcra, Fujioka, 
1985: Niurlla lapilhis, Kool, 1993). 

The relationship between rachidian toodi base widdi 
and central cusp base width also changes widi ontogeny 
but does not change in relation to the anterior-posterior 
location on the radula. Tliat is, within an individual and 
within a size class, lachidian teedi examined from rows 1 
through 81 displav similar scaling of centraf cusp base 
widdi to rachidian toodi base widdi. Rachidian teedi in 
female rapa wlielks tend to have smaller ceiitr;il cusp 
liase width as well as toodi base widths when compared 
to male whefks within the same size class. Fujioka (1982, 
1984) describes similar ontogenetic changes in rachidian 
tootii shape incluiling an increase in central cusp base 
width for Cronia iiuirgaiiticola. Morula miisiva, and Dni- 
pclla sp. in relation to sexual dimoi-phism. The observed 
ontogenetic changes in rachidian toodi shape may reflect 
moiphological shifts designed to accommodate greater 
racludian tooth strike force resulting from the scaling of 
the buccal complex at increased whelk sizes. Pi-esumably 
there is an ontogenetic scaling relationsliip in effect to 
optimize the force provided by die buccal mass muscula- 
ture and minimize the damage to rachidian teeth through 
use that is reflected in the shape of the tootii. 

Rachidian tootii wear, as indicated liv the ratio of 
central cusp width (LI) to central cusp height (L2), 
decreases with increasing distance Irom the anterior 
(oldest) end of the radula. Rachidian teeth in the first 
11 rows of the radula have central cusp heights that are 
less than central cusp heights in rows 21 through 81 in 
all size classes. Caniker et al. (1974) describe rachidian 
cusps tliat have been worn off leaving only the tooth 
base in the anterior rows ol rachidian teeth of Urosal- 
pinx cincira foUci/ensis. In laboratoi'v studies with Tlwis 
bronni and T. cJamgera, Fujioka (1985) obsened that 
rows of teeth worn by feeding occupy 5-15% of die total 
number of radular rows. The whelk Acanfhina spirata 
uses approximately 8 to 20 teeth in each rasping stroke 
as these whelks feed on mussels (Hemingway, 1975). 
These data are consistent with our obsenations for rapa 
whelks, where at least the 1st and 1 Itli rows of the radula 
were used, and the 21st row acted as a transition between 
the part of the radula the whelk was activelv using to feed 
and the more posterior section tiiat was unused. 

The observed changes in rachidian tootii wear may 
reflect ontogenetic changes in predation strategy, diet, 
or possiblv both. Differences in predation strategv' 
are potentiallv reflected in the obsened changes in 

Figure 6. Relationships for female (n = 16) and male (n = 2.3) veined rapa whelks ranging from 20.2 to 174 mm SL between shell 
length I SL ' and radula length ( RL, A), SL and radula width (B), SL and the number of rows of teeth (C), RL and radula width (D), 
and RL and number of row's of teeth (E) with fitted regressions (female = solid, male = dashed) that were used to describe the 
relationships. Linear regression models are plotted for panels A, C, and E. Power models are presented in panels B and D. 
Regression equations and descriptive statistics are given in Table 2. 

Page 224 

THE NAUTILUS, Vol. 122, No. 4 


0,35 : 

0) 03 


TO ^ 

■° ^ 25 

<f> CO 

^' 02 

1 ^ 

c . 










I Petite female (n = 5) « Medium female (n = 4) 04 
t Small female (n = 5) ■ Large female (n = 2) 


i 1 n 

\ 1 * 

\ \ \ I \ \ \ "^ "^ 

T 1 I 

I I r- 


{/) "^ 

D LU 25 
O C/3 

"to c 

i: P 02 

0^:1 15 

D5 ^^ 





J 1 1 f 

! f 


I I T I I 1 I 





ro ^-' 







1 1 

* « 

1 1 

» t « 

1 H 1 M M 1 

t « 

ol— f 

1 . . — I . 1 . 1 1 r 

20 40 

4- r- 

■ t 


60 80 

Rachidian tooth row number 










Petite males (n = 5) «. Medium males (n = 5) 
D. » Small males (n = 6) n Large males (n = 7) 

1 1 h H n [ 

* f ^ ^ i { 5 f ^ 


{ \ \ \ { i I I I 

—I — I — I t — I I I — I I I I — I I I — I — I I 


} t 

1 I f } 

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—I 1 1 I 1 1 I 1 I I 1 I r 1 1 I 

1 z 








= I 



f t 




E ^ 







[ } 



[ 1 













[ ] 











Rachidian tooth row number 

Figure 7. Graphs of rachidian tooth row iiiiiiil)('r in relation to average central i iis|) \M<lth ( LI ) with error bans (standard error ot 
the mean, .SEM) for females (A) and males (D) Irom all size classes, average central cusp height {L2) with error bars (standard error 
of the mean, SEM) loi- females (B) and males (E) from all size classes, and average rachidian tooth base width (L3, SEM) for 
females ((.)) and males (I"; from all size classes. 

M. Hardine et al.. 2008 

Page 225 

Table 3. Suminan' of ANOVA results comparing tooth 
inoipliologs' across whelk size classes and rows within the 
radnlae. Asterisks indicate statistical significance at an alpha 
\alue ot 0.05. .■Xhhrexiations used below are as follows: 1= petite 
size class, 2 = small size class, 3 = medium size class, 4 = large 
size class. NA = Not applicable. Refer to Figru'e 4 for a 
description of LI, L2, and L3. 

Fisher's test 




p \ alue 



Central cusp 

Size class 


4> 3 >2 >1 







width (LI) 



Male > Female 



Central cusp 

Size class 


4, 3 >2 >1 


height (L2) 



1, 11 < 31-81 




Male > Female 




Size class 


4>3 >2> 1 


tooth base 





width (I^) 



Male > Female 




Size class 


4 > 3, 2, 4 = 1 


(ratio L1/L2) 



1 > 21-81; 








Shape (ratio 

Size class 


4 > 2, 3 > 1 









Male > Female 

rachidian tooth wear with ontogeny because the rachi- 
dian teetli are activelv used during shell drilling. Meth- 
ods of feeding which require penetration of prey valve 
shells with the radula (e.g., drilling) will leave more wear 
on the rachidian teeth than non-drilling methods of at- 
tack. Theretore, examination of rachidian tooth wear 
along the radula and differences in wear depending on 
size class may give an indication as to transitions in 
feeding strategies of different size classes of rapa whelks 
(Figure 8). High levels of rachidian toodi wear (L1:L2 
ratio >1) in rows 1 and 11 were associated with the 
petite and large size classes (Figure 8A). The smallest 
whelks (<45 mm SL) typically leave drill holes (i.e., wall 
bores) in die valves of their prey (Harding et al, 2007). 
.\lthough large (>135 mm SL) rapa whelks do not 
always lea\'e predation signatures in prey valves, edge 
bore signatures are left instead of drill holes (Harding, 
Kingslev-Smith, Mann, unpublished data) when signa- 
tures are present. The obsen'ed L1:L2 values for rows 
1 and 11 in the large size class are driven by one male 
(Figure 8B). It is possible that diis indi\idual had been 
using its radula to penetrate prey shells and that the odier 
large whelks had not. Since die large whelks used herein 
were wild caught and had unbiown feeding histories, we 
cannot sav this with certaint)'. Relatively less wear (L1:L2 
ratio <1.2 ) in row 11 was observed in rapa whelks with 
shell lengdis of 45 to 135 mm (small and medium size 
classes) that do not t)picallv drill their prey (Harding 
et al., 2007, Figure S). Similar wear patterns (L1:L2 ratio) 
were obserxed for males and females and may serve as an 
indication that feeding strategies and/or prey may be simi- 
lar between animals of different sex but similar size. 

5 - 

3 - 

A. Females 

• Petite (n = 5) 
A Small (n = 5) 

♦ Medium (n = 4) 
■ Large (n = 2) 

M ! ! f I I 

-I — I — I — I — I — I — I — I — I — I — I — I — I — I — I — I — I 

£ 6 



Petite (n = 5) 

i Small (n = 6) 

o Medium (n = 5) 

° Large (n = 7) 


40 60 

Rachidian tooth row number 


Figure 8. Rachidian tooth wear as indicated by the average 
ratio ot rachidian tooth central cusp base width (LI) to central 
cusp height (L2) in relation to rachidian tooth row number for 
females (A) and males (B) from all size classes. 

It a wlielk is using tiie radula to penetrate the shell, 
the chemical composition of the prey shell may affect 
the level of wear observed on the rachidian teeth. Oyster 
and mussel shells have more calcite and are relatively 
softer than other bivalves with predominantK' aragonite 
shells (Carter, 1980). Drilling through aragonite shells 
has the potential to cause more wear on rachidian teedi 
than shell penetration of calcite shells. Examination of 
the radtilae from rapa whelks using drilling to consume a 
species-speciiic diet is a topic for future research that 
would provide data to address changes in radula wear 
with regard to prey shell hardness. 

Page 226 

THE NAUTILUS, Vol. 122, No. 4 






A. Females 

I I 

• Petite (n = 5) 
A Small (n = 5) 

♦ Medium (n = 4) 
■ Large (n = 2) 


1 { 

0.15 I — I — I — I — I — I — I — I — I — I — I — I — I — I — I — I — I- 








i i 

U I! 

! " " 

Petite (n = 5) 
Small (n = 6) 
Medium (n = 5) 
Large (n = 7) 

-4— ^- 

20 40 60 

Rachidian tootti row number 


Figure 9. Average ratio ot central cusp base width (LI) to 
rachidian tooth base width (L3, vxath standard error of tlie 
mean, SEM) by rachidian tooth row for females (A) and males 
(B) from all size classes. 


This manuscript is dedicated to the late Dr. Melbourne 
R. Caj-riker, whose passion foi- and detailed work with 
Min-icids sei-ves as both a standard of excellence and 
an inspiration. Thanks are extended to all local fisher- 
men that have participated in the X'irginia Institute of 
Marine Science rapa whelk bountA- program since its 
inception in 1998. Melissa Southwordi, Ethan Jestel, 
Catherine Ware, Erica Westcott, Steven Goodbred, 
David Kerstetter, Peter Kingsley-Smith, Amy Bohannon, 
Meredith Fagan, Meghan Harris. Rhonda Howlett, 
Rae Marie Johnson, Comtney Harris, Karen Capossella, 
John Hansen, and Matthew Robinson assisted with 

whelk collections from local fishermen. Drs. Greg 
Capelli, Randv Chambers, John Kraeuter, Ms. Mehssa 
Southworth, and an anon)'mous reviewer provided valu- 
able comments on earlier versions of this manuscript. 
This work was completed in partial fulfillment of the 
refjuirements tor an undergraduate Honors degree 
(SMG) from the Department of Biology, College of 
William and Maiy, Williamsburg, Virginia. This is contri- 
bution number 292S from the Virginia Institute of 
Marine Science, Gloucester Point, Virginia. 


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Carriker, M. R. 1961. Comparative functional morphology of 
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Carriker, M. R. 1969. Exca\ation ol boreholes b\ the sastro- 
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Carriker, M. R., J. G. Schaadt. and V. Peters. 1974. Analysis by 
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T. M, Hardin2;etal.,2(K)S 

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New Jersey. 

THE NAUTILUS 122(4):228-235, 2008 

Page 228 

A new species of Sphaerium Scopoli, 1777, from southern Brazil 
Bivalvia: Sphaeriidae^ 

Maria Cristina Dreher Mansur 

Av. Arlindo Pasqualini, 410 
91760-140 Porto Alegre, BRAZIL 

Claus Meier-Brook 

Sommergasse 10 

D-72119 Ammerbuch-Reusten 


Cristian Ituarte 

Museo Argentine de Ciencias Naturales 

Av. Angel Gallardo 470, C1405DJR, Buenos 




Sphaerium cambaraense new species is described based on 
samples collected in the beginning of tlae summer near the 
headwaters of Taquari River (Jacui River Basin) on the basaltic 
plateau of southern Brazil. This is tlie first record for die genus 
Sphaerium in Brazil and in South America outside the Andes. 
Sphaerium cambaraense is characterized by a relatively large 
and solid shell, a high triangular shell oudine and a solid hinge 
plate. Compared with Sphaerium forbesi (Pliilippi, 1869), 
S. cambaraense has a more strongly triangular shell outline 
and beaks not prominent. Sphaerium lauricochae (Philippi, 
1869), another similar species from Bolixda, Chile, and Peru, 
has a more rounded shell outline. 

Additional Keijicords: Freshwater, Rio Grande do Sul, South 


According to Dreher-Mansur and Meier-Brook (2000), 
the family Sphaeriidae is represented by two subfamilies: 
Euperinae, inchading tlie genera Eiipera Bourguignat, 
1854, and Bijssanodonta d'Orbigny, 1846; and Sph- 
aeriinae with diree genera, Sphaerium ScopoU, 1777, 
Musculium Link, 1807, and Pisidium Pfeiffer, 1821. 

Four species of tlie genus Sphaerium are known from 
South America: Sphaerium aequatoriale Clessin, 1879, 
from Ecuador (Kuiper and Hinz, 1984); S. forbesi (Phi- 
lippi, 1869) recorded from Peru and Bolivia (Haas, 
1949), from Bolivia (Haas, 1955), from Colombia, Peru, 
Bolivian Andes (Kuiper and Hinz, 1984), and Chile 
(Ituarte, 1995); S. lauricochae (Philippi, 1869), from 
PeiTj, Bolivia and Chile (Kuiper and Hinz, 1984), and 
Chile (Ituarte, 1995); S. titicacense (Pilsbiy, 1924), from 
Peru and Bolivia (Kuiper and Hinz, 1984). According to 
these authors, they are concentrated in the Central 
Andes from Ecuador to North Chile including high- 
altitude Bolivia, at 2000 to 4700 m altitude; varying in 
size between 3 and 12.5 min. Haas (1949) reports 

Sphaerium (S.) (Sturany, 1900), from the 
highlands of Bolivia and from a lake in Junin, Peru. He 
examined also material of diis species collected by Sioli 
in the regions of the rivers Maue-Agu and Tapajos, 
tributaries of the Amazon River, which constitutes the 
first record of Sphaerium for Brazil. According to Kui- 
per and Hinz (1984), S. bohviense is a junior s)iionym of 
S. forbesi. 

The record oi Sphaerium obscivationis by Mansur 
et al. (1991) for Mirim Lagoon in southern Brazil is a 
misidentification, as that species is not a Sphaerium. 
According to the revision bv Ituarte (1995), Pisidium 
observationis Pilsbry, 1911. onK occurs in the southern 
Argentina, not in Brazil. 

Sphaerium cambaraense new species is the fifdi 
Sphaerium species from South America and the first spe- 
cies of die genus described from southern Brazil, in the 
highlands near the Adantic Oceair, a location geographi- 
cally verv distant from die Andes and Amazon River 


Specimens were collected with a plastic sieve with mesh 
size of about 0.8 mm. Specimens were sorted from the 
sediment widr fine feather tweezers; anesthetized in 
small vials containing water witir mendiol crystals, fixed 
in a 5% formalin solution for 24 hours, rinsed for 24 hours 
in tap water and presei-ved in 70% ethanol. Soft parts of 
specimens for scanning electron microscopy (SEM) were 
removed with tweezers, shells cleaned with a soft and 
fine brush and rinsed several times in distilled water 
Dried shells were glued on stubs with light-silver glue 
(Porolon Equipment, Herts) or metallic adliesive tape 
(T066 Silver tape 9 mm, Hert-Scotch), coated widi gold, 
and obsei'ved either in a Cambridge Stereoscan 250 Mk2 
or Philips scanning electron microscopes. Shell micro- 
structure was studied by fracturing shells at the middle 
of die height, parallel to commarginal ridges. Terminolo- 
gy for shell microstructure follows Dyduch-Falniowska 
(1983) and Dreher-Mansur and Meier-Brook (2000). 

M. C. D. Mansur et al. 20()S 

Page 229 

Stonuifli nomenclatuic lollows I'urclion (1958, 1960). 
The shape indices, height index [I = H/L] and convexity 
index [Ci = W/H], were calculated according to the ci"i- 
teria followed by Ituarte (1996). 

.•\bbre\iations use in the text are: MACN, Museo 
Argentiuo de Ciencias Naturales, Buenos Aires; MCN, 
Museu de Ciencias Naturals, Fundayao Zoobotanica do 
Rio Grande do Sul, Porto Alegre; MCP, Museu de Cien- 
cias e Tecnologia da Pontificia Universidade Catolica, 
Rio Grande do Sul, Porto .-\legre, Brazil. 

Genus Sphacriutn Scopoli, 1777 

Spliaeiiiim cambaraensc new species 
(Figiu-es 1-20) 


Distinguished b\' the relatixelv large and 
solid shell with trapezoidal tending to triangular shell 
outline, low and wide beaks, subcentrally located, without 
marked nepionic cap, and broad and solid hinge plate. 

Description: Shell: Solid, relatixely large (maximum 
obsencd L: 11.22 mm), slightl)' convex (Ci = 53±5). 
Shell oudine high (I = 85±2), trapezoidal tending to 
tiiangidar Dorsal margin has pronounced cui-ve. Anteri- 
or and posterior margins gradually descending and 

gently curved below middle of total height, without 
marked angle. Posterior end slightly truncated, oblique 
(Figures 1, 2). Ventral margin long and evenly cui-ved. 
Shell surface silky, glossy, witli very fine irregularly 
distributed radial lines, weaker on beaks and irregular 
conimarginal, sometimes coarse, striae (12 or more per 
0.5 mm in the middle of the shell (Figure 3). Outer shell 
surface light yellowish brown; pale browni at beaks, more 
ventrally grayish brown with complete or incomplete 
yellow concentric bands rtmning from anterior to poste- 
rior margins; a large yellow band near ventral margin. 
Beaks without marked embryonic cap. Inner shell sur- 
face white, grayish at muscle scars. Beaks slightly proso- 
gyrous, low and wide, shghtly raised above dorsal 
margin, subcentrally located (Figures 1, 2, 6, 7). 

Hinge plate strikingly cuwed, solid, broad, reaching 
0.5 mm width in middle region in specimens of 11 mm 
length, slightK' narrower at level of cardinal teeth. Hinge 
line arched, paiticularly below cardinal teeth (Figm'cs 6-9). 
Cardinal teedi strong, close to dorsal margin. Right 
cai"dinal toodi, C3, short, strongly cui-ved, posterior end 
enlarged in a grooved cup (Figure 10). Left cardinal teeth: 
outer cardinal tooth, C4, diin, strikingly oblique, located 
immediately behind C2, anterior end slightly overlapping 

Figures 1-5. Sphaeriiim cambaraense new species. 1, 2. Holotype MCP Mol. Outer view of left and right valves. 3. Detail of 
outer shell surface. 4, 5. Inner shell surface, detail of pores. Scale bars: 1, 2 = 4 mm; .3 = 400 |.uii; 4 = 4 (im; 5 = 40 i^m. 

Page 230 

THE NAUTILUS, Vol. 122, No. 4 

Figures 6-11. Spliewnuui cambaracnse new species. Paiatype (MACN-ln .37()(i3). 6. Inner N-iew ot right valve. 7. Inner view of 
right valve. 8. Hinge of right valve. 9. Hinge of left valve. 10. Detail of right cardinal tooth (C3) and ligament. 11. Detail of left 
cardinal teeth (C2 and C4) and ligament. Scale bars: 6, 7 = 2 mm; 8, 9 =1 nnii: 10, 11 = 500 |im. 

C2; inner cardinal tootli, C2, short, high, columnar, deeply 
arched into a V-shape (Figure 11). Right lateral teetli 
.somewhat short, strong, with distal cusps (Figure 8); left 
lateral teeth relatively long, strong, and high (Figure 9). 
Ligament internal but exteriorly visible, slightly protrud- 
ed in larger .specimens (Figures 10, 11). 

Shell Microstructure: Inner shell surface perforated 
by numerous pores (55/400 nm" to 1.3/160 nm") (Figiu'es 
4, 5) representing the opening of tubuli that cross entire 
calcai'eous pait of shell (showai in part in Figiu'e 12). 
Openings of pores on inner surlaee surrounded by a 
funnel-shaped depression and m nionlli circled by a rini 

M. C. D. Mansiir ct ;il., 2(K).S 

Pase 231 







Figure 12. Sphucnuiu ciimbarafiisc new species. Shell iiiicrustructure iruiii a: penostracuiii (topi to t': the eiitlostracuiii (botloiii,) 
Inserts a-g are details of: a, periostracum; b, granular layer; c, diagonal layer forming a composite prismatic stnicture; d, diagonal 
la\er. crossed structure: e, diagonal layer forming a pseudo crossed lamellar stnictiire; f, palisade structure; g, diagonal layer and 
internal surface of the endostracum (arrow). Scale bars: 12 = 50 |im; inserts: a-d = 4 |im; e = 20 \im; 1 = 10 |jm; g = 2 |im. 

Pa2;e 232 

THE NAUTILUS, Vol. 122, No. 4 

M. C. D. Mansur et al., 200S 

Pase 233 

(Figure 4). Shell relatively thick reaching 200 in 
central area (Figin'e 12). Shell structure consists ol' 
fne la\'ers: periostracum, granular laver, diagonal layer, 
palisade, diagonal la\er, and endostracuni. Below the 
2 pni thick periostracum (Figure 12, a), a granular 10 |.ini 
thick laxer appears (Figiu'e 12, b); grains concentrated 
in some points irregidarK' and sparseh' distributed, look- 
ing like cone-shaped bars arranged pei^pendicularly to 
shell surface. These change into irregularly and com- 
pactK' arranged grains of different sizes. More internally, 
shell structure changes gradualK' into a diagonal layer 
(Figiu'e 12, c, d, e), where three different patterns may 
be distinguished: first (about 50 |am thick), composite 
prismatic structure, showing vertical feather-like pattern 
(Figin-e 12, c); second (about 40 |im thick), occupies 
die central part of shell, gradually changing into a 
cross-lamellar stioictnre (Figure 12, d); the third (appro.x- 
imateh' 60 (im), resembles a pseudo crossed-lamellar 
stiTicture (Figure 12, e). Close to tlie inner shell surface 
diere are one or tvvo palisade layers, narrow (each about 
1 |,uu thick) (Figure 12, f). The palisade structure is 
followed bv a diagonal layer with lamellae oriented 
in one direction (Figure 12, g): below, a veiy narrow 
la\-er, the endostracuni (1.0 to 1..5 \.im thick) (Figure 12, 
arrow 1. 

Ax.woMV (Figures 13-20): Inner and outer demi- 
branchs well-developed, outer smaller demibranch 
reacliing half of height of inner demibranch (Figure 13). 
Brood sacs occupy anterodorsal part of inner denii- 
branchs, embnos contained in sacs showed different 
dex-elopmental stages, denoting sequential brooding. 
Up to six developing embryos found in largest examined 
specimen (>11 mm lengtli). Largest, tertiaiy, brood sac 
contained two embi-yos (>1 mm length) (Figure 13); 
secondaiy sac located under largest one, contained three 
or four small embryos. Primary sac attached lower on 
inner demibranch. (Specimens for this study were col- 
lected at the beginning of tlie Southern Hemisphere 
summer (Januan.-, 1994) and many of the specimens 
larger than 9 mm showed brood sacs.) Anal and branchi- 
al openings extended in diverging short siphons, nearly 
equal in size. Anal siphon wider at base and more stre- 
tched out (Figure 13). Two labial palps (Figure 13, 14) 
on each side of mouth, triangular-shaped in lateral view: 
opposite contacting walls with 12 small folds, tapering 
toward distal ends. Mantle musculature (Figure 15) with 
relatix'elv short siphonal retractors; inner radial mantle 
muscles arranged in eight to nine bundles (Figure 15). 
Fan-shaped stomach (Figures 17, 18) bent to right 
side, laterally covered bv digestive gland. Dorsal hood 
relativeh' short and left duct well developed. Stomach 
intemalK- (Figure 17) shows \'er\" simplified structures 

on dorsal hood and right side such as short gastric shield 
under dorsal hood: minor txqDhlosole and rejecting tract 
beginning at right side after descending from dorsal 
hood; and an ele\ation slightly wrinkled between intesti- 
nal groove and rejection tract; anterior fold absent. In- 
testine opening associated to style sac in center of floor; 
major tyj^hlosole arched in front with two expansions 
that end respectively at left and right duct openings, not 
penetrating in ducts, which allocates the stomach to 
Tx-pe IV; left and right duct openings well developed 
and ramified in three secondaiv ducts. 

Intestine short and simple (Figiu'e IS), anterior part 
associated to style sac, broad, descending straight to 
floor of visceral mass; mid-intestine strikingly stretched 
forming single loop; hindgut straight, ascending and 
bending towaril rectum that ends into an anal papilla. 
Nephridia of closed t\'pe; dorsal and outer lobes fused, 
impossible to distinguish in dorsal view (Figures 19, 20); 
external wall of outer lobe ornamented widi many small, 
rounded extrusions; nephridia in young specimens usu- 
ally of open i\rpe and v\dth separated lobes. Fimnel rela- 
tively long and wide; pro.ximal loop with smooth walls 
forming three ascending rings then run backward up to 
posterior adductor muscle, turning laterally in direction 
to lateral k)op. Lateral loop straight, elongated, running 
along outer side and partially covei'ed laterally b)' outer 
lobe. Excretoiy sac sub-triangular, romided in front and 
relatively small. No valve at the entering of the distal 
loop into excretoiy sac. 

Type Locality: Lajeado da Margarida (50'' 15.75' W, 
29"0.87' S; 870-SSO m altitude) on'Camisa River, Antas 
River Basin, considered to be the headwaters of Taquari 
River, tlie main tributaiy of Jacui River, South Atlantic 
Brazilian Basin. 

Type Material: Holotype MCN 38821; Paratypes: 
MCN 33919 (33 specimens), MCP 9109 (6 specimens), 
MACN-In 37063 (two specimens). 12 Jan. 1994. 

Etymology: The specific epithet refers to the Cit)' and 
Municipality of Cambara, close to the type locality. 

Distribution and Habitat: Known only from the type 
localit\'. The Miniicipalit\ ol Cambara is situated in 
northeastern Rio Grande do Sul State, Brazil, in the 
highest part named Planalto Riograndense. From the 
physiographic point of view, this region is characterized 
by a basaltic shield covered by low grass steppes and 
Araucaria Forest mixed with the southeastern limits 
of the Atlantic Forest. The altitude varies from 850 to 
1050 m, and in the winter, temperatures fall below 
freezing and there is the occasional snow. The rivers diat 
cross the region, flanked bv a low galleiy forest, ha\'e 

Figures 13-20. Sphacrium cambaraensc new species. Sclieniatic drawings olsoit anatomy. 13. Gross anatomy (left mantle lobe 
remo\ed '. 14. Folded surfaces of inner and outer left labia! palps. 15. Outer view of left mantle lobe showing the inner radial manrie 
muscfes. 16. Dorsal \iew of the stomach and digestive gland. 17. Floor of die stomach after removing the roof 18. Inner view of die 
organs in the \iscerat mass. 19. Dorsal view of nephridia. posterior adductor muscle and posterior foot retractors. 20. Lateral view of 
left nephridium. Scale bars: 13. 1.5, 16, 18 = 2 cm: 14, 17. 19, 20 = 2 mm. 

Page 234 

THE NAUTILUS, Vol. 122, No. 4 

hard bottoms formed by flattened basaltic stones, and 
currents are strong. The collecting sites were small 
ponds along the river course, where currents were low- 
er, allowing the accumulation of decayed leaves and very 
soft, dark, and fine sand deposits, not deeper than 1 m, 
where specimens settle. Together with the Sphacrium 
samples, many specimens oi one species oi Pisicliuiii sp. 
(MCN 33918), and one of Diploclon sp. (MCN 33920) 
not yet identified, were found. 


Sphacriinn cambaraense is similar to Spliaciiiun forbesi 
(Philippi, 1869) (from Bolivia, Cliile, and Peru). However, 
S. cambaraense has a more decidedly triangular shell 
outline, beaks not full with not marked nepiomc shell; 
in addition. S. cambaraense is larger than S. forbesi. 
Sphaeriinn Iniiricocliac Philippi, 1869), also reported from 
Bolivia, Chile, and Peru, diiiers from S. cambaraense 
by its more rounded shell outline. In relation to shell 
thickness and microsti-uctm'e, S. cambaraense is similar to 
the European species Sphaerium rivicola (Lamarck, 
1818) and Sphaerium cornenm (Linnaeus, 1758); 
Dvduch-Falniowska (1983) reported for these species (as 
well as for Miiseiiliitm lacusfre (Miiller, 1774) and several 
Pisidium species) six different layers, (1) periostracum, (2) 
homogeneous-granular layer, (3) granular layer; (4) diago- 
nal layer (composite prismatic sti^ucture), (5) palisade 
structiu'e, and (6) endo.stracuni. Onlv S. cornenm and 
S. livicola showed a different strvicture tcji' the diagonal 
layer, referred to as "crossetl-lamellar structure" (Dyduch- 
Falniowska, 1983). This structiu'e was also found in 
S. cambaraense. howexer. the diagonal pattern of the 
plates has a different arrangement: in S. rivicola die 
oblique plates show a horizontal herringbone pattern and 
in S. cambaraense some rows of plates are oblique not 
forming a horizont;il pattei'n. These latter resemble in part 
the pseudo crossed-lamellar structiu-e found in the Corbi- 
culidae. The periostracum la\er in ,S'. cambaraense is thin- 
ner dian in S. conienm and the homogenecjus granular 
layer is lacking. The simplified condition of posterior and 
right side of stomach is quite similar to tliat (oimd in 
the Euperinae (Dreher-Mansur and Meier-Brook. 2000). 

The nephridium is relatixelv similar compared to 
S. cornenm (Dreher-Mansur and Meier-Brook, 2000), 
but the excretoiy sac and proximal loop are sliorter and 
the valve at the insertion of the distal loop into excretoiy 
sac is lacking in S. ccmibaraensis; the multilobed surface 
of lateral lobe is observed for the first time in Sphaerium. 

Based on the similar morpliology of the nephridium, 
Korniushin (1998) proposed that South American spe- 
cies traditionally assigned to Sphaerium actually be- 
long to Musculium. According to Park and O Foighil 
(2000), the usefulness of the fine anatomy of nephridi- 
um in the family Sphaeriidae is relative, due to its high 
morphological plasticity. Coolev and O Foigliil (2000), 
based on mitochondrial I6S r DNA gene s('(|uences 
observed thai die Spltaerinm/MitscuUum clade exhibit 
moderate to low Icvc^ls of genetic divergences and 

the same asynchronous or sequential brooding pattern 
(i.e., brooding sacs contain more than one develop- 
ing generations of embiyos). Nevertheless, Coolev and 
O Foighil (2000) recognized Musculium as a monophy- 
letic group. We allocate die new species in Sphaerium 
until more e\idence is available tf) help v\ddi diis ques- 
tion. More recently, a phylogenetic analysis of the 
Sphaeriinae (Lee and O Foighil, 2003) based on a molec- 
ular study of nuclear (ITS-1) and mitochondrial (IBS) 
gene sequences of 15 species from North and South 
America, Europe, Asia, and Australia, reco\ered a 
strongly suppoited monophvletic group of sequential 
brooders [Musculium and Sphaerium). However, die 
analyses indicate that Musculium and Sphaerium sensu 
lato are not natural groups, proposing a new classification 
system comprising five subgenera within Sphaerium: 
Sphaerium sensu stricto Scopoh, 1777; Musculium 
Link, 1807; Amesoda Rafinesque, 1820; Sphaerinova 
Iredale, 1943, and Herringtonium Clarke, 1973. As these 
subgeneric groupings, however well supported by 
molecular data, are not defined from the moiphological 
point of view, it is not possible at this point to place 
S. cambaraense within the new scheme. 


We are grateful to the curators of VICN mollusk 
collection: Ingrid Heydrich and Silvia D. Hahn for 
pro\iding the loan of specimens; to Prof Dr. Cecilia 
Volkmer Ribeiro for the help in sampling and to 
Prof Dr Wolfgang Maier from Tiibingen Lhii\ersit\'; 
Geniiany, for SEM sessions; F;abian Tricarico from the 
MACN SEM unit for his fine work with SEM photography. 
C.I. is a researcher from die Consejo Nacional de Investi- 
gaciones Cientfficas v Tecracas (CONICET), Ai'gentina. 


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of tlie Spliaeriidae (Mollusca: Bivalvia) based on partial 

mitochondrial 16S rDNA gene sequences. Invertebrate 

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tematics of .Sphaeriidae (Bi\al\ia, Eiilamellibrancliiata). 

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nas Gebiete. Archiv fur Molluskenkunde 78(4/6): 149-156. 
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THE NAUTILUS 122(4):236-24.3, 2008 

Paee 2.36 

Some remarks on the gross anatomy o^ Adelomelon fenissacii 
(Donovan, 1824) (Gastropoda: Volntidae) from the coast 
of Patagonia, Argentina 

Maria Eugenia Segade 

Guido Pastorino 

Museo Argentino de Ciencias Naturales 
Av. Angel Gallardo 470 3° piso lab. 57 
C1405b}R Buenos Aires, ARGENTINA 


Specimens of the volutid Adclomclon fenissacii (Donovan, 
1824) were dissected. Tliese were collected during an extraordi- 
nary low tide on Febiiian', 2006, at Playa La Mina, Puerto San 
Julian, Santa Cruz Province, Argentina (49°09' S, 67'';37' W). 
The gross anatomy, along with radulae and shell iiltiastructure, 
are described for the first time. An;ilvsis of these featuies sug- 
gests a close relationship witli the other species of the genus 
Adelomclon IDall, 1906, and raises doubts about the validity 
of Pachyct/inhiohi Iheriiig, 1907, a genus in which it has been 
included hx I'ecent autliors. 

Additional Kci/ivords: Southwestern Atlantic, Mollusca, 

Alxmt 30 uauies have been applied to Voiutidae horn 
the western Atlantic, trom Venezuela to Argentina. Of 
these, 16 are living in Argentine waters. Most are poorly 
kno\\ni, with data deri\ed onlv Irom their original de- 
scription, which generalK^ included \ei"\' little biological 
information. In addition, the tiixonomic status of several 
names awaits clarification. 

Adclomelon fenissacii (Dono\an, 1S24), a locally com- 
mon volute similar to A. Jnrisihaiui, is known only trom 
its shell. Recent findings of egg capsules suggest a compl- 
etel\' different reproducti\'e biolog>' (Penchaszadeh and 
Segade, submitted). 

In this paper we provide new anatomical and ultra- 
structural data for Adelomelon fentssacii derived from a 
large ontogenetic series of lix'e specimens, and compare 
them with otlier species of the same genus. 


The familv Volutidae includes the most conspicuous spe- 
cies among all endemic mollusks of the Southwestern 
Atlantic. This is so not only because of their large size, 
but also because of their interesting reproductive 
biology (Penchaszadeh, 1976, 1999; Luzzatto, 2006). 
Volutes hve in quite diverse euNaronments (i.e., subtidal 
to deep water; soft, muddy, sandy, or mixed bottoms; 
cold, temperate, or warm waters). Several species are 
extremely rare, while others are quite common. At least 
lAvo species, particularly in Uruguay, are the subject of 
extensive commercial fisheries. These art- Zklona 
diifrcsnei (Donovan, 1823) and Adclomelon hrasiliana 
(Lamarck, 1811). Otlier species are likely to be exploited 
in the near luture (Gimenez and Penchaszadeh, 2002). 
In addition, volutids usually are top predators with great 
ecological importance in tlie marine rcnilm. Species of 
volutes studied thus far generally have l)een found to lie 
predators on otlier mollusks and/or scavengers. 

Aiilhor lor corrcspoiidcncc, 


Specimens used in this study were hand-collected dur- 
ing an extraordinary low tide on February 2006 at Playa 
La Mina, Puerto San Juhan, Santa Cruz Pro\ance 
(49°09' S, 67'37' W) (Figure 1). The surface water 
temperature was 15°C. Animals were relaxed in freez- 
ing sea water, presei-^ed in ethanol 70% and dissected 
under a stereoscopic microscope. Radulae were pre- 
pared according to the method described by Solem 
(1972). Photographs were taken using digital cameras 
and images were also digitally processed. Shell ultra- 
stiaieture data were procured from freshly fractured 
colabral sections taken from tlie central portion of 
the lip on die last whorl of several indi\iduals. MACN: 
Museo Arsentino de Ciencias Naturales. 


Family Volutidae Rafinescjue, 1815 

Subftrmily Zidoniuae H. .Adarns and A. ,\il;nns, 1853 

Genus A(7r/<);//</,-/( l);ill, liKili 

Adcldiiiiloii l('i'ii.s\iii-ii (Donovan, 1824) 

M. E. Serade and G. Pastorino. 2008 

Paw 237 

San Matlas 


Buenos Aires "^ 

Puerto Quequen V^ 


Puerto K:^J| \ 
San Juliart\ "S ) 

( ) • 


70= 69= W= 

66= 65° 64= 63 = 

61= 60= 39= 5B= 57= W 

Figure 1. Map showing the stud\" area of Pla\a La Mina, 
Santa Cruz, Argentina. 

Vohita fenissacii Dono\an, 1824, 2, pi. 67; Sowerbw 1846 

1: 203, pi. 46, fia. 7; Reeve, 1849 fi, pi. 10. ficr. 23, 

spec. 39; Dall, 1907: 362. 
Vohita nidis Grav in GriffitJi and Pids;eon, 1834: 601, pi. 

30, fig. 1. ' 
Voliita oviformis Laliille, 1895: 20, pi. 1, figs. 1-2, pi. 2, 

figs. 5;3-56, pi. 7, figs. 121-138, pi. 10,'figs. 4-9. 
Vohita ovifonnis tijpica Laliille, 1895: 20. 
Vohita ovifonnis hngiuscuhi LahtUe, 1895: 20. 
Vohita ovifonnis fraterciila Lahille, 1895: 20. 
Vohita (Cymbiola) fenissacii Donovan. — Strebel, 1906: 

100. pi. 9, figs, 46, 46a, 48^9. 

Adclonichin (?) jcnissacii Dono\an. — Clench and 

Turner, 1964: 157, Pi. 98, figs. 1-3. 
Adehvncdon (sic) fenissacii (Donovan). — Castellanos, 

1970: 110, pi. 8, fig. 7. 
Adchnncdon (sic) fcrnisacii [sic]. — Castellanos, 1970b: 1, 

figs. 6, 9. 
Adclomelon (Paclujcijnihiola) fenissacii (Donovan, 1824), 

—Weaver and DuPont, 1970: 108, pi. 45C, 45D. 
Adchnnehn fenissacii (Dono\an, 1824). — Castellanos 

and Landoni, 1992: 12, Pi. 1, fig. 8. 
Pachijcijmbiola fenissacii (Donovan, 1824). — Poppe and 

Goto, 1992: 116, pi. 38, figs. 3-5. 

niedium in size, up to 122 nnn, 
color grayish-browai. Aperture semi- 

Description: She 

solid, fi.isiforni 

circular, dark-browai within. Protoconch of IV2 smooth 
whorls; teleoconch of up to 4 slightly conx^ex whorls; 
spire low, sometimes some\\'hat upturned: spire angle of 
80°, suture well defined. Columella cun'ed, orange, with 
three to six folds set obliquely to siphonal fasciole, regu- 
larly distributed except for tlie anteriormost one, which 
is separate from the others. Columellar callus usuallv 
weak, but sometimes thick. Siphonal canal fairlv broad 
and shallow. Growth lines span the surface, sometimes 
producing irregular costae. Shell ultrastructure showng 
three layers: a crossed-lamellar aragonite la\er in which 
the crystal planes are arranged peipendicularK' to the 
growing edge of the shell, and also pei"pendicular to 
the middle la\'er (25% shell thicfaiess); a middle laver 
(50% shell thickiiess) of colabralK" ahgned crossed- 
lamellar aragonite, and an outer layer (25% shell thick- 
ness) of amoi'phous calcite (Figure 18). The innermost 
layer varies in tliickness according to tlie sector of the 
lip: along the most cun-ed sector it appears diicker, 
while it is reduced or absent towards the ends of 
die lip. This arrangement is cpiite similar to diat found 
in shells of A. brasihana (Figure 19) and A. bcckii 
(Broderip, 1836). 

EmbiTonic shells veiy thin (at hatching stage), whitish 
in the first whorls and dark browai in the last protoconch 
whorl. Surface covered by S-10 regularly spaced spiral 
threads in the last whorl, no plaits visible. Calcarella 
reduced, weaklv pronounced. 

Foot, head, and siphon are finely mottled puq^le 
in color. The contact surface of the foot is whitish. Foot 
and shell length are similar, (3perculum absent. Head 
broad and flattened, with two short tubular tentacles 
that separate the lateral lappets from die central one. 
Eyes are very small and located near the base of the 
tentacles. The penis emerges directlv beliind the riglit 
cepliahc lappet (Figure 21). The siphon, also well pig- 
mented, is muscular widi paired and symmetrical sipho- 
nal appendages emerging from the base of the siphon 
and spanning half its length. 

The bipectinate osphradium has apro.ximateK' 100 
equal leaflets. The ctenidium is I'/a times as long as the 
osphradium. The h)'}3obrancliial gland is thin. The pro- 
boscis shows the same color pattern as the foot, head 
and siphon. Mouth opening is triangular. 

Page 238 

THE NAUTILUS, Vol. 122, No. 4 

M. E. Segade and G. Pastorino, 2008 

Page 239 

Figures lS-19. Shell ultmstnicture. 
Scale bars = 500 |.im. 

18. Aclcloiuclon fcnissacii (Donovan, 1S24). 19. AddomrJon hiyisiliiiitn (Lamarck, 1811). 

Radiilai- ribbon naiTow, up to 17,4 mm in length (n = 13; 
X = 11.7: DS = 1.97), witli 49-74 rows (propoitional to shell 
lengtli) with one tricuspid centi-al tootli per row (Figm"e 22). 
The radulae increase die number of teedi witli age (Fig- 
ure 32). Centi'al toodi diin and long, anterior profile con- 
cave witli a N-entral-posterior tliickening (Figiu-es 23 and 
24). Lateral cusps of die rachidiim toodi are similar in size 
to die central one. Each cusp widi a dorsal shallow indenta- 
tion or gi-oo\'e present, where die corresponding pre\ious 
cusp imbricates (Figures 24—25). EmbiYonic radulae 
showing about 15 rows of teedi where lateral cusps are 
shorter dian central cusps (Figure 26; Table 1). 

Sahvar\' glands (racemose glands) large and irregu- 
larly shaped. Accessoiy salivaiy glands (tubular glands) 
\-en' long and extremely convoluted, distally expanded, 
o\erl\ing dorsal surface of salivaiy glands. The tubular 
glands can easily be separated from the racemose 
glands. Ducts of the accessory salivary glands descend 
lateralh- to oesophagus and are sub terminal with respect 
to die nioudi. Thev never fused and end separately. 
Ducts of sah\'an' glands become embedded in the 

oesophagus anterior to small \'alve of Leiblein. The an- 
terior esophagus runs behind the proboscis, and passes 
dirough die nen'e-ring where an externally inconspicu- 
ous valve of Leiblein is located. The gland of Leiblein is 
veiy long and extremely convoluted (Figure 20). The 
posterior oesophagus continues to the stomach. 

Material Examined: (D = specimen was dead when 
sampled; A = alive) Punta Cavendish, Puerto Deseado, 
Santa Cruz province, 2 D, collected in 5 m depth 
(MACN-In 31354); Patagonian coast, 1 D (MACN-In 
113S5); Playa Cabo San Pablo, Tierra del Fuego, 4 D 
(MACN-In' 12532); Punta Sinaia, Tierra derFuego, 
Expedicion Facultad, 5 D (MACN-In 12531); Ushuaia, 
Tierra del Fuego, 1 D (MACN-In 9441); Punta Maria, 
Rio Grande, Tierra del Fuego, 9 D (MACN-In 35113); 
Bahi'a San Sebastian, Tierra del Fuego, 1 D (MACN-In 
35393); San Sebastian south, Tierra' del Fuego, 4 D, 
Exp. Facultad de Ciencias (MACN-In 12530); Playa del 
Rio Grande, Santa Cruz pro\ince, 1 D, (MACN-In 
6647); 52^^20' S, 68° 18' W, Station 28, Cabo Vi'rgenes. 

Table 1. Dimensions (mm) of some adults oi Adelomclon ferussacii from MACN collection. 











1 1 

















Shell lengdi (mm; (SL) 














.\perture lengdi (AL) 




























Spire angle (°) 














Radula lengdi (RL) 


11. .3 












Number of radular teeth 














Figures 2-17. Shells of A. fenmacii and A, brasiliana. 2-13. Adelomelon femssacii (Donovan, 1824). 2-4. MACN-ln 37013, 
Puerto San Julian, Santa Cruz! 5-6. Holoty-pe oi Valuta nidis Griffith and Pidgeon, 1834, NHM 19920177. 7-10. Embiyo shells (at 
hatcliing stage) MACX-In 37014. Puerto San JuUdn, Santa Cruz. 11-13. Juvenile shells MACN-ln 37012, 3 km North of Puerto San 
Sebastiln. Tierra del Fuego. 14-17. Adelomelon brasiliana (Lamarck, 1811), embryo shells MACN-ln 37015, off Mar del Plata, 
Buenos .Aires. Scale bars = 10 mm. 

Page 240 

THE NAUTILUS, Vol. 122, No. 4 

Santa Cruz province, A.R.A. Bahia Blanca, 1 D, in 11 m 
(MACN-In 24080); Balii'a Laura, Puerto Deseado, Santa 
Cruz province, 1 D (MACN-In 9199-16); Baliia San 
Sebastian, Tierra del Fuego, 1 D (MACN-In 21154); 
Estancia "Viamonte", Rio Grande, Tierra del Fuego, 
2 D (MACN-In 27219); Isla Quiroga, Puerto Deseado, 
Santa Cruz province, 1 D (MACN-In 26199); Punta 
Arenas, Chile, 1 D (MACN-In 9040-27); Rio Grande, 
Tierra del Fuego, 24 D (MACN-In 12529); Playa 
La Mina, Puerto San Julian, Santa Cruz Province (67° 
37' W, 49° 09' S), 10 A in low tide (MACN-In 37487). 

Distribution: Adcloinclon fcnissacii is a tv}:)ical com- 
ponent ot the Magellanic province, ranging from the 
province of Santa Cruz to Southern Chile. More north- 
em citations of the species (e.g., Clench and Turner, 
1964; Castellanos and Landoni, 1992), are here referred 
to A. brnsiliauii. No specimens of A. fenissacii from re- 
vised collections (MLP and MACN) were found outside 
of this range. 

Distribution According to Previous Records in 
the Literature: Puerto Gallegos, Punta Arenas, (Stre- 
bel, 1906); Santa Cruz coast, (Lahille, 1895); Southern 
Patagonia and Magellanic region (Carcelles and William- 
son, "l951); Golfo San Matias to Straits of Magellan 
(Clench and Turner, 1964), however diey only examined 
specimens from die localities of Puerto Deseado, Cabo 
Buen Tiempo, Rio Gallegos, Bahia de la Posesion and 
San Gregoiio in Chile; Golfo San Matias to Magellan 
Straits (Weaver and DuPont, 1970); Santa Cruz (Castel- 
lanos, 1970b); Southern coast of Buenos Aires province 
to Magellan Straits (Castellanos and Landoni, 1992); 
Southeastern coast of Argentina, south to the Magellan 
Straits (Poppe and Goto, 1992). 


Clench and Turner (1964) suggested the inclusion of 
Vohita fenissacii in the genus AcJeloinelon Dall, 1906, 
because of shell similarity with A. hrasiliana. However, 
at that time they examined no complete specimens to 
confirm such generic placement. Weaver and Du Pont 
("1970) mentioned that no live specimens were collected. 
Later, Castellanos (1970) illustrated the radula confirm- 
ing the generic placement suggested by Clench and 
Turner (1964). However, data on the gross anatomy had 
not previously been reported. 

The last comprehensive taxonomic revision of the 
family Volutidae from the southwestern Atlantic was 
prepared by Clench and Turner (1964, 1970). They de- 
scribed the new subfamily Odontocymbiolinae and the 
new genus Odoiitocymblola, and finally resolved the 
confusion of previous authors (e.g., Pilsbry and Olsson, 
1954) between Adelonielon ancilla (Lightfoot, 1786) 
and Odontocijmbiola tnugeUanica (Gmelin, 1791). These 
authors also described two new species: O. pescalia and 
A. nasi fClench and Turner, 1964). The latter was in- 
cluded ill the new subsenns W'viircnd . After their work. 

Figures 20-21. Adchmclon fenissacii (Donovan, 1824). 20. 
Anterioi' alinieiitan' system. 21. Dorsal view of head, siplion, 
and penis of a male specimen, ae, anterior esophagus; asg, 
accesory salivaiy gland; cl, cephalic lappet; dasg, duct acce- 
soiy salivaiy gland; e, eye; gl, gland of Leiblein; nr, nene ring; 
p, penis; pd, penial duct; pe, posterior esophagus; r.s, radular 
sac; s, siphon; sg, s;ili\'an' gland; t, tentacle. 

several additional new species were described, in partic- 
ular from Brazil (e.g.. Leal and Bouchet, 1989; Leal and 
Rios, 1990). 

Most of the hterature dealing with southwestern At- 
lantic volutids is primarily taxonomic. Anatomic;d data 
are scarce and usually dra\vn from one or relatively few 
specimens, sometimes incomplete. Exceptions are the 
papers by Novelli and Novelli (1982) and Ayyaguer 
(2002), particularh' the latter, in which the authors de- 
scribed in some detail the anatomy ofZidona dufresnci. 

Clench and Turner (1964) and Ayyaguer (2002) men- 
tioned that Adelonielon hcckii, A. aiicilla. A. brasiliinni. 
and Zidonii dufresnci all ha\'e characteristic racemose 
salivaiA' glands IikiscK intci I\\ iiicd in the Inbniar 

M. E. Segade and G. Pastorino, 2008 

Page 241 

Figures 22-31. Radulae oi Adelomelon species. 22-26. Aclelomelon fenissacii (Donovan, 1824). 22. Frontal xdevv. Scale bar = 
2-50 jim. 23. Lateral \iew. Scale bar = 200 (im. 24-25. Rachidian teedi. Scale bar = 200 [im. 26. Radula of an embiyo. Scale bar = 
50 Jim. 27-31. Adelomelon brasiliana (Lamarck, 1811). 27. Radula of an embryo. Scale bar = 50 |im. 28-29. Rachidian teeth. Scale 
bar = 200 um. 30. Lateral \ie\v. Scale bar = 200 (rm. 31. Frontal view. Scale bar = 250 |im. 

Page 242 

THE NAUTILUS. Vol. 122, No. 4 


j= 60 


Z 50- 

I 40 H 

Z 30 

40 60 

Shell length (mm) 

Figure 32. Rt'lutionsliip hctweeii radiilar and .slicll Icngdi 
(R = 0.9178). 

not match those oi A. ferussacn. In fact they look more 
similar to those of other species oi' Adclomclon. Also, the 
gross anatom\' re'vised here does not show conclusive 
features to include A. fcnissacii in a different group as 
suggested previously. The only character that could 
stand as a difference are those of shell nioipholog)'. 
Pilsby and Olsson (1954), Scarabino et al. (2004) and 
Del Rio and Martinez (2006) concede that shell shape 
place A. fcnissacii close to Pachi/ci/inbiola. The)- do not 
mention that a calcarella is reduced, present in the 
young of A. ferussacn (Figures 7-13), but absent in 
those of A. hrasiJiana . At a closer look even the shell 
shape is substantiall)' different, since A. brasiliana has a 
more flattened spire and pronounced shoulder tubercles 
more similar to A. hecki than A. ferussacli. Further de- 
tailed anatomical studies will confirm the relationships 
of these Southwestern Atlantic endemic volutids. 

accessoiy glands. The radula in these species is also 
racliiglossate with a uni([ue central tricuspid tooth. 
These characters, together with several shell similarities 
placed A. fcnissacii in the Zidoninae subfamiK'. 

Radulae of the species A. fenissacii and A. hnisiUana 
are Cjuite similar (Figures 22 and 31). However, A. 
brasiliana has a wider and more convex base of the 
rachidian teeth than A. fcnissacii (Figiu'es 24 and 
28). Lateral cusps of the rachidian teeth are shorter 
and wider in A. brasiliana than in A. fcnissacii. 
These differences are probablv specific characters 
and are also present in the embiyos of both species 
(Figures 26 and 27). 

The name Pachijci/mbiola was proposed by Iher- 
ing (1907: 209) as a subgenus ol Adclomclon for 
A. brasiliana, which is actuall)' the t\'pe species. Pilsbiy 
and Olsson (1954) and later Scarabino et al. (2004) pro- 
moted Pachijci/nibiiild to generic j-ank. and mentioned as 
main characters an ovate shell with a short spire, a free 
oval egg capsule, and a protoconch without calcarella. 
Del Rio and Martinez (2006) also treated Paclti)ci/m- 
biola at the generic rank following the latter authors. 
They described five Tertiary species, three new, under- 
this genus, pointing out as main differences from Aclclo- 
melon the o\ate shape f)l the shell, the short spire and 
the protoconch witlionl calcarella, in agreenicut with 
Scarabino et al. (2004). Adcloniclou brasiliana lias all 
these features and a large free ovoid ovicapsulc, witli 
9 to 33 embiyos per capsule (Penchaszadeh and 
de Mahieu, 197(5; Luzzatto, 2006). On the other hand, 
A. beckii and A. ancilla, which belong in the subgenus 
Adclomclon, liave smaller egg capsules, always attached 
to liard substrates (Penchaszadeh et al., 1999). Adclomc- 
lon fenissacii presents egg capsules similar to tliose of 
A. beckii, which are globose hemispherical and flexible, 
with a white opaque color and a leatheiA' texture, 
altaclied to a hard substrate such as stones or rocky 
substrates ^Penchaszadeh and Segade, in preparation). 

The inclusion oi' A. fcnissacii in the subgenus Padiij- 
cymbiola was firsl suggested based on some external 
similarity to A. brasiliana. However, the egg capsules do 


We thank M. Griffin (UNLPam) and F Scarabino for 
the suggestions that considerably improxed the manu- 
script. Bronw)'n Gillanders helped to improve the 
English. Two anonymous re\iewers made useful sugges- 
tions on an early draft ol the manuscript. This contribution 
was supported by Project PICT No. 03-14419 from die 
National Agency for Scientific and Technical Promotion, 
Argentina. We acknowledge funding by the Consejo 
Nacional de Investigaciones Cientificas v Tecnicas 
(CONICET) of Argentina, to which GP belongs. 


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Abteihmg fur Systematik, Geographic nnd Biologic der 
Tiere, 24: 91-174. 

Weaver, C. S. and J. E. Dupont. 1970. Living volutes. A mono- 
graph of the Recent Volutidae of the world. Delaware 
museum of Natural Histoiy Monograph Series No. 1, 
Delaware, xv + 375 pp. 

THE NAUTILUS 122(4):244-251, 2008 

Paee 244 

A new species and a new record of Muricidae (Gastropoda) 
from Brazil: genera Pterynotus and Leptotroplion 

Alexandre Dias Pinienta 

Diogo Ribeiro do Couto 

Paulo Marcio Santos Costa 

Departaniento de Inveitebrados 

Museu Nacional, Universidade Federal do Rio de Janeiro 

20940-040 Rio de Janeiro, BRAZIL 


Leptotroplion atlanticKS. a new species of Lcploli'oiilioii 
Houart. 199.5, a genus of Trophoninae previously known only 
from the Indo-Pacific Ocean, is described from the northeast- 
ern coast of Brazil, from depths of 240-260 m. Leptotroplion 
atlanticus is veiy similar to Leptotrophon kastoroac Houart, 
2001, but has shorter spines, which are brown colored. The 
shell microsculpture of L. atlanticus was studied using scan- 
ning electron microscopy, which revealed a protoconch sUghtly 
granulated on the last quarter of tlie whorl and a teleoconcli 
with growth striae crossed b\' regularly spaced zigzag spiral 
lines and axial microstriae. The radula of L. atlanticus fits the 
pattern described for the genus, but the lateral/marginal cusps 
of the rachidian teeth are not as independent of each other, 
being somewhat fused into a common base. This study reports 
die first record from die South Atlantic Ocean of the Murici- 
nae Pterynotus havanensis Yokes, 1970, prexiousK' knowm only 
from the Caribbean region. 

Additional keywords: Muricinae, Trophoninae, South Atlantic, 
biodi\ersit\', taxoiionix 


The family Muricidae is represented in Brazil by 49 
species from six subfamilies, of which members of the 
subfamily Muricinae are most numerous, with 24 spe- 
cies in nine genera (Rios, 1994). Other genera of Mur- 
icinae recorded from the western Atlantic have never 
been recorded from Brazil until now. This includes the 
genus Pterynotus, which is found in deeper-water habi- 
tats (Yokes, 1970), with eight species reported from 
Florida and the Caribbean (Rosenberg, 2005), in addi- 
tion to mention records from other regions of tln' world, 
e.g., Ponder (1972) from Australia, Bouchet and Waren 
(1985) from the northeast Atlantic, and Houart (2001) 
from Nc;w Caledonia. 

The subfamily Trophoninae includes 18 genera 
fVaugiit, 1989). It is represented in the western Atlantic 

by five genera, of which only Tropliou Montfort, 1810, 
occurs in Brazil (Rios, 1994). More recently, Houart 
(1995) described the genus LcptotropJtoii, whose 26 spe- 
cies are restricted to the Indo-Pacific region, in New 
Caledonia (Houart, 1995, 2001) and eastern Indonesia 
(Houart, 1997). 

This report presents the lirst record of die genus 
Pterynotus Swainson, 1833 from Brazil, based on Ptery- 
notus havanensis Yokes, 1970, as well as the first record 
of the genus Leptotroplmn from the Atlantic Ocean, 
based on a new species described from northeast Brazil. 


This icport is based on a sample collected on the Cano- 
pus Bank, State of Ceara, in 2005, from a biogenic sub- 
stratum, in 240-260 m depth. Identification of die 
species was based on original and subsequent descrip- 
tions. In cases where fragments of soft parts were avail- 
able, an attempt was made to record some anatomical 
characteristics using camera lucida, especially of the 
head-foot and the pallial cavity. Radnlae were prepared 
for SEM according to die niediodologv described by 
Bandel (1984). 


Family Muricidae Rafinesque, 1815 
SubfaniiK Muricinae Ralinesque, 1815 
Genus Pleryiuiliis Swainson, 1833 

Type Species: Murex j)iniuitus Swainson, 1822 b)' 
subse(|n('nt designation. 

Pterynotus liaianensi.s \'okes, 1970 
(Figures 1-12) 

Pterynotus (Plei'ipuitus) hai(nu-nsis \'okcs, 1970 (new 
name for Murex tristielius Dall, IS89, non Murex 
tristichus Beyricii, 1854.) 

A. D. Pinienta ct al.. 2()()S 

Pase 245 

Figures 1-S. Ptenjnotus havancnsis \'okes, 1970; 1—4, 7-8. M.XllJ 11057, length = 24.4 mm, width = 16.5 mm. 5-6. MXRJ 
1100.3. 1. Shell in apertural \ie\v. 2. Shell m abapertural view. .3. Shell in apical view. 4. Shell in lateral view. 5. Protoconch. 6. Detail 
of teleoconch sculpture. 7. Radulae in dorsal view. 8. Detail of rachidian teeth. Scale bars: 5-6 = 100 \xm\ 7-8 = 10.0 \xm. 

Murex [Pteronotus) tri.^tichm Dall, 1889: 202, pi. 15. fig, 2). 
Pteropui-pura tristicha. — Dall, 1927: 58. 
'Slurex iPtenjnofus) fiistichus. — Clench and Farfante, 
1945: 36. pi. 20. figs. 1^. 

Ptcn/iiofiis phanciis auct.- 
■(non Dall, 1889). 

Ptenjnotus phancus DalL- 
100 (in partj. 

- .\bbott, 1974: 175, fig. 1856 
-Radvvin and D'Attilio, 1976: 

Page 246 

THE NAUTILUS, Vol. 122, No. 4 


Figures 9-12. Ptcnjnntus havancnsis Yokes, 1970 (MNRJ 11057). 9. Head-font in apertural xiew, female. 10. Opeiruluni in inner 
view. 11. Operculum in outer view. 12. PaJial cavity, female. Scale liars = 1.0 mm. Alihre\iations: an, anus; cm, I'olumellar uiusele; 
en, ctenidium; et, cephalic tentacle; ey, eye; hg, lnp<il)r;uiclii;il inland: ml), iii.iiilic- linidc r op, DpcK iilnin: os, dsphnidiuui; ov, 
oviduct; .si, siphon. 

A. D. Pimenta et al., 2008 

Page 247 

Ptcn/iiotii.'i Ixn-inicnsis \bkes. — Fair, 1976: 47, pi. 13, 

■fig. 164. 
Ptcn/notus (Ptenjnofiis) phaiieiis auct. — Harasewych 

and Jensen, 1979: 4, i'ig. 3 (in part, non Dall, 1S<S9). 

Description: Shell of moderate size (up to 25 mm 
in length), fusiform, diin; color white. Protoconch 
paucispiriil, with 1.5 whorls, glass\', slightly bulbous, 
ending in a thin \-ari.\. Teleoconch with up to five 
whorls, somew'hat eon\ex in outline, with three thin, 
flaiing, backwai-d-cuned, wing-like varices per whorl, 
with 5-6 digitations along margin of last whorl varices. 
Axial sculpture consisting of fine growiJi striae, more 
\isible in \entral \iew of \'arices, where diey cross with 
spiral direads, giving it a foliaceous appearance; no 
rnter\"arical nodes present. Spiral sculpture obsolete, 
bai'eK' discernible spiral threads, formed by microscopic 
shallow furrows, somewhat undulated and irregularly 
spaced; abapertural view of varices widi five-six spiny 
digitations per \'ailx on the last whorl and two digitations 
on spire varices; spine on shoulders of whorls largest. 
x\peiture o\al, peristome slightly projected, inner lip 
reflected, attached posteriorly, smooth, with small 
undulations corresponding to varical digitations; outer 
hp smoodi. Siphon canal rather long, shai-ply bent aba- 
perturaUy and to the right side; narrowly open; with 
prexious, old imbricate canal termination, recun'ed to 
die left side. 

R.\DUL.\ (Fem.'vle): Rachiglossate tvpe; radula ribbon 
long and narrow; rachidian teeth with five pointed 
cusps, central cusp the largest, die second largest at die 
margins, each area between central and marginal cusps 
with an acute tin\' cusp; marginal edge rectangular and 
somewhat pointed, base concave; lateral teedi sickle- 
shaped, broader at the base. 

He.\d-Foot (Fem.\le): Head poorly differentiated, eyes 
located laterally and in middle of somewhat long cephal- 
ic tentacles. Foot large. Operculum homy, ovate, cover- 
ing entire shell aperture; external surface widi terminal 
nucleus and concentric growdi lines; inner surface 
attachment area with single horseshoe-shaped scar, not 
positioned centraU\-, covering less than 50% of total area 
of operculum, widi one adventitious layer. 

R\LLL\L CwTH' (Fem.ale): Mantle border smooth; siphon 
long and narrow, widi smooth border, muscle attach- 
ment reaching as iar as gill. Osphradium bipectinate, 
unequal, right side about 1.5 times as wide as left, broad 
and long (about half total ctenidium lengdi). Ctenidium 
monopectinate, long and narrow. Hypobranchial gland a 
thin, poorK" de\'eloped glandular mass covering mantle 
bet\v-een anterior end of gill and o\iduct. 0\iduct occu- 
p\ing about half of palhal ca\ity length, broad. 

^ype Locality": Blake Station 51, off Havana, Cuba, 
between 445 m and 823.5 m. 

Material Examined: MXKJ 11003 (one individual 
and seven empty sheUs); MNRJ 11067 (one individual); 

MNRJ 11057 (one individual); all from Canopus Bank, 
96 miles off Ceara State, 240-260 m depth, from bio- 
genic substratum. 

Geographic Distribution: Off Georgia and Fernan- 
dina (Dall, 1927), Key \Vest (Fair, 1976) Florida, USA; 
Havana, Cuba (type locality); Pleistocene Moi'n Forma- 
tion, Costa Rica '(Yokes, 1992); Golfo of Uraba, Colom- 
bia (Yokes, 1992); Ceani State, Brazil (this study). 

Discussion: Ftenjnotus havanenis Yokes, 1970, was 
originally described as Murex (Pteronotus) tristichus 
Dall, 1889. The taxon was later included in a long list of 
synonyms of Ptenjnotus phaneiis (Dall, 1889) by Hara- 
sewycli and Jensen (1979), based on a wide variation 
found in several shell characteristics such as the axial 
sculpture, intervarical nodes and outer lip. Yokes (1992) 
revahdated P. havanensis, based on the more numerous 
varices on the early teleoconch whorls and the smoother 
shell surface. 

The only available publislied illustrations of P. hava- 
nensis are those of the holotj'pe (DaU, 1889: pi. 15, fig. 2; 
Clench and Farfante, 1945: pi. 20, figs. 1-4; Yokes, 1970: 
pi. 3. figs. la,b; Abbott, 1974: fig. 1856; Fair, 1976: pi. 13, 
fig. 164; Harasewych and Jensen, 1979: fig. 3; \'okes, 
1992: pi. 2, fig. 3), in adchtion to a Pleistocene fossil 
specimen from Costa Rica (Yokes, 1992: pi. 2, fig. 6). 
This study provides die first illustrations of Recent 
specimens (Figures 1-4), except for that of the holotvpe 

The Brazilian specimens herein stuched, all from 
Canopus Bank, about 96 miles off the coast of the State 
of Ceara (240-260 ni depth), are veiy similar to the 
liolot)'pe illustrations, with little sculpturing between 
the varices (Figures 1-2, 4), without the intervarical 
nodes described for P. plianeiis, in addition to axial and 
spiral growth lines, the digitations on the margins of the 
varices and no denticulate outer lip (Figure 1). These 
characteristics, along with die shape and sculpture of 
the varices, clearly distinguishes P. havanensis from 
P. phaneiis. 

Yokes (1992) stated tliat both P. phaneiis and P. hava- 
nensis have denticulations on the inner side of the outer 
lip. Hcjwever, this detail is not stated in die original or 
subsequent descriptions of diis species, as well as in die 
holotvpe illustrations, which show a smooth outer lip, as 
well as in the specimens from Brazil. Such denticles 
reported by Yokes (1992) probably correspond to the 
undulations by die varical digitations. In addition. Yokes 
(1992) considered that P. havanensis bears several vari- 
ces on die early teleoconch whorls, making reference to 
the figure in Harasewych and Jensen (1979). Brazilian 
specimens bear die usual three varices on the first tele- 
oconch whorls. 

The radula herein illustrated (Figures 7-8) has some 
minor differences from the radula of P. phaneiis illu- 
strated in Harasewych and Jensen (1979: 15, fig. 17). 
In P. havanensis, die central and lateral cusps are longer 
in relation to the total leiigdi of the tooth; also, the two 
inner cusps are narrower and shorter. 

Paee 248 

THE NAUTILUS, Vol. 122, No. 4 

Subfamily Troplioninae Cossmann, 1903 
Genus Leptotrophon Houart, 1995 

Type Species: Lcptotroplion caroac Houart, 1995, by 
original designation. Recent, New Caledonia. 

Leptotrophon atlanticus new species 
(Figures 13-26) 

Diagnosis: Shell densel\ spin\': color cream whiti' with 
brown spines; paucispiral protoconch witli slight granu- 
lated microsculptiu'e on last quarter of last whorl. 

Description: Shell up to fi.H mm in lengtli (holotype), 
biconic, and densely spiny; spire high. Color creamy 
white, with light browii spines in li\e specimens. Proto- 
conch rounded, paucispiral, with 1.50 to 1.75 whorls, 
witli slightly granulated microsculpture, forming faded 
spiral cords on last quarter of last whorl; terminal \ ailx 
low. Teleoconch with up to 4.25 slightly shouldered 
whorls. Sutiu-e impressed. Axial sculpture consisting of 
numerous weak, orthocline lamellate grou'th striae and 
spiny varices, regularly spaced, forming spines at inter- 
ceptions ol spiral cords. Spiral sculpture on spire of two 

Figures 13-20. Leptotrophon athinlicns new species. Holohiie, MNRf \ 1004, length = 8.8 mm, width = 4.6 mm). 1.3. Apcrtnnil 
view. 14. Ahapertural view. 1.5. Apical view. 16. Operculum in inner view. 17. Opereuhun in outer view. IS. Radulae in ddrsal \ie\\. 
19. Detail ofraehidian teeth. 20. IDetail of lateral teeth. .Scale l.ais: 16-17 = 500 ym ; 18-20 = 10 \\u\. 

A. D. Pimenta ct al.. 2()()S 

Patre 249 

Figures 21-26. LeptoiropliDii tithniticiis new species. I'arahpe, MNRJ 11009, leiit^tli = <S.2 iiiin, width = 4.4 iiiiii. 21. Abapertiiral 
\ie\v. 22. Detail ot sculpture on teleoconcli whorl. 2.3. Detail ot microsculpture on teleoeoncli whorl. 24. Protocoueh. 2.5-26. Detail 
of protoconeh sculpture. Scale bars = 100 |im. 

strong cords, tlie adapical one located at 1/.3 of whorl 
height below the suture; interceptions between axial var- 
ices and spiral cords form two priman- spines regularly 
arranged in spiral crowns, each crowii with 9-10 spines 
on last whorl of holot\pe; spines tall, channeled, coin- 
monlv adapicalK" and backward-cur\ ed, slightly larger at 
shoulder; spaces bet^veen adjacent spines have a squa- 
mous appearance, due to crossing of spiral cords with 
axial growtli striae; last whorl with four additional spiral 
spim" crowns, regularlv aiTanged at the base, along with 
corresponding spiral cords: secondary squamous small 
spines appear iiTegularlv arranged, interspaced with pri- 
mary" spines; holot\pe with a secondar,' spiral cord appear- 
ing between the two priman- ones closest to the suture, 
on die last half of tlie last whorl. Microsculpture formed by 
growth striae crossed b\" regular!)- spaced zigzag spiral 
lines and axial microstriae. Aperture slightly oval, rounded 
adapicaUx" widi about 1/.3 of total shell height. Columellar 

lip flaring, smooth and adapicallv adlierent. Anal notch 
indistinct. Outer lip smooth, fragile, primaiy spiral cords 
and grovN'tii striae visible widiin. Siphon canal long, nar- 
row, bent liaclcwards, nai"rowi\- open and snuxjth, witli 
seven imbricate old canal terminations. 

Radul.\ (Fem.ale): (if lacliiglossate t\pe; ladiila riblidu 
long and narrow; racliidian tooth trapezoidal, witli li\e 
pointed cusps; the two lateral cusps adjacent to central 
cusp somewhat fused into a bifurcated cusp, outermost 
cusps slightly larger; marginal edges rectangular well 
pointed, area between outer cusps and marginal edges 
with Uvo very small folds; base somewhat sinuous; lateral 
teeth sickle-shaped, broader at base. 

Operculum hornw elliptical, covering entire sliell aper- 
ture; external surface with terminal nucleus and concen- 
tric growth lines; inner surface attachment area with 
single horseshoe-shaped scar not positioned centrally, 

Paee 250 

THE NAUTILUS, Vol. 122, No. 4 

covering about 50% of total area of operculum, with one 
acUentitiou.s layer. 

T^pe Material: Holot)iDe: MNRJ 11004 (8.S mm); 
three paratyijes: MNRj 11009, all from t^|^e locality. 

Type Locality: Canopus Bank, 96 miles off Ceara 
State, 240-260 ni depth, from biogenic .substratum. 

Distribution: Known from type locality only. 

Discussion: The allocation of Lcplatrojthini u\ 
Trophoninae follows Houart (1995), who, when describ- 
ing Lcptotrophon, stated that he was considering 
Trophoninae in a traditional way, to include typical "Tro- 
plion-like" species, diagnosed as small, thin spinose 
shells with a flaring columellar lip. In fact, Kool (1993) 
stated that Trophoninae is probably a non-monophyletic 
group and, thus, Houart (1995) considered that 
future studies would probably show that the genus 
Lcptotroplion would have to be transferred from the 

The new species fits \'en' well in the diagnosis of 
Leptotroplion , and is very similar to several species from 
the Indo-Pacific, including the type species, L. caroae. 

Leptotroplion afkiiiticus bears the characteristic spin\' 
sculpture (Figures 13-15), ronnd-oxate aperture, and 
flaring columellar lip (Figure 13). The radula of L. atlan- 
ticiis (Figures 18-20) fits the pattern described for Lcp- 
totrophon, but the lateral/marginal cusps are not as 
independent of each other, being somewhat fused into a 
common base (Figure 19). In other species referred to 
Leptotroplion by Houart (1995), the lateral/marginal 
cusps are similar to L. athinticus [e.g., L. caroae and 
L. acerapex (Houart, 1986)]. In addition, two small mar- 
ginal denticles may be seen on the outer base of the 
marginal cusps (white arrow in Figure 19), a chai'acteris- 
tic found in other species of Leptotroplion described by 
Houart (1995). 

The protoconch ol the species described In Iloaiut 
(1995) shows considerable variation in shape, witli 
rounded-globose, acuminate, or carinate protoconchs 
present in different species. The tyj^e species has a cari- 
nate protoconch, and Lcptotrophon tilhntficiis has a 
rounded-globose protoconch (Figure 24). 

The most similar species are L. hastiuvac Houart, 
1997, and L. perehirns Houart, 2001. Both species are 
larger tlian L. atlanticiis. Lcptotrophon perchnis has a 
taller spire, more acuminate apex, and more inflated last 
whorl; in addition, it has more numerous and smaller 
spines. The shell shape of L. kastoroac is almost identi- 
cal to that of L. atlanticus, but the spines are longer and 
are more upward-ciuved. 

Previously to this work, no record of sliell miciosculp- 
ture for any species oi Lcptotrophon was available. The 
protoconch is always described as smooth. In fact, the 
protoconch of L. atlanticus lias an overall suK^oth ap- 
pearance (Figure 24), but SEM reveals that only its most 
apical region is smooth; the last quarter area, close to the 
varix, bears spiral sets of microscopic granulations (Fig- 
ures 25-26). A delicate microsculpture also covers the 

entire teleoconch surface, including the spiral cords and 
spines, with axial and spiral microstriae forming a some- 
what reticulate pattern (Figures 22-23). 

The operculum of L. atlanticus is similar in its outer 
surface to those illustrated by Houart (1995) for species 
of Lcptotrophon from the New Caledonian region. 
Houart (1995) did not describe the operculum of Lcpto- 
trophon internally; the operculum of L. athinticus has 
one adventitious layer. 

In spite of the several common characteristics in the 
shell and radulae moiphologs- of Lcptotrophon athinti- 
cus and the species from the Indo-Pacific, the generic 
allocation herein used should be considered as provi- 
sional, due to the wide geographic separation between 
the new species and the other species in the genus 
Lcptotrophon, most of which bearing protoconchs 
that indicate non-planktotrophic development. All 26 
previously described species of Leptotroplion are re- 
stricted to the Indo-Pacific (Houart, 1995; 1997; 2001); 
Lcptotrophon atlanticus new species is the first record 
of this genus outside that region. The bathymetiy 
of L. atlanticus, on the other hand, falls within the 
bathymetric range reported tor the Indo-Pacific species 
(200-720 m). In the future, direct comparisons with the 
Indo-Pacific species, especiallv including characters 
\isible nnK' under SEM, would be most helpful to estab- 
lish the ilegree oi similaritx among these species. 


We are grateful to Vlr. Roland Houart (Institute Royal 
des Sciences Naturelles de Belgique) for conmients on 
the ta-xonomy of the species. \'inicius Padula (MNRJ) 
and Franklin Noel dos Santos (Universidade Federal do 
Para) provided additional bibliography. Dr. Janet Reid 
re\ised the English text. Dra. Noemia Rodrignes helped 
with the SEM photos. We also thank hvo anonymous 
i-ex-iewers for their comments and suggestions. Special 
thanks are due to Mr. Antonio Gil Bezerra and Ms. Elisa 
Grachohl Bezeri'a, owaiers of INACE Shipyard (Indi'is- 
tria Naval do Ceara) for the loan of the fishing boat and 
to Mrs. Jose and Marcus Coltro for financial support to 
the collecting efforts. Conselho Nacional de Desenvol- 
\iniento Cientihco e Tecnologico (CNPq) provided a 
PROTAX pos-doctoral grant to PM. Costa. 


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THE NAUTILUS 122(4):252-25.S. 20()S 

Patie 252 

A new species o^ Chkimi/cJoconcha Dall, 1884, from southeastern 
Brazil (Bivalvia: Chlamydoconchidae^ 

Luiz Ricardo I^. Siinone 

MiLscLi dc Z()ologi;i da Unnc-rsidade de Sao Paulo 
CaL\a Post;J 42494 
04299-970 Sao Paulo, BRAZIL 


The second species in die genus Chlaimjdoconcha is 
described. Clilainijdocoiiclia avalvis new species, occurs oii 
the coast of Rio de Janeiro coast, in southeastern Brazil. The 
new species has very reduced valves and a mantle surrounding 
the entire body, two features of the genus. The outer surface of 
the mantle lacks papillae e.xcept for a single one located close 
to the excurrent siphon. These are distinctive characters of 
ChUimi/doconcha orcutti Dall, 1884, from the eastern Pacific 
coast ot North America, the single other knowm species of the 
genus. Some of the more interesting anatomical characters of 
the new species are: posterior pair of retractor nuiscles of foot 
free from valves, absence of adductoi' muscles, gastric st\le sac 
totally separated from intestine, and the presence of a single 
(excurrent) siphon. 

AdditioiKil kri/ivords: Anatomv, western Atlantic, Rio de 


The geniLS Clilamxdoconcha Dall, 1884 (tspe species by 
original designation: C. orcutti Dall, 1884) has l:)een 
known to be monotypic. Chlamijdoconcha orcutti occurs 
from California to western Mexico (Carlton, 1979; 
VIorton, 1981). The species is characterized by reduction 
ol the shell, which is restricted to the anterior region of a 
spherical mantle cover; the mantle outer surface has 
many, somewhat equidistantly distributed papillae. After 
the original description, furthei' anatomical studies of 
C. orcutti were done by Bernard (1897) and Morton 

A sample collected by Ijiologist \'inii'ins Padula on the 
coast of Rio de Janeiro was sent to the autlior for study. 
The analysis ol the material revealed a new species of 
Cidaintjcloconcha, formally described herein. This paper 
is also the first discoveiy of the genus in (lie Atlantic 
Ocean, representing the second known species in the 
genus. The present description also includes a detailed 
anatomy, which is discussed in comparison lo C orcutti 
(Bernard, 1897; Morion, 1981). 

The tiLXonomie allocation ol the genus Chhtuufdo- 
concha has been pj-obleniatic. It has been included in 
the Galeommatidae (Morton, 1981), but full family 
status has been assigned (Chlamydoconchidae, Bernard, 
1983), as part of the Galeommatoidea. Full superfamily 
status was also considered (Chlamydoconchacea, Keen, 
1969). The Galeommatoidea, are mostly moUusks with 
usual bivalve shells, but may also include highly modi- 
fied, slug-like animals, y\'ith iutei'nal and reduced shells. 


The specimen was delivered presened in 70% EtOH. 
A photo of the li\ing specimen was taken before pi'e- 
seivation. The dissection of the presened animal was 
performed by standard teclniiques, under a stereo 
microscope, with the specimen immersed in the alcohol. 
All dissection steps were also photographed (e.g.. 
Figures 3-5). Drawings were made with aid of a camera 

Abbreviations used in ligiu'es are: an, anus; au, auri- 
cle; by, byssal gland; cc, gill ciliaiy connection; ce, cere- 
bral ganglion; co, cerebro-\isceral connecti\'e; tld, ducts 
to digestive diverticulae; dh, dorsal hood; di, inner 
demibranch; do, outer demibranch; es, esophagus; fg, 
gill food groove; fni, posterior loot retractor muscle; fr, 
anterior foot retractor muscle; ft, foot; ga, genital aper- 
ture; gi, gill; go, gonad; gs. gastric shield; in, intestine; 
ip, inner hemipalp; ki, kidne\; nib, m;mtle bojilcr; mo, 
mouth; mt, niantic; ne. iiephiopore; op, outer hemi- 
palp; pa, pedal apeitmc of nuintk'; pc, pericardium 
pg, pedal ganglia; pi, p:illial papilla; pni. pallial muscles 
pp, palp; rt, rectum; sh, shell; si, excurrent siphon 
ss, st\li' sac; st, stomach; ty, t\phlosolis; ve, \'entricle 
vg, visceral ganglia; vm, viscenil nuiss. 

Institutional abbreviation: MZSP; Musen de Zoologia 
da Universidade de Sao Paulo, Brazil 


C.ldauuidocoiuhii avalvis new species 
(Figures l-2()j 

L. R. L. Simone, 200S 

Page 253 

Figures 1-9. Chlaimjdoconcha aiahis Holohpe photos. 1-2. Living specimen, dorsal and lateral views, photo \inicius Padula. 
3-5. Preserved specimen. 3. Dorsal \ievv. 4. \'entral view. 5. Left view, right mantle lobe partially removed and deflected anteriorly, 
right gill deflected upwards. 6. Right valve, outer view (transversal section aitificially done). 7. Same, inner view. 8. Left valve, inner 
view. 9. Right valve, ventral view of its posterior, concave region. Scale bars = 1-.5 = 2 mm; 6-9 = 0..5 mm. 

Pase 254 

THE NAUTILUS, Vol. 122, No. 4 

Figures 10-1.3. Clih/iiiijflocoiiiliii aoalvis anatomy. 10. Left view oi entire animal, riglit nianlli' iolir partiallv removed and 
deflected anteriorly (right in Figure). 11. Region of right valve, internal surface of mantle removed, showing pallial nnrscles (pm) 
originating in valve, some adjacent .structures also shown. 12. Gill, transversal section in its middle region. 13. Right palp, outer 
heniijialp deflected anteriorly, a slioit portion il nmcr (leiiuliiaiich also shown. Scale hars = I nun. 

Diagnosis: Species with a single papilla close to exciir- 
rent .siphon. Anterior pallial gland shallow. Internal shell 
size about 10% of mantle surface; with rounded, almost 
squared posterior end. Anterior paii' of pedal retractor 
muscles -.vitli a branch originated from shell. Gastric 
main chamber and style sac narrow and long. 

Description: Shell (Fr:ures 6-9): Reduced, iueijui- 
valve, occupying about 1/10 <>( mantle, embedded into 
mantle anterior region (Figiuc 10, sh). Length appro>a- 
maleK 4 times wddth. Color white, opaque. Outline softly 
irregnlai'. Both xalves a.symmetrical; left vaKe about Va 
shorter than right valve (Figure 8) (this may be 

L. R. L. Simoiie, 2008 

Page 255 


Figures 14-20. CJihiim/cIocoiidia aviilvis anatomv. 14. Left \ifw of entiie animal emphasizing location of digestive tract and 
topolog\" of main muscles, ganglia and glands, animal artificialh i-epresented as transparent. 15. Midgut as ()i sitti, right view. 16. 
Same, sht longitucUnalK' to expose inner surface. 17. Renopericai'dial structmes and region, right aiiricle artificiallv disconnected 
from gill and deflected upwards, a transversal section of indicated level of right kidney also sliowm, 18. Cerebral ganglia, posterior- 
slightlv right \iew. topologx- of esophagus also indicated. 19. Pedal ganglia, right and slightly posterior view. 20. Viscerid ganglia, 
right and sHghth' posterior \iew. Scale bars = 1 mm. 

abnormal). Shape somewhat deformed and irregular; 
flattened, planar. Prodi.ssoconch rounded, sub-termi- 
nal: located in middle of anterior fifth of \'al\e length: 
shape semispherical. witli small dorsal bulging portion; 
0.26 mm long. 0.31 nun height. Outer surface some- 
what irregular with strong comrnarginal undulations 
and witli rounded, concave impressions; with ventral 
edge elevated (Figure 9). Calcareous concretions close 
to peripherv' on right valve (Figure S). Periostracum 
extending about 1/3 beyond calcareous portion of each 
valve, vvider dorsallv: color vellowish. transparent. Hinge 

edentulous. Ligament small, restiicted to umbonal 
region, relatively wid