THE NAUTILUS

Volunw 116, Niiiuhcr 1
April 4. 2002
ISSN 0028-1344

A quart erhj devoted
to vuilacolo^ij.

voocis Ho:-' ;

APR 1

Wooes t-ioc, ij.M 02043

EDITOR-IN-CHIEF

Dr. Jose H. Leal

The Baile\-Mattliews Sliell Museum
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The Bailey-Matthews Shell Museum
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EDITOR EMERITUS

Dr. M. G. Harasew'Ych

Department of Invertebrate Zoolog)'

National Museum of

Natural HistoiT
Smithsonian Institution
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CONSULTING EDITORS
Dr Riidi'j;er Bieler
Depart ment of ln\ertebrates
Field Museum of
Natural Histon-
Chicago. I Lfi0605

Dr. Arthur E. Bogan

North Carolina State Museum of

Natural Sciences
Raleigh, NC 27626

Dr. Philippe Bouchet
Laboratoire de Biologic des

Invert^bres Marins et Malacologie
.Museum National dTIistoire Naturelle
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C;enter forCon.servation Research

and Training
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Dr Robert T.Dillon. Jr.
Department of Biology
College of Charleston
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Dr. Eileen H. Jokinen
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Dr Douglas S. Jones
Florida Museum of Natural History'
Universit)- of Florida
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Dr Charles Lydeard
Biodiversit\- and S\stematics
Department of Biological Sciences
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Los Angeles C'ounty Museum of

Natural History
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Department of Living Invertebrates
The American Museum of Natural

History
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Museum of Zoolog)' and Department

ol Biology
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Florida Museum of Natural History
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Mr Richard E. Petit

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Dr Gar)- Rosenberg
Department of Mollusks
The .Academy of Natural Sciences
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Los Angeles Count) Museum of

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Department of Ceolog)'
University of California at Davis
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.Aquatic Ecology Laboratoiv
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Dr. John B. Wise

Houston Museum of Natural Science

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THE €7 NAUTILUS

CONTEXTS

\'()Imiic 116, Niunlx'i' 1

April 4. 2002

ISSN 002H-1344

Eugene \'. Coan RfCt-iit castfrn Pacific species ol SiiiiLiiiiiuildrid and

Psrnuiilillti i BixaKia: Psamoliiidae ) 1

^^'allel• Narclii The aiiatoiin and hinctional ni()i"pli()I(>i;\ ol Tu rhi

Fabio di Dario vciitnci>\ii (Ciraw 1S.3S) (BixaKia: X'cncridac) 13

Berntl R. Sehone Usi> nl ilNIDS ilif\ainctli\ldisila/anc) to dn oii;anic

David Bentlev microstnictures in etclictl hi\al\i' iiiolliisk and lianiat'lc

shells 25

Philippe Bouchet Gone with the wind: a pelaij;ic marine spi'cies dcsci-ihed as

an endemic land snail from the Bahamas 32

Note

Miehael Rex Bioo;eoy;raph\ of the ileep-sea i^astropotl I'lihizz'ui pliiiiorhis

(Dall. 1927): an micominon form ol rarit\ 36

Notiee 24

'I'liis piililic-atidii is sponsored in

pai'l l)\

tile Stall' ol I'loi'ida, Dcpartnu'iit

of State.

Dixisiou ol (Cultural Ailaiis,

and

tlu' Florida Arts Coiiucil

THE NAUTILUS llfii 11:1-12. 2002

Page 1

Recent eastern Pacific species of SangiiuioJaria and Psammotclla
(BivaKia: Psammobiidae^

Eugene V. Coan'

Department ol Invertebrate Zoolotn.'

and Geolog)'
Caliiomia Academy ol Sciences
San Francisco. CA 94118-4599 USA
g;ene.coan@sierrachil).org

ABSTRACT

There are three Recent eastern Pacific species ol Sdii^iiiiinlcir-
ia, S. ovalis Reeve, 1S57. .S, tcUinoides A. Adiuns. 1S5(), and .S.
tenuis Olsson, 1961, and one species (if P'idiniiKitclla. I' hciiini
(Pilsbn and Lowe, 1932). A neot\]i(' lor /' hcrliiii and Iccto-
hpes lor S. tellinoklcs and its junior s\iion\ni S'. purpurea are
designated herein. The distributions of the species are docu-
mented, along with their fossil occurrences and their relation-
ships to other Recent and to fossil species.

INTRODUCTION

Ha\ing discussed the eastern Pacific repi'esi'utatixcs ol
tlie genera Hvtcrodonax (Ct)an, 1973: 46—46) and Gnri
(Coan, 2000). I herein complete the rexiew ol tlie lamilv
Psammobiidae with treatment ot four speties tliat have
previously been assigned to tlie genus Saiij^uiuol/iriu.

Previous reviews of San^uinolaria are those ol Reeve
(1857) and Beitin (1880). Trv'on (1869) listed tlie then-
known species. Dall (1898, 1900: 978-979) and Wilhui
(1993) di.scu.ssed the genera of the Psammobiidae.

Thus tar. there are no papers on the anatnmv or bi-
ologv of S(i)i^ui)iolaria or Fsaiuinotella.

M.ATLRIALS AND METHODS

In tlie following treatment, eacli valid ta\on is followed
bv a sviionvniv. information on tApe specimens and tvpe
localities, notes on distribution and habitat, iind an ad-
ditional discussion.

The svnonvmies inclutle all major accounts about the
specie.s, but not most minor mentions in the literature.
The entrie.s are arranged in chronological order under
each species name, with changes in generic allocation
from the previous entn. if anv, and other notes given in
brackets.

' Mailing aildress: 891 San Jude .Venue. Falo Alto. CA 94.)()(v
2640, US.A; also Research .\ssociate. Santa Bad)ara Museuni
of Natural Histon' and Los .\ngeles Countv Musenni ol X.it-
urai HistoA.

The distributional inlormation is based on Ht'cent
specimens I have examined, except as noted. Habitat
information is scant, because most material has been col-
lected in beaclidrift. Fossil occiuTenees are taken from
the literature.

References are provided in the Literature Cited lor
all works and ta\a mentioned.

Moi-jjhological Characters: -\ combination of shell
shape and color, .uid various aspects of the shape oi the
pallial sinus sidfice to distinguish among the four species
discussed here: these arc detailed in the tlescrijitions and
in Table 1.

Abbre\iali<)ns: The following abbreviations are used
in the text: ANSP, .Academv of Natund Sciences of Phil-
adelphia, Philadelphia, Penn.svlvania, USA; BMNH,
British .Museum (Natural Iliston) collection. The Nat-
ural Iliston- Museum. London, England; C.\S, (.'alilor-
nia Academv of Silences, San Francisco, California,
USA; ICZN, International C!onnnission on Zoological
Nomenclature; L.\{'M. Natural Iliston Museum ol Los
Angeles Countv. (iaiifornia. USA; PRI. Paleontological
Research Institution, Ithaca. New York. USA; MCZ.
Museuni of Comparative Zoologv. Hanard Uiiiversitv.
Cambridge, Massachusetts. USA; MNHN, Museum na-
tional d'llistoire luiturelle. Paris. France: SHMNH. S;m-
ta Barbani .Museum of Natural ilistorv. S.inla Bariiara,
Calilorni;i. USA; SDMNII, S;in DiegoMnseum of Nat-
ural Ilisloi-v, S;ui Diego. California. USA; UMML. Uni-
vcrsitv of Miiuni \huine Laboraton-. Rosenstiel School
of .Miirine and .\tmospheric Science. Miami, Florida,
USA; USNM. United St;ites National Museum collec-
tion, NatioiKil Museum of N;itural llistorx', Smith.sonian
Institution, Washiuglou. DC. US.\; Z.MC;, Zoologisk Mu-
seum Copenlnigen. Denmark.

.\hilciial in the private collections ol Carol C. Skog-
huul. Phoenix. .\ri/ona, US.\; anil Kirstie L. Kaiser,
Puerto \ ;illarta, Jali.sco, Mexico, was also examined.

Pagf

THE NAUTILUS. Vol. IIH. \,.. 1

Table 1. Kc\ (lilTrR'ntiatins; cliaractcrs. si/r. ,iikI tn'i|iiciic\ (il Ea.stcni Pacific Sini'j_ini)i<liirin and Psiiuituofcllii

Color

Shape

allial smiis

MiLKimum No. lots
.size, mm studied

S. OKulls

S. tcUUioUlcs
S. tenuis

P. hciiini

white, with
beaks

purplish rci

while

purplish re(

pink etjuivalve, ovate, equilateral

e(jiii\al\e. oNate-elonsjate.

Ioniser posteriorK
e((ui\al\'e. o\ate-elouijate,

longer anIeriorK'

right \aKe more iiiHated.
elongate, lontrer posteriorK

gieatk expanded, pointed

dorsallv, meets pallial line at

appro.x, 50° angle
expanded dorsallw meets pallial

line at 90° angle
greatK' expanded, pointed

dorsallv, meets pallial line at

approx, 75° angle
elevated, rounded dorsalK, meets

pallial line at approx. .30° angle

34

20

72

79

34

1

93

98

studieil:

198

.SYSTEMATICS

FamlK' Fs;inini()!)iidae Fleming. 1S2S
C.eiius SaiiiiuinohirUi Lamarck, 1799

Sanguiiudaiia Lamarck. 1799: S4.

Tvpp .species (li\ m()uot\p\ ^; Soltii sdii'j^iiiiiohnlirs (imelin.
1791: 3227,'

Dt'scriplion: lv|ui\aKi'. with a narrow posterior gape.
Periostraciiin (liin. Pallial siiurs deep, mocleratelv to ven'
elexatcd dorsalK, its doisal line with an expanded nniscle
attachment area just anterior to posterior atlduetor. Pos-
terior enieiform mnscle scar without a small anterior sat-
ellite scar Hinge n.irrow to moderate in width: teeth
small; n\ mnh weak.

Sau<iui)U)laria oralis l-leexf.
l'"ignres 1. 2. 9

hS57

Saiiiiuiuiilnria ill alis Hee\e. i S57: pi. 1. fig. 2; Nhircli. hSfid:
185; CariXMiter, 18«4: 563 [LS72 reprint: 49| l;is a possihh>
suionym ofS'. inininla]: Tivon, 1869; 78; Bertin, h880: 84;
iball. 1898; 61 |as a s\iionv'm ol'S. tcllinoide.'ih Keen, 1958;
188, 189, fig. 460 |as a separate species): Keen. 1971; 243.
(ig. 610, 244 [not to be confused with lliuliihi otalis Ber-
lin. 1880; 92, pi. 4, fig. 5a, h, a species ol Solrlilliiui de-
.scrihed from -.m unknown) loealit\|.

S<tnii,\iin<ilana vcspciiiiui Pilshiv and Lowe. 1932; 90-91. I II
[;ls "Si-iiiclc" Vf.spciiina]. pi. 12. figs. 3, 4; llertkin ;ind
Strong, 1950; 220; Keen, 1958; 188 |as a .sviiouym of ,S.
(HY/Z/.s]: Ols.son. 1961; ,349, 558; pi. 85. fig. 5'fas a'sep;n-ate
.species]; Keen, 1971; 244 [as a svnon\ni of S. (iiiilis\:
Men/ 1986: 41.

Description: Ovate. etjuivaKc. linn, approxiuuiIeK"
efinilateral (heaks at 4S-50'7c I'rom ;mlerior end); an-
terior end ronnded; posterior end aeutelv ronnded.
without a radial sniens. Pallial sinus large, much ex-
tended and pointed dorsallv, meeting pallial line at an
appro.ximately 5()- augl.. prochicing a short extent of
noii-fonHnence l Fig. 9) Sculpture of fine eommarginal
striae. Color pink iicr uniliunes. heeoniing white to-
ward ventral margin; color \isihle botii cxteriorK' and
interiorly. Length to .34.2 mm |PiU 25921; Ciil'mico.
Panamal.

Type Material and Localitie.s:

S(in<s,iiinol<iriii ornlis — BMNI 1 1957 7 15. 1 . holotxpe. with bro-
ken right valve; length, 22.2 nun; height. 14.0 mm; width
(left valve), 3.1 mm (Fig. 1). "CJeutral .Vmerica", Hugh
('uniing The localitx is here elarified as being San
Juan del Siir, Rixas Province, Nicaragua 11.3°N!. tol-
lowing KJZN Cixle Recommendation 76A.1.4.

Sinifj,iiiiu>liiria vc.spt'iiina — ANSP 155013. holot\pe, pair;
length, 31.1 nnn; height. 20.9 nnii; width, 1L3 mm (Fig,
2). ANSP .398873, paratype; length, 26.4 mm. SDNHM
50773, paratvpes, 6 pairs, 2 vahes. San Juan del Sur, Rivas
Province, Nicaragua (1L.3°N); H.N. Lowe, 193L ANSP
154663, paratvpe, left \al\e; length. 29.7 mm. Corinto,
Chinandega Province, Nicaragua (12..5°N); H. N. Lowe,
1931.

Distribution: (;u;i\ui;is. Som)ra. .Mexico (27.9°N)
[CAS 1.54.3fi9], to (aianieo, Los Santos Province, Panama
(7,2°N) [PHI 2.5921]; from the intertidal zone to .37..5 m.
.Material examined: 20 lots.

Material trom the Golfo cie Panama that has been
misidentified as this species [USNM 962S.3. 962S7.
96353, 9fi.36L 96383], or labeled as Siin<:,iiini)liin(i s/j.
[Kaiser collection], while eloselv resianhling a Saiigiiin-
olarid. Ikis conspicuous lateral teeth, a low, elongate pal-
lial sinus, and slightlv oblique connnarginal seuljitnre.
This material is instead Tclliiia [Utiiclliiia^ uinu/diid

Ihrllcm ,111(1 Strong. 1949 (pp. S.5-S6. 97
26 i I see ;ii,so: Ols.son. 1961; 409).

il. l.f

1gS.

Discussion: This sjiec ies is vei"v similar to its western

.Miaulic I lologue. .S. stin<nii\ioicnt(i (Cwiielin. 1791:

322.5 — as Solent |svnon\ins; Solcii fiiiYilti.s S[)engler.
1794: 111: Tclliiia lihucuiula Hciding. 179S: 1S6: .SV;ii-
iiuiiiolaiid rosea l.:uiiai'ck. ISOl: 12.5: Lt'liaria rosacea
Schumacher. 1S17: 122-12.3, pi. 6; Saniiniiiolana iiirca
.Morch. IS5.3: H)|. which occurs from Florida and Texas,
through the WesI Indies lo Hiazil. Snti'^uiiiolaria oralis
diders in being more rounded iuid less pointed jiosle-
rioiK. in h;i\ing :i n.trrower hinge pkile and liner lunge
teeth, and in not altaining as large a size (S. saiiiiiiiiio-
Iciita can attain at least 4.'>. I mm in length i. .\ddition:illv.
the palli;il simis ol .8. saicj^iiiiiolciila meets liie p:ilh.il line
;il ;l 90" ;ingle. ;ind il does not rise ,[s hu" dois:ilI\. w illinul

E. \'. Coan. 200;

Pacre 3

Figures 1, 2. S<nii^iiiiii>hiii(i oidlis Kccm
Lowf. Iciititli 31. 1 iiini.

1. ||iil(i|\pi' (if S iir(;/(s, l(-ii<;tli 22,2 nun. 2. Ilcili)l\pr ut .S. icspciiina Pilshn and

a.s shaqi an an<j;le at its summit (Fiii;iiri' 10'. Tliis wcstrrn
Atlantic species niav also develop tliickci- slielK lliau an\
eastern Pacific material of .S. oKilis Si/irj^uhinliiriii msca
Lamarck is not preoccupied In Sulcii mscm {imiliu.
1791 (p. 3227). which was based on a (i'j;nie in ( .'heninit/
(1782: pi. 7, fig. 55) that seems to he a Si'lililliiia This
.species was attrihuted to the Hed Sea l)\ Bertiii 1 ISSO;
98). It is also not preoccupied In' Tclliiui ruM-ii Cmelin.
1791 (p. 3238), which is based on an illustration in Knorr
(1771: pi. 9, fig. 3) that uia\- be of a specimen ol Asripliis
dvjiomta (Linnaeus. 1758: rS87, as Wiuis).

San^iiinohiriti lifiva Deshayes. 1855 (p. 32fii. de-
scribed from an miknowni localit}' (see also l^eexe. 1857:
pi. 1, fig. 1 I, has been suggested to be an additiou:iI
s\non\ni of .S. sau^uinolcnta, one based on light-coloied

material iCosel. 19S9: 715h (losel based this conclusion
on two lots in the MXIIX from Ncnicrn/. \cracru7,.
Me\ic-o. However il' material in the Sli.MNII (133229,
345687) and the (i.\S (152575' from ni'ar \eracruz is
correctK- identified as S. i i/rcir this is a diiierent species,
indeed. Diill ( 1898: 58) gave the distribution of .S". vilira
as being from Texas to ("olon. I^anama, but (liis needs
to be verified, 'i'lic Sl^\l\i! and (.'.\S material is white
autl translneeut. with |);illi:il sinuses that are not \ei"v
dorsalK extended iiud that meet the pallial line almost
veitieallv I LiL^ure I i : the largest specimen is 52.4 nun
in leiiiith. 'Hie Ivpi' lot iil .S 1 Una in the I^MNII should
be examineil to be certain the species has been correctK'
iiiteipreted.

S(in<iiiiii(il<iiin tiiirciuiiiclri .Martens. 1S79 (p. 744)

Pasie 4

THE NAUTILUS. \ .

Ifi, \(

[synonvni: S. africmia Cosel. 19S9: 711-715; pi. 1. fig.
K: pi. 7, figs. 26, 27)]. is a similar West .Mrican species.
(A still earlier name that ma\ a]iply to tliis species is
Tcllimi arhatina Spengler, 1798: 100.) In (iescrihing ,S'.
africana. Cosel (19S9) differeiitiati'd it Irom .S. saufftin-
olenta as l)eing larger and more elongate, with less
brightly colored beaks. These characteis would also sep-
arate S. mtrcot'incta Irom ,S'. oiidis.

San^uiiioliiria tclUnoUlv.s A. .Ailanis, 1S50
Figures 3-6, 12

Siin<liiiri(il(iri(i Iclliiu'iilrs \. .Xdaui.s, 1851): 170, pi. (i, fia;. (i;
Kecxc, 1,S.57: ])l. 1, fig. 3: CailH-nter, IS57a: 286, 301:
18571): .31: Morch, 1860: 185; Caipeiitcr, 1864: 563 [1872
reprint: 49J; Tnon, 1869: 78 [as "S. lelliiiidcs"]- Bertin,
IcSSO: 84; Dall, 1898: 58; Meitlein and Strong, 1950: 219-
220; Keen, 1958: ISS, 189, fig. 462; Olsson, 1961: 348,
550, pi. 77. figs. 10. 11; Keen, 1971: 244, 245, fig. 611

Tellina miiiwlri C.onU. 1851 : 90; 1853: 397, pi. 16, fig. "l; Gould
and Cai-peiiter. 1857: 199: Caipenter. 1857a:' 226, 231,
245, 301: 18571): 547 (in Appendix as a senior ,s\iionvni of
.S. piirj>iin'(i]: Miircli, 1860: 185 [as "muwacca" and a svii-
onvni of S. icUinoidcfi]; Gould, 1862; 212; Caqienter,
1864: 537, 541, 543, 549, 563, 668 [1872 reprint: 23, 27,
29, 35, 49, 154]; Tr\'on, 1869: 78 [as a svnonvni of ,S.
iellinokh's]; Bertin, 1880: 84; Dall, 1898: 16 '[as a .synonym
of .S. tcUUwidcs]- lleitlein and Strong, 1950: 219 [as a
s\iu)n\in of S. Iclliiioides]- Johnson, 1964: 110; Keen,
1971: 244 [as a s\non\ni ol .S. Icllinoidcs]

Sanguinolaria puqwmi Desha) es, 1855: 346; Reeve, 1857: pi.
1, fig. 5; Gould and (Carpenter, 1857: 199 [as a svnonym
of S. miniatii]: Caipenter, 1857a: 226, 231, 245, 301, 352;
lS57c: 31, 548; 1864: 563 11872 reprint: 49]; Tiyon, 1869:
78 [as a synonym of ,S. frilinoidcs]; Dall, 1898: 61 [as a
synonym of .S. trlliiioidcs]: llertlein and Strong, 1950: 219,
251, pi. 2, figs. 5, 8 [as a separate species]; Keen, 1958:
188, 189, fig. 461; Keen, 1971: 244 [as a .synonym of .S.
Icllinoidc's]

Sanffiiiu>hiriu lellinoidc.s cloufidtd Morch, 1860: 185; llertlein
and Strong, 1950: 220 ]not preoccupied 1)\- Ciiri (Pstiiii-
moUicna) clonfialn (Lamarck, 1818: 514 — as Vs(nnmohia),
which is widespread in the Indo-Tacific (Willan, 1993: 61-
64)]

Description: Ovate-elongate, equivalve, some\\h;it
tliicker-shi'lled than ,S. ovalis at a similar si/.e, becoming
sturdy in large specimens; posterior end somewhat lon-
ger (beaks at 40-42"t from anterior end); anterior end
roimded; posterior end somewhat produced, set oil l)\
an indistinct radial sulcus a[)proximatelv Kvo-thirds dis-
tance Irom end generally most evident in large speci-
mens. Pallial simis large, proiluced and pointed dorsallv,
meeting pallia! line at 90' anule, its ventral margin thus
completelv confluent with pallial line (Figure 12). Sculp-
ture ol fine ccuiniargiiial striae, (.'nicilbrm muscle sears
inflated in large spcciin.Mis I. ;ferior color pink to pur-
ple, sometimes with d;nkcr curnmarginal color bauds:
interior often dark ]-iuiplish-re(l Length to 71.5 mm
(ANSP 220.326: Aeapiilco. C.uerrero. Mexico).

Type Material and Localities:

Siiitiiuiiiolarin liUinoidcs — B.VLMl 1966540/1, pair lectotype
here designated; length. 32.6 nun; height. 19.2 nun:
widdi, 8.4 mm (Figm-e 3). BMNH 1966540/2, paralecto-
t\pe; length, 32.5 mm. Gulf of California. The locality' is
here clarified as being GuaMiias, Sonora, Mexico
(27.9°N). loHowing ICZN Code Recommendation 76A.1.4.

Tellina minUitd — .\1C;Z 169258, holotype, pair; length, 51.0
nun; height, 33.0 mm; width, 14,3 mm (Figure 4). San
Juan [del Sur, Ri\as Province, Nicaragua] (11.3°N); Lieut.
T. P. Green. The localitv was mistakenK given hv Johnson
(1964: 110) as being in Orange County, Califonii:i,

Stnii^iiiiKildiiii imrpiircd — B.VINII 19f)6539/l, leclotvpe here
designated, pair; length, 50.0 nnn; height, 31.7 nun;
width, 15.1 mm (Figure 5). BMNH 1966539/2, 3, para-
lectotypes, pairs, lengths, 48.4 mm, 44.4 mm. The original
specimens came from the collection of Hugh Cuming, but
the t\pe loealit\ was gi\en as unknown and is here clar-
ified as being Guaymas, Sonora, Me.vico l27.9°N), fol-
lowing ICZX Code Recommendation 7(i\.1.4.

SdUf^uiiioldiia tellinoides elon^dld — ZMC unnumbered, holo-
tvpe, pair; length, 52.4 nnu; height, 30.2 nnn; width, 13.3
nnn (Figure 6). "Realejo" [Corinto, Chinendega Province,
Nicaragua] (12.5°N); A. S. Oersted, 1846-1848.

Distribution: Punta Pescadero, Pacific coast of Baja
Calilornia Sur (23.3°N) [USNM 22964], into the Golfo
de California as far north as Punta Diggs, Baja Cidifornia
(30.9°N) [CAS 150381], and Puerto 'Libertad, Sonora
(29.9°N) [ANSP 184183], Mexico, to Cojinu'es, Esmer-
aldas Province, Ecuador (0.4°N) [PRI 259208]; intertid-
a! /one to 14 m. Material examined: 79 lots. San^uitio-
laria tellinoides Ikus been reported from the Pliocene
|ania Formation at Puerto Jama, Manabi Province, Ec-
uador (0.2°S) (PiLsbn- and Olsson, 1941: 72). and (as
"(iff.") from the late Miocene Gatun Formation on the
Atlantic coast of Panama west of Colon (W'oodring, 1982:
673, pi, 115, fig. 12).

Discussion: The pallial sinuses ol this species become
more tlorsalK' pointed in large specimens. This species
can be distiuguislied from i'saiitiiiotclla heifini in being
e(]ni\aKe. and 1)\ its more rounded outline, less atten-
tuate jiosterior enil, :ui(l more (l()is:il!\- extentlcd and
pointed pallial sinus.

Sdiiiiniiuiliiri/i leiiui>: Olsson. 1961
Figni'c 7. 13

Sdiipiiiioldiid leiiuis OLs.son, IS)61: .349, 558, pi. 85, fig. 6;
Ke<'n. 1071: 244 [as a sviionvm of S. (wnlis; not a hom-
ouMii With Psdiioiu'hid tennis Deshaves, 1855: 320, a ,svii-
nuMii 111 ihc I IK Id- Pacific (.Idri dnonudd I Deshaves, 1855:
320— as I'sdnwiohid) (Willan, 1993: 22), nor with Soletel-
lirid tennis (Desha\es, 18.55: .349 — as Capsd (Cdpsella))
from the I'liilippinr Ishuuls (Willui. 1093: 77)[

Description: Ovate-elongati'; e({ni\aKe; ant(>rior (>nd
somewhat longer (beaks at 44'X from anterior end); an-
terior end rounded: postciior end I'ounded. with :i slight
(race ol ;i radial sulcus at two-thirds ol wa\ to postciior
end. Piillial sinus large, exlendi'd dorsalK', meeting ]iallial
at an ■ij)pi'i)\iui:i(e 75" angle (I'^igure 13). Scuiptnic ol

E. \'. Coan, 2002

Pai^c 5

Figures 3, 4. SiniiS^iiinnldriti ullininilcs A.
{ )i>ul(l: li'iiiidi. 51 ,0 mill

Adams. 3. I ,cilnl\pc ol' .S. tclliiiniilcs. lcii<j;tli -i^fi mm. 4. Ilolotypc (il Tclliiia iiiiiiialii

fine foiiniiai^iiial .striae. I'Alciini' cnloi' wliitc, willi a
lit^lit pink railial liaml a little anterior ot midline; while
inteiiorK. Lensjtli t(i oo.S nnii (li()l()t\pe).

T>pe Nraterial and Localily: AXSF 21Si)il lioh,
t\pe. pair: leiiiitli. 3.1S nnn: lieit^ht, 26.1 inni: widtii,
10. U nnn ( Fii^nre 7l: ( lanoa. Manabi Proxinee. Kenadnr
(0.5°N); A. A. Olsson, 1958. An additional fraiinientai-\
specimen cited h\ Olsson (1961) from Pnnta Montafiita.
Gnavas Province, Ecuador (1.8°.S). would he a paratypc;

il lias not been located in tlie r.MMI;. I'nlortiinateK;
none ol Olsson s specimens ol Suiiffiinolmid lia\i' \ct
been located in tlie UMML (\. \'()ss, e-mail. 2-3 May
2001 1,

Distnbiition: Tims far known onK from the original
specnneiis — ( .'anoa. Manabf Pro\ince (0.3°\ * ilioiotxpe),
presnmabK lo i'lnita .Montafiita. (Jnaxas Proxinee ( 1.8°.S)
(specimen missing), Ecuador. Botli specimens were col-
lected in beaclidrilt.

Pa<'e 6

THE NAUTILUS, Vol. 116. NO 1

Figure'. .5, (i. Sdii-j^'iinnliinii tclliiioidcs A. Adams. 5. Lfctolvpc (iT .S', imrpiircii l)cslia\cs. Icii'j;lh oDd niiii. (>. Il(il(il\]ic nl S.
icllinoUlis rlnu^aUi NKircli. linolli .52.4 iiini.

E. \". (Joan. 2002

Figures 7, 8. 7. Siiiii^iiiiHilariii Icniiis (iIssdii: Imlnlxpc liiiulli >-vS iiin
Tcllina hanlci/i Bertiii; leiit^tli. fiT.o mm.

S. I'sinitiiKitclld IkiIiui I'ilshiA aiu! I.iiwc : iu(il\ |ii' iil

Page S

THE NAUTILUS, Vol. 116. NO. 1

Discussion: Tliis is tlic rare Soiitli Aiiit'rifan alK ot S.
oifilis. It cliflers in not lia\inti rose-tolored l)eaks, in be-
ing thicker slielled, and in having a more rounded pos-

terior end.

Genus Psaminotclhi

Icrniiannsen.

1S.52

Psammotella Ik'rrniainiscn, 1S.52: 114, c.v "Fsaiiiiiiott'lle"
Blainville, 1S28: .541 | vernacular],

T\pe species (bv nion()t\pv): 'T. nifcscens Chenin.", = TcUina
nifescens Dillwvii, ISIT: 85, t'.v Chemnitz ins, = TclUna
cntcnta [Lightfoot], 1768: 10 [as Tellina "cmcntac"]. .58
[as T. cntenta]'^: first reviser: Rehder, 1967: 7. Recent,
western .Vtlaiitic. [mm Psammotella H. .4danis and \. Ad-
ams, 1S.56: 39.3, c.v Deshayes ms, = Psammotdlina R-H.
Fisciier, 1887: 1105, new name, a subgenns o{ Can and
perhaps a senior SNiion\ni of Psammotaena Dall, 1900:
976 (VVillan, 1993: 60)]. '

Description: Shell inequivalve; right valve more in-
Hated; posterior gape narrow. Pallial sinus deep, mod-
erately elevated dorsalK, without an expanded muscle
attachment anterior to posterior atlductor muscle scar.
Posterior crncitorm muscle scar with a small anterior
satellite scar Hinge moderatcK lieavA'; teeth small:
iniupli weak.

Discussion: I here rank this ,Ni'w World taxou as a
genus because of its major differences from Sau^uhm-
luria — conspicuous differences between left and right
\al\(>s. plus dillerences in its pallial sinus and cruciform
muscle scars.

Psammotella hcrtiiii ^Pilsbn and Lowe, 1932)
Figures 8, 14

Tellina hanlet/i Bertin. 1S7S: 268-269 Inoii Tellina haiilei/i

Danker 18.53: 53-54. pi. 10, figs. 4-6|.
Sanfifiinolaria heriini Rilsbrv' and Lowe, 19.32, new n.inu' lor

Tellina haulei/i Bertin. 1880, non Dnnker 1853; I'ilsbiv

and Lowe. 1932: 91. )43. pl. 10. figs. 7. 8: Hertlein and

.Strong, 19.50: 220-221; Keen, 1958: 188, 189. fig. 4.59;

Olsson, 1961: .349, .5.50, pi. 77, fig. 8: Keen, 1971: 244.

245, fig. (5l2: Hertz, 1986: 36.
Tellina nijesceiis "Chenniitz", auetl.. non T mfeseen-s Dillwvn,

1817, ex Chemnitz ms; Hanlev, 1846: .307-308, 332. pl.

63. fig. 213: Caipenter, 18.57b: 32; Rtimcr 1S72: 111-112,

pl. 27, figs. 1-3 [mm Tellina nifeseens DilKwn. IS17: S,5,

ex (;licinnil/ iiisj.

Description: h^loiigale; right valve conspicuously
more inflated th.m left; posterior end longer (beaks at
40-48'^ from and lit;; .•■idi; :uiterior end rounded; pos-
terior enfl attenuated., set off bv a radial sulcus near end

in riglit \aKe, tip truncate; posterior end siiuious in right
vahe. tip suhtmncate. Pallial sinus deep, its dorsal mar-
gin in right vaKe ele\ated, rounded to bhmtK' angular;
ventral margin of sinus meeting pallial line at an ap-
proximate 30° angle, confluent with pallial line for ap-
proximately 75% of its length (Figure 14). Sculpture of
fine, irregular conunarginal striae, strongest on posterior
slope; right \al\-e with radiid striae, strongest along \en-
tral margin and in large specimens. Cniciform muscle
scars bulbous in large specimens. Color pink to puqile.
sometimes with darker conunarginal bands. Length to
92.8 mm (LACM 71-179.12. Punta P.-qucfui, Baja Cd]-
ifornia Sur. Mexico).

Type Material and Locality: Tellina lianlciji Bertin
was based on the figure ot Tellina nifescens "(Jhemnitz"
in Hanlev (1846; see abo\e). which presumabK came
from the onl\- localit\- mentioned — Tunibes. Tumbes
Province, Pei"u (3.5°S). This specimen has not been lo-
cated in the BMNH (J. Pickering, e-mail, 7 June 2001).
nor in the Leeds Museum, present location of a sub-
stantial ;unount IIaule\s material (A. Norris. e-m;iil, 13
)uue 2001). The specimen selected by Pilsbrv and Lowe
(1932) as iiolotvpe" of their new name— ANSP 152068
from ./\capulco. Guerrero, Me.xico (16.9°N) (Figure S) —
would normalK have no type status, because the txpe of
;i renamed junior homon\ui remains the original tvpe of
that taxon (ICZN Code Art. 72.7). Howex'er.' because ( 1)
Tellina nifescens was long used to refer both to the west-
ern Atlantic species now knowii as Psammotella cnienta
and to the eastern Pacific /' heiiini. (2) we cannot as-
cert;iin wliich ol these two species Hanlev actually illus-
trated, (3) there is no good material t)f the eastern Pa-
cific species in coll(>ctions from Tumbes, Peni. and (4)
PilsbiA ;uul Lowe's "txpe" is ;i fine spetiinen long ;ic-
ctjriled t\pe status, this specimen is here designated
neot\pe of Tellina liaiilei/i Bertin. It is ;i p;iir of \aKes
measuring 67. .5 nun in length. 28. 1 mm in height. ;uid
12.4 nun in width.

Distrihution: L;ignn,i ())n de Liebic [Scinunons La-
goon], H;ij;i California Sur(27.8°\) ]ANSP 22.5928], La
Paz, Baj:i C;iliforuia Sur !24.2°\) [SBMNH 24586.
24587], ;ind Kmpalnie, Sonora (27.9°N) ]SBMNH
1.35133], Mexico, to M;ineor;i. Tumbes Province. Peru
(4.rS) |CAS 1.54.370; SBMXII 125767], and evidently
as far south ;is (^)l;in. I'iin;i Province, Peru (5.0°S) (Pa-
redez and ( !;uxlo/o ms; Mnsco de llistoria Natural,
UTiivcrsickid N;icioual .\I;i\()r de S;in .\I:ircos. Lima.
Peru); inlerlickil /one to 17 ni. \l,ileri,il e\:nnined: 98
lots.

Figures 9-15. Li:ii.',t ,:■
scars ol left and righl ■■.•'. 'I.
(Gniehn):CAS 152576: \h,i. i,
Veracmz, Mexico; lenL'th, 46 " ;
.S. tennis Olsson; liolotvp": liii'.'i!;
length. 66.6 mni. 15. P < nienta

■■■■•a] views ol slu'lls ol San^liitnohiria ,un\ I'stnnninh llii sliewnig liiii'j,!'. palli;ll sinns, ;incl .iildnetor

■ -./'v Reeve: CAS 1.50375: .\e,ipiileci (aieirero. Mexico: length, 26,2 mm. 10. S. saii'diiinolenta

'..u.iiha. Bnr/.il; length. 46, I hum IL ,S litrea Desliaves; C.'^S 152576; Boca ik-1 j-lu). \'ei-;icniz.

\1. .s lellimmles A. Adiims; .WSR 220.'i26; .\c;i]inIeo, Cnerrero. .Mexico: length, 41.5 nun, 13.

■J S miM M. Psammotella heiiini iPilsbn ;md Lowe); C.\S 1.50.380; ,\capnlco. Guerrero. Mexico;

.i'4lilloMf . ( olon. P;in;iin;i I'rovinee, I'linam.i; length. (il.O nnn.

E. \\ Coan. 2002

Page 9

10

11

12

13

14

15

Paire 10

THE NAUTILUS. Vol. 116. No. 1

Discus.sion: Psaiiiini>tvlla licrliiii is \v\\ similar (o its
western Atlantic lioniolos^ue. /' cnicnia lll.iiilitloot].
17S6: see nncler genus) [.synonyms: Tcllina opcrciddta
Gnielin, 1791: 3235 (as "T. opcrculalds" in Lugcluni
printing); Tcllina nifcsceiis nill\\-\n. I SI 7: S5. c.v Chem-
nitz ms]. The last sviions m is naii 7' rulrsrcns (imelin,
1791: 3238. which was based on (;ualtieri (1742: pi. 25.
fig. C), .seemingly a V<'ncnij)is. perliaps best regarded as
one of the nianv s\non\iris ol V. dectissatiis (Linnaeus,
175S: 690 — as Venus), as suggested bv Carpenter
(1857b: 32). PsaininofcUa rnicnta occurs throughout the
Caribbean to Hnizil. PsuiniiioivlUi rnicitta dilters from P.
bciiini in being more iuecjiiilateral. with a llattener left
valve and a more inflated right \al\e, and a less dorsallv
expandi'd pallial sinuses in both \aKes (Figure 15). Oth-
er supposed dilTereiices that have been suggested \an
too much among populations to be useful. For example,
Hertlein and Strong (1950) throught tliat /' rnicniii is
narrower posteriorlv, but this does not seem to be (he
case ifCnougli lots are studied. Thev also said that the
pallial siTiuses ol /' cniciita were "more arched posteri-
orlv" (?e.\tending further posteriorly) and "confluent
with the pallial line lor a greater distance." but I laii t
see nuich difference in these parameters.

Psaininotclla smitlncoodunnli (Maur\-, 1917: 393-394
[ = 229-230], pi. 64 | = 38|, figs. 1, 2— as San^iiinolaria
{Psaiiiuu)t('lla)). from the late Miocene Cercado For-
mation ol the Dominican Hepublie is presuniabK ances-
tral to these two species. .Vlann' pointed out its affinits'
to P. herliiti. but tlid not compare them. The original
figures are insulficientlv clear to see anv differences
from either Recent species. Weisbord (1964: 372) noteil
that this Miocene species differs from Recent material
in lacking radial striae in the right \aK('.

Psammolclla alottafii Olsson (1922: 432-433 | = 260-
261], pi. 32 [ = 29]. figs. 5, 6 — as S(iu<iiiiiioltiria {Sam-
motcUa) |sic] from the late Miocene Catun Formation
at Banana Hill on the Atlantic coast of Costa Rica, was
biiseil on two poorly presenetl \al\i's. The sjiecies was
said to be more elongate posteriorK than /' cniciilii and
to lack radial striae in the right xaKc. Weisbord if964:
372-373) added that the anterodorsal margin of the rigli(
valve and the posterodorsal margin of the left \al\i' were
luore concave tiian in /' cniciila. Both of these fossil
species are tlins far (oo |)oorl\ known lo draw an\ useful
conclusions.

I'or a comparison with Saii'^iiinoldrid Iclliiinidcs, see
undt-r this species.

L.XCLUDKD TA.XA

Saii'^nhudari:! (alifcrtiiaitd Conrail. 1837 (p. 231. pi. 17.

fiii. / ) I.-, ■liliirnirii. atirlt.. lunn. tiidi). is a s\non\ni of

Mannr r ■! (Liimaeus, 175S: 677 — as Tclliiid)

i^ • •-■ •17).

S- (!■ p. 81), is now regarded as a s\ii-

onvm ol • ■':; (C:oiuad, 1837: 230-231; |il.

17, fig. () :. -^'K I 426).

S. fli.sca {Ha\. :'''■ as J ,-,c/;/i»io/;/c/). a combiniilion

ol some ;iutliors, is aSMloinni of Mucoiiui hullliicd (Lin-
n;ieus, 1758) (Coan et'al.. 2000; 417).
.S. <^rdiidis Caipenter, 1857, c.v Gould ms, a ikhiuh uii-
dtini in Caipenter (1857a: 228, 349), is regarded as a
prolnible sviionvm of Xuttnllid iiuttaUii Conrad, 1837
(Coan et aL, 2000: 426).

S. mitldlla Conrad. 1837 (pp. 230-231; pi. 17, fig. 6) is
now placed as Xnttdllia iiuttidlii (Conrad, 1837) (Coan
et al.. 2000: 426).

S. paciftca (C^onrad. 1837: 241. pi. 18. fig. 13 — as Psam-
iitohid) is a combination bv some authors for the species
now known as llvtvrodoiuix pacificus (('onr;id. 1837)
(Coan et al. 2()()0; 428).

S. ndirnradidfd Caipenter, 1860 (p. 1). c.v Xuttall or
(Conrad ms. is a iioiiicu nudum now regarded ;is a prob-
able s\non\in of Gaii cdlifornicd (Clonrad, 1849; 121)
(Coan'et ;i[.. 2000 426: Coan, 2000: 3).

ACKNOWI.EDCMENTS

I ;ippit'ciate tlie help of the following curators, other
personnel and their institutions, who niaile specimens,
literature, and information a\ailabk'; .^dam J. Baldinger,
Museum of Comparative Zoologw Hanard Uni\ersitv,
Caiubridge, Massaclnisetts, US.\: Ruck) xon Cosel, Mu-
sc'-um national d'Histoire naturelle, Paris, France; Lind-
se\ T. Groves and James H. McLean, Natural Histon
.Vluseuni of Los Angeles Counts, Los Angeles, ("alifor-
nia, USA; Elizabeth Kools, Department of Inxertebrate
Zoology. California Academv of Sciences, Golden Gate
I\nk, San Francisco, California, US.\; Da\id R. Lind-
berg. Museum of Paleontolog\, Uni\ersitv of California,
Berkelev. ("alifonii;i, USA; Adrian Norris. Leeds Mu.se-
uiu, Leeds. I'aiglaud. UK; Joan Pickering. The Natural
Histoi-v Museum, London, England, UK; T\juana Nick-
ens, Natiouiil Museum of Natural Histon, Washington,
DC, US,\; GaiT Rosenberg, The Acadeiu\ of Natural
Sciences, Philadelphia, Penns\i\auia, US.A; Paul \'aleu-
tich Scott. Sant;i l^;irb;u';i Museum of Natural Histon.
Santa B;uiiara. (!alilornia, USA; Ole S. Tendal and .\nnie
Lone \'e(lelsl)\. /oologisk Museum, Copenhagen, Den-
uKirk; \;me\ \'oss, Uni\ersit\' of .\liaiui. .Miami. Florida,
ISA ( !arol ('.. Skiiglimd and kirstie L. Kaiser gener-
ousK m;ide ;i\;iil;ible materials or information from their
collections. .\!;m R. K;ib;it provided some ;ul\iee. Lind-
se\ T (iroN'es. Ke\in L;nnprel!. C;n()l (,'. Skoghmd. Paul
\alentieh Scott, ;uid Richard \\ illan ukkIc helpful com-
ments (III llic iiMiiiiscript. SIku'oii \\illi;mis helped pre-
p;il"e the phUes.

i.iii'Mail BF, CITFD

Adams. .\. 1850. Dcsc rijiliiins of new species ol sluils Iroiii (lie
("iiiiiiii>;iaii cdllection. I'roceeciings ol the Zoologiciil So-
ciel\ (if I. (.11(1(111 lor 1849 (200): 169-170, pi. 6 |ripriiitecl
in: XiiiiaK and Ma'.;aziiii' of Natural Histon i2>7(37): 69,
1S51I,

Adams, II and A. .Aihinis, 1S56 [1S53~1S.")S|, The ueiicra of
l^ccciit Miilliisca; arraiiilcd according (o tli(ir iir(.iaiii/.a-
tidii \aii \ii()rsl, I.oihIiiii liii). 3M— tl2, Ncn. 1S.')6|.

E. \'. C;()aii. 2002

Paw 11

Bertin. \'. ISTS. Revision des tellinuk'-s dii Mus(''uiii ill listoiir
Naturi_-lli_'. Noiuellt's Arrlii\c,s, Mum'muii iiatiniial
d'Histoire iiaturelle (2)1: 2()1-:561. pis. S, V),

Bertin. N'. IS.SO. Resisioii des sjaridees du Mnsenui d llisloiic
Xaturelle. Nouvelles .ArcliiNes. Musi nm iialinn.il
d'Histoire naturelle (2)3: .57-129, pis. 4. ,">.

Blain\ille. H. M. D. de. 1828. Telliiia. F. C. Lc\ranlt. vA.. Dit-
tionnaire des Sciences Natnrelles. . . . 52: 528-559. I ,v\ -
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Caipenlei. R R LSoTa. Report on tlie present state ol our
knowledge with regard to the .Mollnsea ol the west coast
of North America. Report ol the British .Association lor
the Adxancenient of Science 2(ilior lS5fi|: 159-.368 + 4
pp.. pis. fi-9.

Carpenter P. P. 1857h. Catalogue ol tile collection olMa/.atlan
Shells, in the Britisli .Museum: collected In Frederick Bei-
gen. London (British Musenm'. \ii + .552 pp. [.some coll-
ies as i-i\' + ix-wi] [also puhlislied sinmltaneonsK as ( 'at-
alogne ol the Reigen Collection ol .Ma/atlan Mollusca, in
the British Museunr Oberlin Press. Washington. \iii + \ii
+ 552 pp.] [reprinted: Paleontologic:il Hese:Li(.li liisliln-
tion, 196T.]

Caqienter P P 1860. Check Hsts of the slielK nl North \mer-
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ha!. West coast: Oregonian and ( ,'aliloniian proxiiue. 4
pp: No. 2(li), West coast: Mexican and Panainic proxince.
1.3 pp.

Caqienter P. P. 1864. Snpplementan report on the present
state ol (inr knowledge with regard to the Mollusca oFthe
west coast ol' North America, Report ol the British .Asso-
ciation for the Advancement of Science .'53 [lor 1S63[:
517-686. [reprinted: Caipenter 1S72: 1-172]

Caipenter, P. P. 1872. The mollusks of western North .America.
Embracing the second report made to the Britisli .Asso-
ciation on this subject, with other papers: reprinted In
permission, with a general index. Smithsonian Miscell.i-
iieoiis Collections 10(2521: xii + 325 + 1.5-121

Cliciiinit/., ]. 11. 1782. Neuer sxstematisches Concliylieii-Ciab-
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Coan, E, \', 1973, The northwest ,\mericaii Ps:inmiobiidae, The
N'eliger 16: 40-57, 4 pis.

(.'oan. E. \'. 2000, The eastern Pacilic Recent species ol die
bivahe genus Can (Tellinoidea: Psaiiimobiidaei, with
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1-29.

Coan. K. \'.. P X'aleiitich Scott and F R. Biaii.ud 200(1, BnaKe
seashclls of western North .America .Maiiiie ImaKc iiiol-
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\iii -I- 764 pp

(Conrad. T .\. 1837. Descnptions of new marine shells Iroiii
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227-268, pis, 17-20,

C:oiirad, T ,\, 1849. Description ol Iwo new genera ;ind new
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121,

Cosel, R, \(in. 19S9. TaxononiN of tropical wcsl American bi-
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D.ill, W U, 1900, Clontribntions to the Tertian fauna of Flor-
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groups treated ol ,iiid their .American Tertiarx' species,
P;irt \'. ■|'eleoilesiiiacea: Sdlcii to Diplodonla. Transactions
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Deshaxes, C. 1', 18.5.5, Descriptions ol iKnv- shells Iroiii the col-
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321-:336. i'281l: .337-3,52, (282i: .•3.5;3-368, (283): ;369-37L

Dillwxn, L, W, 1817, ,A descri]5ti\(' catalogue of Recent shells.
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Ciiallieii. N. 1742. Index lestarniii conchvliorum qiuu' adser-
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Ilaiilev. S. C T. 1846. ,Monoiira|)li of the genus TcUinu C. B.
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Herrmannsen, A, N. 1852. Indiiis genernni nialacozoorum
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Paee 12

THE NAUTILUS. \'ol. llfi. NO. 1

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Hinds, R. B. 1842. Deseriptions of new shells, .\nnals and
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Keen, A. .M. 1958. Sea shells of tropieal \\esl .\nierica; marine
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Keen, .'V. .VI, 1971. Sea shells of tropiciil west ,\meriea; marine
mollusks from Baja California to Peni, 2nd edition. Stan-
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Knorr C. W. 1769, 1771. X'ergnugi'n der .^ugen und des Ge-
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Linnaeus, C. 1758. S\s(ema naturae per regna Ilia natiu'ae . . .
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Martens, K, E. von. 1879. Ubersicht der son Herru W. Peters
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(1860;, (5i: 19.3-213 (lS61i.

Olsson A. A. 1922 The .Miocene of northern Costa Ric:L uilh
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124.

THE NAUTILUS 1 lfi( 1 ):13-24. 2{)()2

Pam- 13

The unatom\' and functional nioiphol{)g\' of Tivcia vcnitricosa
(Gray, 1838) (Bi\al\1a: Veneridae]

Waller Narehi
Fahio cli Dario

DcpurUiiiR'iito (.If Zoologia
liistituto de Biociencias
Uni\er.sidade de Sao Paiiln
BHAZIL
wiiaifliiCS'usp.br
lddario(S'ib. usp.br

ABSTRACT

A tlctailcd sIikK oI slicll. organs ol tlir iiiaiillc ia\il\, siplioiis,
and digcstixe tract of TivcJtt n-iithco.s/i is pre.st'ntcd, and lol-
lowed bv conipari,sons with other .specie.s of Tivrla. Tiiclii i i-ii-
tricosd occurs from Espi'rito Santo State in Brazil to La Palonia
in Urugnav. Indi\idu;ds li\e in .sand\ bottoms (■\p<ised to
poiniding .surf, where usualK- a large aiiionnt ol nialrnal in
suspension is present. The anatoniv and Innctional nioiphologv
of T. ventricosa are compared with those ol congeneric species
occurring oH the coast of southeastern [Brazil. The si[ihons ol
T vcntricosii are of t\pe B of Yonge. the ctenidia ol t\]ic C.i 1 1
of .\tkins, the relation between the labial pal]is and the etenitlia
belong to categoiT II of Stasek, and the stomach to t\pe \' ol
Pnrclion.

INTRODUCTION

Tlic \ I'licroidca is a large superfaiiiiK ol liixaKcs lliat
lias rachatt'cl hroailK worldwide, t\picall\ into soil, iisn-
all\' intertidal sediments. The genus Tivcia Link, ISIIT.
occurs on the .Atlantic and Pacific coasts of the .American
continent (Abbott, 1974; Coan et ah, 2(K)()).

Isolated references to Tiichi iciifhcoMi [('•vd\. bSo.S)
are foimcl mainlv in regional checklists (Lange de .\Ior-
retes, 1949; Gofterje. 1950; Bncknp. 1957; Hios. 197(1,
1975. 19S5, 1994), hut tliere are no icports ou ibe lune-
tioiud inoipholog\- of the species. I'iicld iciiliici'Mi oc-
curs Iroui southeastern Brazil to Urugua\ iHios, 19941
.Along the Brazilian littoral, the species is recorded in
the states of; Espi'rito Santo (Rios, 1970), Rio cle Janeiro
(Rios, 1970), Sao Paulo (Rios, 1970i, Parana (Cofferje,
1950; Rios, 1970) and Rio Grande do Sul (Bncknp. 1957;
Rios, 1970, 1994). The species was rec'orded in Itios
(1970, 1975. 1985, 1994) from the littonil of Uruguay
and Broggi (1970) collected hissil representali\es ol the
species from the Qiierandino FoiiiKitidu, Uruguax. and
described the distribution of the fixing animal Irom San-
tos (Bnizil) to La Paloma, Rocha (Uruguayi. Tiichi n-ii-

//■;V(<sy; was not recorded li\ (.'arcelles (1944' and (.'ar-
celles and Williamson (I95|i Irom |Ik> coast ol Argen-
tina.

.Although not sold ui llie coinmereial seafood markets,
the species is an import;nit food item for coastal po])u-
lations, tourists, and campers on the coast of Sfio Paulo
State, Brazil, The main goal of the present paper is to
proxide basic inlonuation on the ,matom\ and some as-
pects ol the lii<ilog\ ol the species.

.M.ATEfilALS ANf) MI"ni()l)S

Li\ing specimens of T niilricd.sri (Figure 1) weri' I'ol-
lected on the intertidal region of beaclies in the localities
of Sao Mcenle. Bertioga. and Ubatuba. all on the coast
of Sao PiLulo. Bnizil. Specimens were lound buried at
depths of appro\iiuatel\ 5 cm in sand oi' nuiddy-sand
substrates, in areas with strong dist)n-|)anc<' of bottom
deposits, wbicli end up suspended in large amounts in
tlu' wati'r column. Specimens wert' collected during low
title, when lhe\ e<inld be detec-ted, at the surface of the
substrate, b\ ,i little circular mound deposited around
the tips of liie siphons. Some of the study animals were
kept aliM' np lo I months.

Drawings wi^re m;ide from relaxed and pri-seneil
specimens. .Vhignesium sul|)liate was used as a relii-xing
agent, ('iliarv currents wei'e studied with the use of
Carborundum (F), carmine suspensions, or suspensions
of .A(|uadag. Organs and tissues were fi.xed in Bouin's
fluid, and sections (i-S |jLm in thickness stainetl with
Ehrlich's baematowlin .uid eosin. .\z;m. and .Shillon's
triple stain.

N'oiichcr specimens are dc|)osiled in the Museu (ic
/,oologi;i. Uni\ersid;ule de Sao Paulo. M/Sl' 3294S.
.32949.

RESULTS

.Shell (Fijiiires 2—4): 4"bc shell of T iciitricosti is
e(|ui\;iKc. ei juikitciiil. trigonal, with subcentral nndxi-

Pai^c 1 t

THE NAUTILUS, \o\. 116. X(

r<'>D^ or^o 0 ,

■'O-Q <

/fo .0*-J.c'?C>>-r

0

^

■i O oo , O^O oCi, ,,>^^0 Or, 9 0°

C^'^oCP^,

15, Po

g O O 0,2 O/
Co

:o^^ ■

0 o(>?<^"^ ao'5^&«^^o^:V^d 0,

^ - q;?^6<^^>^^ ; ^^^^^^o c> V 0

<5b

0-

Figure 1. Tivchi icnlrirosa. Aiiiiiuil iii iiatmal li.iiiital: cvtci-

rial \ie\v Iroiii the left side. Arrows show the direeti I ihc

iiRiirrent and exeurrent currents. .Scale line = 2 cm.

lies and beaks located close to eaeli oilier Tlie external
surface is smootli, oxcrlaid 1)\ a tliiek. <j;loss\ [lerios-
traenm. which gix'es a varnisiiccl appearance to the shell
vaKcs. The external color is cream with a xariahle pat-
tern of radial or zig/a<4 hrown streaks (Fimire 2). Sonie
specimens show alternating ime(|nal ra\s of straw and
chestnut color, with the rav color on llie ri'^lit \al\e
olteii more \ivi,l than that on the left. The interior ami
respeeti\e Iree inartjiu ot'tiie shell haxc a smooth whiti'
surlace. The pallial sinus is short, ohiiijue. U-siiaped.
The .scar of the anterior adductor imisele is o\al and
more impressed than the posterior one. which is almost
circular in outline (Figure 3). The hinge is strong, with
3 weli-dexcloped cardinal teeth in each \alve (Figure
4). The uiedi.m toed, is n.irrower. hiadedike: the an-
terior and posterior sets are hilid. with the posterior
one coar.seK .serrated. Thi^ knoh-like anti'iior lateral
tooth on the left \al\e fits into a deep dejjression on
the riglit \ai\e. Along tlie posterior dorsal margin of

the right \al\e there is a large grotne and a smaller
groove is present on the anterior dorsal margin of the
teft \al\e. Hie shell of the largest studied specimen
measured 7.U cm in length. 7.4 em in height and 5.2
cm in width.

Siphons (Figures 5, 6): Thi' siphons are short and
fused at the base. They are formed bv fusion of the inner
fold with the internal surface of the middle fold of the
mantle margin. The aperture of the incurrent siphon is
surroimded bv branched tentacles, and that of the e\-
iiirrent siphon has a tapered siphonal membrane with
simple tentacles at the basal region.

The incurrent siphon (Figures 5, 6) has about 200
tiiitacles deployed in 5 sets surrounding the aperture.
The tentacles are directed toward the aperture when the
animal is pumping water in. which allows for sieve-like
trapping of particles. The most e\tern;il set consists of
small, digitiform tentacles, some of which displax" a bi-
furcated tip. The four internal sets consist of branched
tentacles that decrease in ninnber and increase in length
in each succeeding set toward the internal region of the
siphonal opening. A U-shaped pigmented dark band is
present at the base of the group of three most internal
sets of tentacles. The incurrent siphon is \"er\' sensitiw
to touch and \ibrations and witlidiaws in part oi' com-
pk'teK upon stimulation.

The excurrent siphonal aperture (Figure fi) is smaller
than the incurrent siphonal aperture, .\bout 4S simple,
external tentacles of niilk"x-white color surround the base
of the siphonal uieiiibrain\ These tentacles are pig-
mented at their bases.

Siphonal Menihranes (Figure 7): .\ pair of siphonal
membranes is prest-nt at tlu' basal opi^ning of the incur-
rent siphon; these membranes eoiitribute to regulate the
water flow into the pallial ca\it\. Two other additional
si|)honal folds are present and unitetl at the dorsal side
of tlu' incurrent siphonal opening; thev eontimu' \en-
tialK and in anterior ihrection as expansions of each
iiiaiitle lobe.

Organs of ihe Mantle Canity (Figures <S, 9): He-

mo\al of the left shell \al\c and respectixe mantle lobe
exposes the organs in the iiumtle c;i\it\- (Figure 8). The
ctenidia ;n"e dorsalK loe;ite(l. The inner deiiiibrancli is
kirger than the onlei one; this latter bears .i snpraxi;il
extension, Tlie imiei iiumlle siirf;ice is coxered with
slioit cilia, except in a narrow border on its free edges.
The mantle edges are fusi'd for ;i short extension ;mte-
ii<ir to die iiK'iirreiil siphon, 1 )ors,il!\ lo these fused \en-
Ir.il m;i|ij;ins. the lii'j,lil\ mobile ;idilitioiial siplimial iiiein-
bi;iiies delimit a eaii.il, Ihe Iree edge of the iiiaiille li.is
4 folds and a large number of ramified tentacles, (hose
ill the proximal and distal regions ol (he pedal gape di-
\uled into seeondai'X and (erli,ir\ biaiiehes, 'I'hese short,
simple tentacles uia\ ;ilso be ;ibsent in the niedi;m region
of the pedal gape. When tlie ;miiiial is resting, the ten-
tacles on the ielt side .liteniale with those on ihe right
side, so til, it lhe\ prevent sand gr;iins Irom entering IIk'

W. Naalii and F. di Darici. 2002

Page 15

Figure 2. Tiicla rciitrirnsa- Extfiiial xirw <A slicll \aKcs sIhiwIih,; \ariali(iii in cdliir pallcrii. Scale line - 2 cm.

mande ca\it\-. When the foot is protruded, tlie tentacles
touch all its sides, forming a protecti\e latticedike stnic-
ture around it.

Cilian currents on a wide area of the mantle surface
are ventralK- directed toward txvo main rejection tracts,
one on each \entral margin of the mantle lobes. The
main rejection tracts drixe and accumulate psentlcjiaeces

into the xcntral canal; this latter protects undesirable
material from being washed back into the mantle i-ham-
ber (Figure 9). Upon accnmnlalion of a gixcn amount
ol pseudotaeces. the adductor muscles snddeuK contract
to pusli watci" out ol tin' mantle ca\it\ causing the si-
|ihonal membranes to open and allowing pseudolaeces
to be eliminated through the incurrent siphon.

Page Ifi

THE NAUTILUS. Vol. 116. No. 1

Figure 3. Tiiclti nittiicusd. Iiitriiial \U'\\ (if tlie riij;lit shell
\;iKe, .slumiiig tlic pallia! .siiuis anil the scars (il the adductor
muscles; an. auterior adihutor nuiscle scar; /), pallia! liue; pa.
posterior adductor uuiscle scar; ps. palli;il sinus. Scale liue =
3 cm.

Mu.sculature: The anterior adductor mu.scle is larger
liiiui ll]c postciior one. The anterior pedal retractor
nuiseles are inserted into the shell \al\e posterior to and
at a short distance from the anterior adductor nuisele.
Some ol its fibers spread superficially into tlie anterior
and dorsal region of the foot, while most spread in pos-
terior direction: thest' latter have tlu^r t'xtremities in the
foot intcrnallv to the fibers coining IVoni the posterior
pedal retractor muscle. DistalK, within tlie foot, the fi-
bers from both sides join together.

The posterior ped;il retractor muscles are inserted in

Figure 4. Tiuhi i,;,fn,,n,i luterual views of the shell
valves, details of teelli Top. left \aKc; bottom, right \aKc
Scale line = I cm

Figure 5. I'iiflii loitiiriKd FulK exleiided incuneut siphon
projecting out ol the shell \al\es, showing the intricate dispo-
sition of the branched tentacles. Scale line = 0.2 cm.

the shell \al\e in .uitciior direction and dorsalK to the
posterior ;iddnctiii" muscle. The letractor nuKScles on
each side unite inuler the pericardiiun, where bundles
of fibers Imm the light nnrscle cross with those from the
left one. Where the nuiscK's enter the foot, their bundles
once more dixcige and their fibers radiate in anterior
directinn ;uiil xenli.ilK nitn ihc loot.

The pedal retnictor umscles constituti" the extrinsic
muscuhiture ol tin' loot. In ;id(litiou to the extrinsic nnis-
culalurc. ;i \;n-\ing amimnl ol intrinsic nnisciilature is
also picscul in the pidximal ;md dist;il portion ol (he
loot. Innnedi.ilrix liclciw the epithchnni iil the \iscer.ii
mass in the pi()\im;il p;ut ol the loot there are circuhu'
muscle strands lh;it run in ;mtcro-posterior direction and
completeK suiT'ound llie \isccr.il in;iss. The distal Iree
[)art ol the hiot is patki'd with bundles ol fibers running
transversiilK, ihe same occiuiing in llie \iscci;[l nuiss,
wlieic transverse libel's ;u"e less nnnicnins.

Foot: The loot is hirge. wcdge-slKi|)ed. ;uid fileralK
comjiressed, which laiilitales burrowing in I'ehitivcK soft
substiiituni, where il di<fs with rlixllunie movenicnfs. .\

\\\ Xarclii and F. (]i Dario. 2002

l'ae<* r

sm

Figure 7. In i In i nifriidsd Internal \ icu iil llir ci[)('i]lii<j;s ol
the ineiirreiit ami r\riinent siphons in the mantle ea\it\': f/.v.
additional siphonal nienihrane: in. inenrrenl siphon; ins, ineur-
lerit siphonal nienihrane; c.v. excnrrcnt siphon; ]>ri, posterior
atldnetor nnisele; shl siplioiuil nnisele. Se;ile line = 1)5 em.

Figure 6. Tiiclii vcntiicosn. ExttMuletl sipli(ais seen Irom
al)o\e. Top, diaijrani emphasizini; disposition ol tiii' 5 sets ol
tentacles of the inenrrent siphon; c.v. exciirrent siphon; in. in-
eurrent siphon; I, II III. I\'. and \'. sets oi tentacles. Scale line
= 0.5 cm; liottoni, detail of the openiini; ol the exenrrent si-
phon showing the openinii nK'inliniiie and tentacles. Sc;ili' line
= 0.2 cm.

its.'ir

e(iiii|

)letfl\

specimen ol 4.S (.■iii shell lenu;tli hiiried it
in 7 minutes.

.Specimens h\e compIeleK liuried at a depth ol ap-
pro.ximatelx" 5 em: olten specimens can lu' limnd with
tlie shell pointiii'j; out nl the substratum.

Ctenidia (Figures 8, 10-12): The shape ol the cte-
nidiii and the current patterns on them are illusti;ited in
Figures 8 and 10. Each ctenidium is Formed l)\ t\ui de-
mihranchs, the inner demihranch hi<j;lier than the outer
one, particularK in anterior direction, and i:;ioo\ed iilon'j;
its free margin.

Tlie lamellae olliotli dtanihranchs are plicate ;md liet-
erorhalxlie (Figure 1 1 ). Plicae are present alonii the larg-
er part of both lamellae, ranging from a mininmm ol 29

to a maxiiinmi ol (iU in the outt-i' and imiei' demibr;uiehs.
The filaments (Figure 12 1 bear lateio-trontal (fi |Ji.m
long) and frontal cilia (10 |jLni long), the latter replaced
bx' increasingk- longer terminal cilia (uj-) to 40 |xm long)
in the distal free edge of the filament. Throughout the
lateral sides of the filaments lateral cilia produce a pow-
erfid respiratoiA and feeding current.

On tlie ascending lamella ol the outer demihiMiieh.
downward eili;ir\ eiiirents were obsened on ;ill filaments
of the eicsls iiiid lioiighs of the plicae, (.'ollected mate-
lial is iiKiiiiK eon\e\{'d around the free margin of the
outer ilemibraneh onto its descending lamelhie where
frontal cilia e;iri-\ |i;irticles onto an acceptance, month-
directed current on the etenidi;il .i\es.

There is no food groo\e along the free edge ol the
outer demibranch, but a group of terminal cilia along
the oiilei' fiee delleets large particles in anterior direc-
tion. These p.iilicles are carried lor a short distance
;iloiig the free iiKUgin. and then tr;uislerred to the outer
surface of the iiiiiiM- demibranch. On the Irontal region
of the outer deinibrancli tliere are \t'n large cilia that
cle;m the ctenidium b\ lenuning partit'les as large ;is
sand ^niins,

'file lroiit;il cilian eiirri'iits on both l.miell.ie ol the
inner demibranch are almost alwa\s directed toward the
marginal grooxc, e\ce]ition made to thv proximal region
of the descending Limellae, where cilia be;il in dors;il
direction to eoii\e\ material to the aecept;nice tract
aloii'j, the elellidium ;ixis.

Labial Palps (I'igiiri's 13, 14): 'fhe l.ibi.il palps ari'
flat, triangular, with the inner faces tleepK plicate and
the outer faces smooth (Figure 13). The \cntral tips ol

F;l<TL' IS

THE NAUTILUS. \ol. 116, Xo. 1

U

ex

Figiirt' S. Titi'la icnlrirosii. Oiwaus of thf iiiHiitlc ( a\it\ \iiwr(l Ikhii llir hit side alter R'liiox.il (iltlic Iclt shell \.i\w .mil Ictt in. mile
lobe; (III. iuiteiior adductor iiinscle: ti.s. atklitional siplioiial iiieiiilnane; ex. cxcuiTenl siphon; /. loot; id. inner deniihianeli; ilp. inner labi;il
palp; ill. incniTcnt siphon: m. mantle edije; oil. outer deniilinmeh; nip. outer labial palp; ji/i. posterior ;i(lductor ninscle; sr/c. supra-axial
extension ol the outer ileniil)r;uieli: //. iiuilio; i, \entiiele. Anows iudiciite the direetiou ol the eurrents. Se.ile line = 2 em.

tilt' most anterior filaments of the inner deniiliniiuli ;ire
inserted into and Insed Id the (list. [I m-.il ;j,iii()\e ni ihe
palps. On tiie antero-doisal ri'tjion nl the external snr-
faees ot tlie palps, ei!iar\- enrrents move particles aionnd
tlie dorsal margin to the internal sinfaees. The following
ciliai"\- currents on the |)alps (I'"ignre I li weic I'eeog-
nized:

1. Acceptance currents (a) that condnct parti( les to-
ward the nionth I'roni one crest of the plic-ae to liie
next ib: dois.illv directed on the ahorai surface of e;ic'h
plica, and eL on the lateral region of the folds, a vvu-
trally directed current is present on the crest of eacli
ridi^e.

2, Hej(eiii,M lurrenls id), on the lloor of troughs he-
txveen .idj.ieeni plleiie :ui(l (e) on the lower half of the
sides oi til'- plii .1 ^\her(■ particles are driven onto the
smooth \<(iir,il eil'^e .)f ihe p.ilps. Here, a strong rejec-
tion ciM-renI e. ,uiir,els m;ili'ri;il towards the free tip of
the palp.

Mn.scnlar, as •ai I! :is i ilian ;icti\itx- nmst he taken into
account when tin lelion of the laliial pali^s in sortimj;

lood particles is considered. The palps ol T. vcninrosa
;U(' i"<'hiti\el\ ;ieli\{'. The plicae on the iiuiei" snii.iee nia\
he erected or \mi.\ Hal. ;ni(l the p.ilps uui\ lie contractetl
and twisted into ;i spii;il shape, h I'i n ti;i n 'j; th<' smooth xcn-
ti;il ed<j;e .uul icspeelixc lejet-lion c'nrrent in contael with
the ridged sui'hice.

When siu;ill amounts ol [Kulieles .tie presented to the
p;ilps. little sciiting (;ikes phiee. The lidges are I'elaxed
and oM'rlap. Ivxposed to the ;iccept;mce currents, the
majoritx ol [)articles are c.uried rapidK' toward the
month.

Laige amounts of iiarticles on the palj)s induce dil-
lereut deijrees of nmsenlar acti\it\; which results in o[i-
tinud soiling ellieiencw I'.xcess ui;itei"i;il e;uises ,i nms-
enlar reaction In contraction ol the organs into ;i s])iral
shape and most material to he ri>jeete(l.

.Suliuiilted to moderate amounts ol uiateri;il. the \ar-
ions sorting anil icjectiou cmicnts on the plie;ie are ex-
posed. Large particles fiilling into the troughs ol the
gro()\es are rejected, while ihe sm;dler ones ;ii-e trans-

W. Narelii and F. di Dario, 2002

Faw 19

sae

Figure 9. Tinlii icntricosti Iiiiu-r siirlaci' of the ri^lit inaiitlt'
lobe alter reni()\;il nf the lelt shell \al\e and part i)f tiie left
mantle lobe sjiowing eiiian- cieansiiis; currents and additional
siphonal membranes. Arrows indicate the direction ol the cur-
rents. Scale line = 2 cm.

porti'il In the sortiiiLi; currents: oiiK pari (il the original
material reatiies tlie oral i^rodxe hetween tlic palps.

Alimeiitai-\ Canal (Figures 15-lfi): A short isoph-
ai;us opens nito the anterior wall ol the stomach. The
cornhined st^le sac and intestine opens into the poster-
oxt'iitral region ol the stomaeli. and passes l)ack"\\ard and
downward as a wider tulie into the loot. The mid'j;nt
continues Ironi the distal end ol that widei' tube, coils
tisihtk about fixe times on the xt^ntral side ol the stom-
ach, then passes downwartl and back\\ard to ascend as
die hintl<j;ut in posterior direction to the st\le sac. TIr'
hindL:;ut passes through the pericartliuni, wheic it is sur-
rcjuudcd 1)\ the ventricle antl posterior aoitie bulb, and
lings dorsalK' the posterior adduitoi' muscle, to end in
the anal papilla.

The stomach (Figure 16) has its esophageal opening
defined In a transverse rim. The minor t\phlosole ends
on the right side, close to the orifice of thi^ combined
st\le sac and intestine in the floor ol the stomach. Tlie
major tsphlosole. associated with the intestinal gjoove,
penetrates the stomach and passes loiAxaiii. then down-
ward into the right caecum. Tlie riglit caecum receives
six ducts Irom the digestive diverticula. Emerging iroui
the right caecum, the major tvphlosole passes to the lelt
across the floor of the stomach, forming a flap that pro-
jects in dorsal direction toward the esophageal orifice.
The major tvplilosole then is directed deepK into tlii'
lelt caecum, returns and ends on the caecum opening.
The left caecum receives 4 ducts Irom the digestive di-
vi'rticula. The origin of the intestinal gi'oovc lies on the
lelt anterior floor of the stomach, close to the opening
ol the left caecum.

CiliaiT currents in the lelt caecum diicet jiaitick's

alod

a I id

Figure 10. Tii i-hi i citlntOMt l)i.ii;raimnalic vertical section
through tlie cteiiidia m tin- stvle ol .\tkins ( 19.37b) showing the
direction ol particle IIoh bv llic Irontal cilia: alid. ;LScending
lamellae ol the inner demihnincli: alod. ascending lamellae of
the outer demibrancli: (Uid. descending lamellae o( the inner
deniibranch: dlod. desciMiding lamellae ol the outer demi-
brancli: g. marginal groove ol the inner demibrancli: sar. supra-
avial extension ol tlu' outer deniibraiuii. .Arrows indicate the
direction of the cilian currents.

:iw:iv Irom the orifices ol the ducts earning them toward
the intestinal groove.

A group ol 5 ducts lioin the digestive diverticula en-
ters the left pouch. A soiling area King on the floor of
this pouch prolongs onto the right side ol the stomach,
where il exjiands and lornis a bi-aded swelling. Cilia on
this sorting area beat backward along the grooves and
awav Irom the openings ol tile ducts.

.\ wi'll-deveioped dorsal hood projects upwards Irom
the rool ol the stomach ami cunes over toward the left
side. .\ sorting ari'a on its rool and anterior wall prolongs
over tlie esojihageal opening, then on tlie right wail of
the stomach to finisii posterior to tiie orifice of the right
cai.'cinii. Material rejected Irom this sorting area is
passed into a rejection tract, wiiicli arises witliin the dor-
sal hood and discliarges into the intestinal groove. Tlie
:interior margin ol the rejection tract is fornu'd hv a fold
v\ilh sevenii small ridges.

'I'lie gastric shield covers an extensive area on tlii' left
wall ol the stomach, and sends a flange into the opening
ol die dorsal liood, and anotiier into tlic lelt poucii. Vhv

FiU'e 20

THE NAUTILUS, Vol. IIH. N(

%«^

d>ill§§

Fij^urc 11. Tin'Ia vcnlricDsa. Tninvfisf sixtioiis cil tlic outi-r
(l('iMii)iaiKli. Top, niL'clian plicate, lieterorhabdic condition of
the ctcnidiuni. Scale line = 0.1 nun; center, detail of the plicate
deniihrancli. Scale line = ().2.t nnn: bottom, di.stal non-plicate
condition ol (lie onter deiniliiancli. Scale line = 0.1 nnn.

^

l^l«,

s — "'

/ I ' / ^ I

h\

Figure 12. Til , /
Ix'ft. diagram oftlH
rij;Iit. detail of two li'
current; /r. froiilal cili,
re,, larjie frontal cili;i; /'
direction ol tlie ciliarv cnrn

ilialion p;ittern on clenidia.

' ' snri;ice ol the ctenidinni;

• llie lat(Mal cilia; a. oral

'i.i, lfi\ laterolrontiil cilia;

'la. .Vrrow.s in(lic;ite the

.. • lini- = .50 (iin.

Figure 13. iiitld imlricdsa. Cilian cnrreiits on the labial
palp.s. Top, external .surface of the labial palp; bottom, intennil
.sin'face of the labial palp; din. dorsal mary;in: (7/j, inner labial
palp; ()//j, outer labial palp; viii. Nfntnil margin. Sc;de line = 3
cm.

ga.stric sliicld li:is ;i small toiitli-likc projection ;it tlic an-
terior eiul

There is a .sorting area on the anterior wall ol the
storn;ieh. het\\t'en the aperture of the esophagus and tlie
tr;ins\-erse section ol the intestin;il grooxe. (Mlia on this
area beat upuaril, toward the esophagus. This sorting
area extends to the right caeeuni. where cilia heat along
the grooxes from rigiit to left. ;uul proh.iliK eon\e\ p;u"-
ticlt's toward tlie dor.sal hood.

i:)is(;ussK)N

Tilda I'ciilricii.sii is restricted to sand\ heaches direetiv
exposed to wave action. The species is no( lound in shel-

^^....i-i iuV-:y':-'i''Vi

Figure 14. Tiirlti vciitricosd. Diagnnnnwtic representation
ol the ciliinA mecli;nnsnis on the lolded inner snrhice of the
l;ibi;il palp, sliouint; the \;irj(ins cili;ir\ tnicts. .\nlcri(ir is ;il Icif

W. Narchi and F. di Daiio. 2002

Pat'c

Figure 15. T'ncUi icntrircsii AliiiicntaiA caiiaL sirii Ikhii
tlic lelt side; nil. anterior atkliictor iiiiiscle; up. anal papilla: </(/.
dii;esti\e di\erticula: ilh. dorsal hood; c. esoplianns f. foot; /ii,'.
liindgiit; mo. mouth; pa. posterior adduetoi' ninselr; ; . ventri-
cle: .s.v, st)'le sac; .st. stomach. Scale line = 4 cm.

tcrt'd lia\s or other en\ironin('nts lacking strong; wave
action. Tiiis sii<itie.sts that a liiti;li tleifree of :u'ration,
made possihk' l)\ stroii'j; niixiiiti at the uater/air iiitei-
lace, i.s essential to tlie species. (loderje (1950) lound T.
vcntiico.'ia on open seashores, co-occurring witli the \e-
nerid hWdWv Aini/inti'i piirpunitiis (Lamarck, ISIO). Til-
da rciitriciisa. like Tilda stultonnn (Mawe, 1S2.'5), as
tlescrilieil h\ Wevmoudi (1920, 1923), is exposed to con-
stant sin-f pounding, whicli its thick shell is ahle to with-
stand without cracking. The species apparentk thrixes
in what might he c'onsiilered as an inil;i\i)r:il)le cnxiron-
ment.

The large, liea\A', massi\e shell ol 7' Kiiliicusa ex-
ceeds in size all other species of Tilda occurring oil tlie
Brazilian coast. The largest specimen collected during
the piesent stndx' (7.9 cm in li'ngth: 7.4 cm in lieigiit.
and 5.2 cm in width) exceeds the largest one registered
by Gofferje (1950) (3.5 cm in lengdi, 3.0 cm in height,
and 0.9 in widdi).

Armstrong (1965) uiulertook sonic experiiiiciits willi
10 species of'bi\ai\es, including Tilda slullaniin. .Study-
ing the position of the animals in relation to die w:i\c
front and tlieir lielia\ior in the sulistratinii. tli:it .uitlicn
found that the direction of \va\e action is related to the
//( sitit orientation of that species. Such behavior was not
observed in tliis stmK' for T. icatriccsa. nor lor 7'. inac-
froidrs (Bom, 177.S) '.studied by Narchi tl972i. .Ml lol-
lowing references to T. mactroidcs are from Narchi
(1972) unless othenvise noted.

The siphons of" T! iciithcosa aw ol T\pe 15 of Voiige
(1948, 1982). The extremek moliile siphonal membrauc
of the excurrent siphon interferes with the flux of water
passing through the ctcnidia: when opened and extend-
ed outwaiil, the membrane controls and directs the I'x-
halant current. .As observed in T luactnudcs. there is no

leiitiKiilar ring loiined In the iiicdian numtk- fold
around the excurrent :ind incurrent :ipi-rtiires in T. icii-
Irifiisa

The siplioiis of 7' iciilricdsa are similar in stmctiire
and shape lo diose described lor T. sluUoniiii (W'ev-
iiioiilh. 1920. 192.3) antl T iiiaclroidc.s. The incurrent
siphon in T. icnlricosa has branched tentacles disposed
in 5 sets surrounding the a])ertine, unlike T miHiroidvs
where tliev are disposed in 3 sets. Thi' excurrent open-
ing of i. icnlricosa has about 48 simple tentacles, which
are darkK jiigmented at tluir l);ises, wliile in T. inactro-
idc.s the excnrrenl opening teiitacli's are :ibout 20. and
niilkv while in eoloi.

Narchi :iiid I .opes (19981 recorded for tin- hrst time
the presence of the additional sijihonal membranes in T
icnlricosa. In this sjiecies, the tx\o additional sijilional
membranes e.ui be elev:ited and angled toward each oth-
er isolating llie pseiidolaixes ami functioning a,s a txpical
\vaste caiKil. :is Kellogg (1915) descrilied for different
species of bivalves.

The siphonal meiiibr:ines iiuiv be raised lo IreeK ad-
mit the iuciirreiil sli"e;im. or iiiav be drawn {k)wnward
to direct the stream low.ird the inantli' edges, and away
from the gills, such as when much sediment is present.
In this last configuration, a ri'lativeK' large amount of
sediment would be dejiosited ventralK on the mantle,
and i|iiicklv lr:iiispiirled posteiiorK.

In T icnlricosa. the siphonal membranes control the
water current that passes through the mantle caxitx;
while the additional folds can be elevated and angled
toward each other until tliev meet in the summit of and
enclosing the inciirrent siphonal opening. This is proi)-
ablv an adapl:itioii to large amounts of snsjiended sedi-
ment in the water bv directing pailicles awa\ from the
ctenidia and aiding in tluir rcmoxal.

.As seen in some N'eneridae (.\nsell, 1961), the tree
edge,s of the m:mlle li.ive lour lolds. .According to Yonge
(1957) and .\nsell (1961). the presence of four folds is
accomplished In the duplication of the middle mantle
fold.

.\ large n ber ol brancheil tentacles in the margin

of die iiKinlle was described tor T. cra.ssatdoidcs (Kel-
logg, 1915) and T niactroidcs. In T icnlricosa there are
;ilso branched tent:icles ])rolecting the organs ol the
mantle c:ivitv Iroiii inli-usion of large particles ol sand
and possible injiuv. Manv specimens showed torn or re-
jxiired oiilei (leiiiibi'.mchs as apjiarent conse(|iiences of
such injnn, as (aien'in and Narchi (2000) also siiowed
for I'rololhnca ( Lcnkoniai pcclorina (Lamarck, 1818).

The surface of the mantle in T. vcntiico.ta has \-en-
trallv direcled cle:iiising cilian ciuTent.s. similar to that
observed bv .\iisell lOOL in soni<> Wneroidea fnjm
Kngland and in T inaclroidcs. In T icnlricosa aiul T
niaclroidcs. minor cilian cleansing currents pass from
the free margin of the mantle in the posterior region,
passing radialK inward to join the main rejection tract.

.Ansell (1961) described some \ariation in outer de-
mibranch confignration, betxveen species, and e\-en be-
txxeen specimens of the same species due to the strength

Pa-e 22

THE NAUTILUS, \ol. 116, No. 1

sa6

af

Figure 16. Tivda ventrirosti. Interior of stonuicii. seen From rigiit side alter opening bv ineision in tiie right wall: rif. anterior
lokl: lis, beaded swellings; chl. digesti\e diverticnla; (III. dorsal Jiood; c, esophagns: gs, gastie shield: ig, intestinal grooxe: Ir. left
caecum; /;;, left pouch; ml, minor t\phlosole: n: right caecum; nii. mouth ring; sr/,. principal sorting are;i ot the d(>rs;il hood; sii,„
sorting area of the lel't pouch; «;-, sorting area below esophageal orifice; .v.v, style sac; ty. t\plilosole.

of tlip current at the free edge earning particles tow aids
the palps. .According to Ansel! (1961), Vcocnz/j/s aitrca
(Gmelin, 1791 . \'(>unij)is rhoinhoidcs (Pennant. 1777).
Mijsia innlfila Peimint. 1777) and members of Vt';i(/\
Linne, 175S aiit; Di'^iuia Scopoli, 1777, present an in-
cipient current directed toward the month at tlie free
edge of die oiiii i de.rl'DJiincli.

The pattern and general paths ol the etenidiacnrrents
of T. ventricosa are i:; agreement witli the results ol
Ridewood (190.3) and Atkins (19.37a. b) on the mor-

[)hol(ig\ of the ctenidia :uid their cihation in some \'e-
neroidea. Tlie ctenidia in T. ventricosa belong to T\pe
V.{\) of .'\tkins (1937b\ with one groove along the free
ventral margin of the iniu-i' deniibranch. as \;ui'hi
(1972) and Xarchi and Ciibrii'lH ( 19S()> described for tlie
venerids Aiioniahx'iinlid hnisiliana iCmelin, 1791) and
Chionc suhroslrdid (Lamarck, 1818). respectively. In T.
Iliad roidcs the eteiiitli;il eiliation is of T\pe C(2) of At-
kins i 1937b).

The hibi;il palps of T i ml ricosn sliiiw the s;mie b;isic

W; Xairlii and F. di Dario. 2002

Paw 23

features of ntlier species oIN'eneroidea studied In Tliiele
(1886), and are also similar in stnieture and niuseular
acti\it\ to tliose of T iiiiictroidf.s. The \(-ntr,il tips of the
most anterior hlann'iits of tlie inner demiliraneli are in-
serted into and fused to a distal oral groo\c: the asso-
ciation of ctenidia and laliial palps belong to ( ^alegon II
of Stasek (1963).

The configuration of the alimentan canal of 7' icn-
tricosn is similar to that of T niiutrnidcs. differing
uiainK In the nioic coii\oluted midgut, with 5 closeK'
packed coils in the first, anil with oiiK one loose coil in
the latter.

\\'here\er known, the aiiatomx of the stomach is gen-
eralK' similar throughout the X'eneridae. The left and the
right caeca of T. ccnthcosa receive 4 and 6 ducts from
the cligesti\e di\erticula. respecti\elv. Six and .5 ducts
were respectixek' recordetl for T niactnudcs In 7' vni-
tricosa another group of fi\e tlucts from the digestive
diverticula enters the li'ft poucli without being associ-
ated with the major t^plilosole or the intestinal gnxne,
as described h\ Furchou 1 19fS()) for C.ajninum iniiiiniiiin
(Montagu, fS47) and 7' iiiiictniiilcs.

The stomach of 7' iciitiifasd has the structmc of a
tvpical suspension-feeding cnlamellibranch. It is similar
to those of the vfuerids Olossus liuiiKniWi (I.-inne, 1758),
G. iniiiiviiiin. \Cnii\ ciisiiKi Linne. 1758. and 7" iiKKiro-
idi'.s, as described In Owen (1953). Purchon (1960). .\u-
sell (1961), and Narchi (1972), respectiveK. Within the
stomach, food particles are kept in motion In thi' com-
bined action of the rotating cnstalline stvie and the cil-
iated walls, and are subjected to sorting maiuK in the
posterior sorting area, as in T. luactroidcs. The stomacli
of T. ventricosa luav be capable of handling luanv par-
tick'S at one time diii' lo lis complexity.

.AC'KXOWLKDC^MUXTS

To Conselho Xacional de I.5esen\c)l\imento (,'icutifico e
Tecnologico (CNPf|). Bra/il. for the Support (irant nniii-
ber 300490.

LITER.ATUHK (TFKD

Alilidtt. 1-i, T U)74 .\iiiriR'aii .Seashells. 2ikI, Kdilion \an \i)s-
traiid l^einliold. New York. 663 pp.

Ansell, -\. D. 1961. Tlie tiuictional rnoipliologx ol Bnlisli spe-
cies of \'eneracea (Eulaniellihranchia). Joiini.il ol llir M.i-
rine Binlotjical .\ssociatiiiTi (if the United Kiii^diiiii )li2 :
4Sy-.517

Arinstrcina;, L. H. i9(i.5. liurniwiiin liiiiitatidiis in I'llcrspnda.
The Wliiier 7: 19,5-2I)(1,

Atkins. D. 1937a. On the cilian inechanisnis and interrelation-
ships of laniellii)ranchs. Part II: Sortint; devices on the
gills. QuarterK- Journal ol Mitroscopical Science N..S. 79:

Atkins. D. 1937li, On the eilian mechanisms and interrelation-
ships of Lamelliliranch. Part III: T\pes of fnnellihraneii
gills and their food currents. Quarterl\ Jdurnal ol Micro-
scopical Science N.S. 79: 37.5-421.

Broggi, J. 1970. Sobre el hallazgo de ciuco nue\ as especies para

el Qnerandino Unignavo. Comunicaciones de hi .Soeieckid
.\I:ilaeoloiiiea del Uruguay 2iIS»: 427—131

Hneku[i. I., 19.57. C'atalogo dos moliiscos do .\Iusen Piiim:m-
(k'lise de ( liencias .Naturais. Ilieringia 1: 1—10.

Carcelles, \ K 1944. (.'atalogo de los moluscos marinos de
Puerto One(iuen. Re\ista del Museo de Pa Plata, \ne\a
Seiie. Zoolot;ia 3: 2.53~.509.

Carcelles. A. H. and S. 1. Williamson. 1951. Catalogo de los
moluscos marinos de hi Prorincia Ma<;,illaiiica. Ke\istadel
Instituto Xacional de Investigacion de las ( ienc i:is Natnr-
ales2l5): 22.5~.3S3.

Coan. K. \'. I' Nalenfich Scott and F. R. Bernard. 2000. Bi\aKe
seashells ol western North .America. Marine l)i\alve mol-
liisks From .\rctic .\laska to Baja California. Santa Barbara
Museum of Xatunil Histon- Mono<;rapli 2, Sant:i Barbara,
\iii + 764 pp.

Golterje, C. .\. 1950. CJoutnhui^ao a zoosjeografia ila makico-
fauna do litoral do estado do Parana, .-^ninivos do Museu
Paranaense 8: 221-2S2.

Gneron. C. O. C. and W. Narchi. 2000. .\natona:i luncional de
Prototluicii (Lctikttnui) pectunna (Lamarck. ISIS) (Bi\al-
\na: Wneridae). Re\ista Brasileira de Zoologia 17: 1007-
I ( i: >9

Kello'.;^. I I,, 1915. ( !ilian, mechanisms ol lamellihranchs with
desenptiiiii (j| anatonn. journal ol .Moi]iIiolog\ 26: 62.5-
701.

Lauije de .Morretes. F. 1949. Ensiiio tie cat:ilogo dos moluscos
do Brasil. ,-\njiii\os do .Museu Paranaense 7(1): .5-216.

Narchi. W. 1972. Comparati\e stndx" of the hmctioiuil mor-
pliolot^ ol AiuiiiudiHiirdiii hrasiliiinii ((anelin. 1791 :ind
Tivcld iiinciroidi's (Born. 177S! (BivaKia. \cneridae Bul-
letin ol Marine Science 22: 643-670,

\:irc 111. W, and M. .\. CJahiielli. 19S0. Sobre aiiatomia lunc'ional
de CliioiK stdirostratii (Lamarck, ISIS). Re\ista Xordes-
tina de Biologia 3: 25-46.

Narchi. W. and S. G. B. C. Lopes. I99S. On the meiirreiil
siplional membrane oiTivcla vcniricoso (C^raw IS3St (Bi-
vaKia: \'eneridae). Jn: Bielen R and P. M. .Mikkelsen
(eds.) .-\hstracts of the World (a)ngress of .\lalacok)gv.
Washington. D.C.. PJOS. Unitas -Malacolos;ica and .\mer-
icaii Makuologieal .So(ietx. \\'asliini;ton. p. 2.37.

Owen. G. 19.5.3. On the hiologv ol Glitssiis humiinus i L. i i/.v-
ociirdki cor Lam.'. |oiinial ol the Marine Biolosjical .Vs-
.sociation ol the United Kingdom .32: S.5-106.

Purchon. R. I). 19(iO. The stomach in the Eiilamellihr.iniiiia:
stoiiLieli l\|ies l\ .iiid \. Proc(>e(lings ol the /.oolot;ical
Sotielv of London 1.35:.!): 431-4.S9.

Ridewixid. W C 190,5, On the structure ol die >;ills ol the
Lamelliliraiicliia. Philosophical Transactions ol the Ro\al
Societv of Loiukin Ser. B. 195: 147-2S4.

Bios. I'', (.'. 1970. (aiastal Brazilian Seashells. I'uiidavao Cidade
do Kid Gr:iiide, Fundayao I'nivcrsid.ule do Rio (iraiide.
Rio Cr.iiide. 2.55 pp. 60 pis.

Rios. L. C. 1975. Bra/ili:in Mariiii' Mollnsks Iconoiiiaphv. I'iiii-
dayao Universidade do Rio (inmde. Rio (a'aiide. 331 pp..
91 pis.

Rios. F. C. 19S5. .Seashells of Bni/if Isl, edition. l-iiiida(,'ao
Cidade do Rio (iraiide. F'uiida(,ao Universidade do \\w
(Jrande, Miiseu Oceaiiografico. Rio Grande. 32S pp., 102

Rios, L. C. 1994. Se:isliells ol Brazil, 2iid, edition, l-undavao
Cidatk' do Rio Grande, Instituto Ac()ua R.J.. Museu
Oceaiiografico Prof E,('. Rios. Univ(>rsidade do Rio
Grande, Rio Grande, .36S pp., I 13 pis.

Paee 24

THE NAUTILUS, \ol. llfi. NO. 1

StiLsek. C, H. 1963. S\niopsis and cliscnssion of the association

of ctenidia and labial pal|)s in tlic l)i\aKcd Mollnsca. The

\eliger 6: 91-97.
Thiele, J. 1S.S6. Die Mundlappen der Laiiullihi.uichiaten.Zeit-

sclirif't fiir wissenschal'tliclie Zoologie 44: 239-272, 2 pLs.
WVviiiouth, F. W. 1920. The edible elanis, nuissels and scallops

of Calilornia. Hnlletin ol the (California State Fish and

Game CConi mission 4: 3-72, 19 pis
Wevmouth, F. VV. 1923. The lile-histon and u,ro\\lli ol the IMs-

mo clam {Tirclti stitllonitn Mawe). Bnlletin of the Cali-
fornia State Fish and dami' Connnission 7: 5-120.

Von<j;e. (;. .\1. f948. Formation of siphons in I>amellibraneliia.
" Nature 161: 198-199.

Yonsie, (;. M. 1957. Mantle fusion in the 1 ,ainellibranehia. f'ub-
blica/.ioni de la Sta/ioue '/ooiot;ica di Napoli 29: 151-171.

Yonge, C M. 1982. Mantle inar<;ius with a re\'ision of siphonal
t\pes in the Bi\aKia. |onru:il ol Miilluscaii Studies 48:
102-103.

Notice

THE 2002 R. T. .\BBOTT VISITING CURATOHSlilP

Tlic |-5ail('\ -Matthews Shell .Vluseuin is pleased to invite applications lor the 2002 R. T. .\liliott Nisitiin^ ( Jiratorsliip.

The Ciiratorship. e.stablisliecl oriwinallv in accordance with tlie wislies ol the late Dr. R. Tucker Abbott, Foundint;
Director of the Shell Museum, is awarded annually to enable mollusk .s\steiiiatists to \isit tlie museum for a period
of one week. .Abbott Fellows will be expected, by performing collection-based research, to assist with tlii' curation of
porticjns ol the Museum's collection and to provide one evening talk for the general public. The Museiun collection
consists of Tuarine, freshwater, and terrestrial specimens. A large percentage of oiu" hoklings have been catalogui'd
tln'ougli a coniputerizeil database managi-ment svstem. .\ substantial portion of the time will be available for research
in the colk'ction, but field work in southwest Floritla can be arranged. The R. T. Abbott \isiting (!iu-atorship is
accompanied bv a stipend of $1,500.

Interc\sted malaeologists are invited to send a copv ol tluir ciiiiieuhnn vitae together with a letti'r tletailiu'j; their areas
of taxonomic e.xperti.se and resi.'arch objectives, and to pinvide a tentative subjei-t for tluir talk. Si'ud materials to:

Dr. Jose 11. Leal, Director

The Bailev-Matthews Shell Museum

P.O. Box ioSO

Sanibel, Fi,. 3.3957

jleal@shelhmiseum.org

.Applications for the 2002 X'isiting Curatorship should be sent no Liter than .Mav .30. 2002. The aw:nd will be aimouni'ed
by late June. Questions about the \'isitin<j; ( 'malorship slmnld be sent to tlie e-mail address above, or liv piinne at:

(941) 39.5-22.33: lax (941) .395-0706.

THK NAUTILUS 1 Ifii ] ):25-:3I. 2002

Paw 25

Use of HMDS (hexamethvldisilazane) to dn- omiiiic
microstructures in etched bixaK e niollusk and barnacle shells

Beind R. Schone'

Dt'purtiiieiit of Geosciences
Universitrv" of Ariz.ona
Tucson, AZ 85721 USA
l)enKl..schoene@excite.coiii

Da\id Bentley

l5i(it('(liiioloin,- Imaging Facilit\
Uiii\cisih ol' Arizona
Tucson, A/ S5723 USA

ABSTRACT

The organic framework of molfnscan and iiarnacie sjiclls \ields
clues to Ijiocalcification processes. Sliglit demineraiization of
the shells reveals the fragile meshwdrjs of insoluble organic
fillers and membranes, which tend to collapse, wrinkle, and
shrink when air-dried from water Gomparison of different diT-
ing teclmiijnes on etched bi\al\e mollnsk iChionc flti(tifr(i'j,ti)
and baniacle shells (Chtlunudlus sp.) reveals that hexanietli\kl-
isilazane (HMDS) produced results qualitatiscK superior to
critical point dning or dning from eth\l alcohol or water
H.MDS dries stnictural details of the organic iiicsliwork e\ccl-
lentK and facilitates the recognition of faint growth nicrements
for growth pattern analysis (sclerochronolog\). The H.MDS
method is cost-effecti\'e, sa\es time and can lie used as a rou-
tine substitute for drying microstructures in slightK etched
molluscan and baniacle shells.

INTHODl CflOX

Organic matter plaws an es.sential inle in the formation
of molhi.scan and baniacle shells. It has been realized
that the organic matri.x proxitles the structural frame-
work ("template"; Clark, 1980) for hiomineralization and
influences the mineralogical and cA'stallographic prop-
erties (Mann, 1983: Simkiss and \\'ilhur, 1989; Cren-
shaw, 1990 and literature therein; W'atahe et al., 1993).
Demineraiization of skeletal hard parts mixcils the un-
derKing three-dimensional organic microstrmtures,
which ma\' vield clues to hiocalcificatioii processes.

Organic matter is also a major constituent of some
growth increments (e.g., Koike, 1986). Slight deminer-
aiization of cross-sectioned molhiscan and InuiKicle
shells re\eals a three-dimensional relief of insoluhli' or-
ganic components and dilferentialK dissoKcd cnstals (as
a result of different crxstal sizes and orientation). In
sclerochronological studies (growth ;iual\ses), supei'fieial
etching is commonly used to aid in the identification and
measurement of internal growtli ini'rements in niollns-

' Current address: Institute and Musiuni lor Ceologx :ind I'a-
laeontologN, |ohaiin Wolfgang (ioethc L!in\ersity. Senckeii-
berganlage 32-34, 60054 Frankfurt am .Main, (iermany.

can and Inuiiacle skeletons (Khoads and Lutz, 1980;
Scluine et al., in pi'ess). The etching time varies for ilif-
terent species and depends on, hir instance, the shell
strnctinv, mineralogv, and organic content. .Although
growth patterns in mollnsk shells are the focus of im-
ineious studies. ouK lew papers deal with the growth
patterns of harnack's. Unliki' most crustaceans, lianiacies
do not replace their hardparts. Both mollnsks (e.g.. Dav-
enport. 1938; Fannelhi and .MacCJlintock. 1968) and bar-
nacles (e.g., Bourget, 1980) grow bv periodic accretion
of skeletal material producing circadian growth incre-
ments (see sketches in Figure 1 and 2: direction of
growth). In baniacles, the growth lavers are best \iewed
in the she;itli Liscr (I'"igure I), and in most bi\alM- mol-
lusks in the outer shell laxcr (Figure 2i.

Like most solt tissues, the shell organic framework,
including the orgauii--rich growth increments obseiM'd
in cross-sections, is jirone to collapse, shrinkage, and
wrinkling when ;iir-dried le.g., .Anderson, 1951; Nation,
1983; Clark, 1980, 1999). Hre\cnting these unwanted ef-
fects requires special cluMiiical treatment, which (k>h\-
drates and hardens the fragile organic structures. Several
techniques are used to di\ biological solt tissues, .al-
though extremeK time-consuming and quite dangerous
(highl\- pressurized chamber), critical point drviug
(CPD) is b\ far the most common method (Anderson,
1951: using liquid COj, e.g., Clark. 1980 or Freon 13 as
a transitional fluid, e.g.. Koik(\ 1986). On average, pre-
paring one sample In WD re(juires full attention o\er
1.5 hours. The basic (.'PD e(juipment costs se\-eral thou-
sand dollars, (iood results were also achiexcd with the
sublimation deli\di;uit Peldri II ( K(>n!ied\' et al.. 1989).
lIowcM'r, prepar;itiou lollowiug this technique takes
more than t^\■ice the time as (;PD, and Peldri II is no
longer axaihible because of environmental hiizards. Flu-
ids with low surface tension (ac-etone or propxlene oxide,
Bovde and Wood. 1969) sometimes produce reasonable,
artifact-free results for biological soft tissu(>s. Some
workers prefer the extremeK hazardous osmium tetrox-
ide technique iOnattlebaum and Carner, 19801.

.\ reliable ;uid sim]ile dning techni(|ue. which pro-
duces results (|u;ilit;iti\el\ eompai:ible or superior to

Pac{e 2tt

THE \AUTILUS. \<)1. 116. Xo. 1

2 mm

2 mm /

\

/

cross-section

'^

parietes

Figures 3-5

1 cm

Fieures 14-16

''^

inner shell
surface

\

Figures 6-8

commissure line (outer rim)

Figures 9-13

cross-section

B. R. ScIkhk-

D. Bcntlfx. 2002

Paffl"

CPD. was introduced In Nation (19S3). After tlelndra-
tion in a series ot graded ethanol solutions, tlie samples
are innnersed in hexamethvl-1, 1. 1. 3, 3. 3 disilazane
I hexanieth\klisikvzane. HMDS), an organic reagent with
tlie clieniical formula [(CH5),Si],NH, for approxiniateK'
fi\e to 30 minutes, and allowed to air-dn' at room tem-
perature. The low-cost HMDS dning technifjue does
not require full attention during processing. It was suc-
cessf ulK applied to dn- \arious kinds of" soft parts in dif-
ferent organism groups, e.g., in insects (Nation. 1983;
Rumph and Turner. 199S). soft tissues of niollusks (.Leal
and Simone, 1998), \ertebrates (Heegaard et al.. 1986;
\\'er\ha et al.. 1990), microorganisms (Dekker et A..
1991; Hochherg and Lit\aitis. 2000) or pollen (Chissoe
et al., 1994). .\lthough ;ipplied to deniineralized human
teeth (Perdigao et al., 1995; CaniJlio et al.. 1996). to
our knowledge HMDS h;is not been used as a transi-
tional soKent for dning molluscan and baniacle shell
microstnictures.

This stniK' e\aluates the usefulness of ll.MDS for lin-
ing microstnictures in etched barnacle and bi\ al\e mol-
lusk shells using scanning electron microscopy (SEM).
Special emphasis is gi\en to the use of HMDS in scler-
ochronological studies. We compare the HMDS tech-
nique to 1) the CPD method. 2) ;iir-dning immediately
after etching (.\IR), and 3) a combination of dehydration
in a series of graded alcohol baths and ;iir-di"\ing i.\LC).

MATERI.\LS AND METHODS

Material and general preparation: Durint; field
trips in 1999 and 2000. we collected biirnacles and bi-
\iJ\'e mollusks in the intertidal zone of the northern Gulf
of California, Me>dco. For this stud\, we used one bar-
nacle specimen (Chthainahis sp.) collected ali\e at Isla
Sacatosa in December 2000 (N31 29.60, \M14 50.85;
specimen no. ST11-A18), three Cltionr fiiiclifrana (Sow-
erbv, 1853) specimens collected ali\e at North Orca in
November 1999 (N31 32.60, WH4 52.78; specimen no.
NO3-A105, NO3-A106, NO3-A108), and three C. fluc-
tifms^a specimens from Isla Sacatosa (N3I 29.60. \\ 114
50..8.5; specimen no. ST12-D1. .ST12-D2. ST12-D3; De-
cember 2000). All specimens are housed at the Depart-
ment of Geosciences, The Uni\ersit) of .Arizona. Clollec-
tion Flessa, Subcollection Schone.

.After remo\al of the soft parts, all specimens were
cleaned in Chlorox (5.25'7r soilium Inpochlorite) to dis-
sol\e remaining superficial organic matter, rinsed \\ith
water and dehvdrated with ethyl alcohol (EtOH). Coat-
ing with ]-B KWIK Wekr^' pro\ides a protecting co\er
for the shells during cutting. The fragile barnacle re-

(juireil I'mbcdding in |-B" Epow. |-B° Epow is not ad-
vcrseK affected b\ immersion in EtOH or H.MDS.

Cross-sections of Ijarnaclc and lji>al\e inolliisk
shells: We obtained three cross-.sectioned slabs (thick-
ness 0.5 uun) from the barnacle shell (ST11-A18; Figure
1) and from the left \aKe of each of the three C. fluc-
nfmoa shells no. ST12-D1, -D2. and -D3 (Figure 2) us-
ing a low speed (Buehler'^' Isomet') saw. Use of an ul-
tratliin saw blade (0.4 mm thickness) assured that the
effects of different preparation techniques (see below)
on the organic microstnictures could be studied on id-
most identic;il portions of the shell. In this stud\ we
focus on the composite prismatic shell la\cr of C. flttc-
tifraga. because the\ are of primarv' concern in sclero-
chronological studies. In the cross-,sectioned shells of C
fnctifni'^a growth increments are clearK' de\eloped and
easv to measiue. Moreo\er, etching and dning results
ma\' \an' with respect to different hpes of shell inicro-
stiTicture. Focusing on one tvpe of shell la\er allows
comparisons of the effecti\ eness of different dr\ing
techniques on similar shell microstnictures. The cross-
sections were mounted on petrographic slides, ground
on glass plates (600 and 1000 grit -AbO,, powder), and
polished on laps (9, 6, and 0.3 |jLm .Al^O, powder). Ul-
trasonic cleaning bet\\een e;ich polishing step assured
that no grinding powiler was left on tlie surface.

Inner and outer sliell suriaces of the bivalve: Shell
fragments of three C. Jiiictifraga specimens (NO3-A105.
-.A106. and -.A 108) were mounted on a petrographic slide
to expose the growing surface at the inner shell edge.
Two shell fragments of specimen ST12-D3 were mount-
ed on petrographic slides with die outer surface exposed.

Chemical treatment: etching and dicing: All sam-
ples were then etched in a 0.25 inol EDT .A solution (eth-
\lene diamine tetraacetate. pH 7.95. buffered with
NaOH) and carefulK rinsed in de-ionized water The
shells w-ere etclied for \aning amounts of time (Table
1 ) to obtiiin different degrees of demineralization: shglit
supei-ficial etching and decalcification of the upper 1 mm
of shell niat(Mial. In previous studies we canied out a
series of tests and anakyed the effects of an array ot
etching times and EDTA concentrations on shells of C".
j{uctifm<i(i and Chthainahis sp. (Schone et al., 2002. and
unpublished data). For the presentation of microstnic-
tures in C. fiuciifraga and Chtliaiiuihis sp. the approach
used herein pnned to be most appropriate. We com-
pared the following lour dr\ing techniques (Table 1); air-
dning from water (AIRi. dning from EtOH (.ALC). dn-
in<r from H.MDS and CPD. For ALC. HMDS, and CPD

Figures 1. 2. Positions in the shelLs. wliere tlie samples (Figures 3-16 1 were taken, d.^s; = direction ot growth. cUrecfion ,n winch
sulisequent crrowth increments were added. 1. Cross-section through the p;iri.-tes (shell plates) of tiie lianuicle Chthamalu.s sp. The
sheath laver (distal portion of the parietes) exhibits faint dailv growtli stnictiires T-igiires .3-.5). 2. Chiomj{m-t if rn<ia sheW. Samples
for Fienres 6-S were t;iken from tlie grouiiii; edge of tlie inner shell surface, samples for Figures 14-l(i from tlie outer sliell

lecome appan-nt in the outer shell laser (Figures 9-13).

lor rignres

surface. In radiiil cross-sections, growth p.itteins

Pa.'e 2S

THF NAUTILUS. \()1. 116. NO

B. R. Stlioiie and D. Bentlex. 2002

Page 29

the samples were rinsed in a sfrit's of graded EtOH
(40%. BC^f, 80f7r, 5 min each; hvo times lOO'-L 30 iiiin
eacli). The HMDS technique inchides iiniiicrsion ol the
dt'h\ih'ated samples in 97'^?- hexametln Idisilazane
lAldrich'^' chemical; two times for 30 min each! and
subsecjuent air-dning. CPD was performetl in ;i Polaron
CPD E32()0 apparatus. The delndrated samples were
placed in the CPD apparatus with alcohol, the chamber
cooled to 10°C, and CO^ allowed to enter the chamber
Free alcohol was flushed from the chamber until dn' ice
was seen exiting. The sample remained in CO, for 45
minutes flushing the chamber e\"en' 15 minutes. At the
end of the exchange, the chamber was sealed and the
temperature raised above 35°C (critical temperature for
CO. = 32°C). CO^ \-apor was then released.

Coaling for SEM and photographic documentation:

.Vlimiinnm stubs were glni'd to the pctrogr;iphic
shdes. The samples were sputter coated immediateK
with a 30 nm gold la\er in a Bio Rad Polaron Di\ision
SEM coating svstem. The samples were stndietl with a
Cambridge Instruments Stereosciin 120 SEM at accel-
eration x'oltages of 15 and 25 ke\' Black and white pho-
tographs were taken on Polaroid Positi\"e/Negati\e
4X5" T\pe 55 Instant Sheet Film toi- ilocumentation.

RESULTS

Barnacle shell microstructure: Under 750X mag-
nihcation. the 15-min-etched cross-section of the sheath
[lortion of the barnacle parietes (Figure I) sliows partly
dissohcd calcite cnstals in a crisp three-diniensio]ial
mesh of organic fibers when dried from li.MDS (Figure
3). CPD results in a less well-dried organic framework
(Figure 4). Apparently, some delicate organic fibers col-
lapsed resulting in a slightK' bhurcd image. Even worse
is the drying in ALC samples, l^resniiuibly. as a result of
a complete collapse of the organic iiiatiix, single ciystals
can bareh' be discerned (Figure 5). Some growth layers
disappear parth or entirely when immediateK air-dried
after etchiuE; (Fiiiure 5).

Bi^al^e shell microstructure:

Inner shell sire.vce near nii < ommfsm ki : I! \il )S

treatment of the slightK etclieil growing surface at the
iimer shell surface (Figure 2! reveals a thin organic
mesliwork residue in between ;md on top of the polyg-
onal pattern of the composite prismatic stnictnre (Fig-
ure 6). CPD-treated shells are not depicted, because the
results are companible to the HMDS d(>siccation. .\L(J
produces a soiuewhat collajised oi'ganic framework (Fig-
ure 7). The rod-like surface is compIcti'K coxcred in
.\IR samples (Figure S).

CROSS-SEfTlON, Ol TER COMPOSIl E PRISNLVIIC SMELL
UWER: .\ cross-section etched for 15 min and dried
fn)m H.MDS clearK shows the composite prismatic
structure of the outer shell layer iFigures 2. 9). Organic
membranes of daih' and even subdaiK' growth layers are
intersected b\ peipendicular running fibrous mem-
branes (Figure 9; for exiierimental studies on the timing
of growth pattern foiination in (' flii(iifra<ia see Scluine
et al.. 2002). CDP dries the organic mesliwork less well
(Figure 10 i.

.\t lower magnification (375Xi the differences be-
txveen the HMDS (Figure 11 > ;md C:PD i Figure 12)
techniques are less olnions. .\ reli;ible comitini; ;mcl
measurement of narrow dailx growth increments laid
down during liot sunmier conditions can be conducted
on HMDS ;ind CPD s;implcs. but b;ircl\ accomplished
on die sample treated with the .M.C method (Figure
13!. In the latter case the collapsetl organic matter cov-
ers most parts of the faint growth patterns ;ind does not
allow for detailed growth pattern ;inalysis.

Outer shell surf.voe: Outer shell surfaces exposed
for t\velve hours to the 0.25 mol buffered EDT.\ solu-
tion reveal a well-dried, dense, nnilti-layered organic
network when dried from H.MDS i Fignr(> 14). Growth
lines cannot be seen iin\ more, .\pplying the .MR meth-
od, the organic fnunework collapses and covers the
shell's microstructures as a mass without clear structures
(Figure 15). Durintj; the vacuum in the sputter coater

Figures .3-16. SENi iscanniixj; electmii iiiicroscopv ) inuincs iil .tclieil hdn\M\f •ClillKnndlii-, sp.) ( Fis^ires 3-5) and bivalve moliusks
shells (Cliit'iiv fiuctifraoa) (Figures (i-t6). Working (list;iTice in iniu iWDi. atccleratioii vnltasje in ke\' l.W), ptcliing time in min
(ET), and scale are'provided in cacli fit;iire. .•}. Best diving of the orijanic mesliwork (\> and single cnstals (cs) is achieved when
the etclied barnacle shells are immersed in H.MDS prior to air-diviim. I'mminent lid-ies are daih iiiwvtli lines (dgi!. 4. CPD (critical
point dniiia) dries the organic network paitlv. hiil the sinjile ciyst;ils cannot be discerned well .5. .Ml structural details are gone
when using^'ALC (;iir-dning from cthvl ;ikohi')b. Some faint Growth lines iell over to iidjacent srrovvth lines (arrow). 6. The grovving
edge of the inner suriac* exliihits crisp poKnoiuil cnstal structures in the H.MDS treateil specimen. 7. CPD dries most of die faint
structures as well. 8. ALC results in a blurred image. 9. Dailv and subdailv (sgi) sriovvth increments, organic membranes (onii as
will as peipendicular ruimint; orijanic fibers (of) are well dried in H.MDS-treated cross-sections aU'Mmc fluctifras.(L 10. but less
well 111 CPD samples. 11. Daih' <;iovvth lines (diji) laid down iliiriii-.; hot smuiiier ;iie best viewed in HMDS or 12. CPD prepared
cniss-sections. 1,3. ALC does not'allow for a detailetl analvsis. because organic structures colkipse. 14. Outer shell snriaces of C/n«/K'
fillet if rw^a reveal a dense, three-dimensional insoluble cailaiiic meshwork when immersed in HMDS pnor to air-dning. 1.5. .-MK
(air-dnine;! results in an undifferentiated mass of collapsed oiaaiiic stnictures. 16. \\\mS treated samples mav shrink and wrinkle
under'vacuum during the coating process or in the SEM dies = non-etched slu II siiri;ice. don = demineralized organic network).

p. me 30

THE NAUTILUS, \ol. llfi. \(

Table 1. 0\enie\v ol' s;imi|)1c licatmcnt.

l'R'ser\atioii t

ecliuiinie

Species Spi'eliiirn # l-'tcliinjj; time

HMDS

CPD

ALC

MB

Chllwlamiis fastis

ST 11 -A 18

13 mill

Figure 3

Chione jiuc1ifr(i<!fi

N03-A1().5

3 mill

Figure 6

NO3-A108

3 mill

NO3-A106

3 mill

ST12-D1

15 mill

Figures ^

STliTW

12 li

Fii'ures 1

Fisrure 4

9, 11 Figures 10, 12

14, K-i

Figure 5
Figure 7
Fiiiure 13

Figure S
Fisiiire 15

and the SKM, however, the organie nehxork .slieet wrin-
kles and shrink,s (Figure 16j.

DISCUSSION

Acl^anla^es of llio HMDS leclinitjue: Iniinersion in
II.MUS prior to air-(h"\iiig th'ie.s more clearlv the true
arrangement of the insohihle organic microstmctures in
etelied l)i\al\e inollusk and liarnaek' sheiks than ck)es the
eomnionK' used (,'PD teehniijue. CieneralK, the organic
h-amework of slightK' etched cross-sections or etched
surface samples is well presencd in its original three-
dimensional condition when dried lioiii HMDS. This
dning (jnalit^• allows high precision anaKsis of the struc-
tural organic framework in the shells.

CJrowth patterns in IIMDS-treati'd, slightK deniiner-
ali/.ed shells appear innch clearer than in AIR or ALC
l)rocessed specimens. In the latter cases, narrow growth
increments are eonipletel\- covered l)v collapsed organic
material. Under the hiirdeu of the collapsed organic
incshwork, fine and organie-rich iiicrcinents ma\' fall
ox'er to adjacent growth increments. Tiicsi' ad\erse ef-
fects prexent sclerochronologists from counting and
measuring the growth increments correctK. In speci-
mens dried from IIMJ^S, e\'en snh-daiK grnwili patti^ns
can be ri-cogni/ed with ease.

II.VIDS dr\ing does not re([uire I'xpensivc lab e(|nip-
mcnt or additional skills like CTD. Sample preparation
is ea.sy, takes only a few minules and does not re(|iiirt'
constant monitoring during the immersion or the dning
process. Moreoxcr, the sample size is not restricted In
the small vacuum chamber like in the CPI) apparatus.
but only by diffusion constraints. II.VIDS dmng is less
dangerous than CPD, becau.se it (kies not re(|nire high
pressures ilining ]-)rocessiug. which ma\' represent some
potential vonrc( of hazard to the operator.

Disadvanlages of ilic FiMDS technique: Ihick or-
ganic 1,1'. I is shrink and uriiikle. These artifacts max re-
sult Iroi.i the :,nmersion in HMDS itself or from the

high xacuiiui '..;,•:
lem is also v i-m
soft tissues f '■ , '
ditional fixatm
ingand HMDS ;
organic mesliwork i
maliu re|i()rtedK ha

• caimg and SEM. The latter prob-

X.: 1. r CPD and Peldri II treated

■ ■ ' .!.. llWfii. It is not clear if ad-

^ . iii iriatri.x prior to delivdrat-

' ■ III staliilize and harden the

'! .. 'I liniiian dentine with for-

s ; ' ■ -I I'lli'cts on the stnicdiral

arrangement of the organic fabric (Canalho et ak, 1996).
We assume that longer infiltration and dning times max'
reduce the aforementioned, unxvanted effects.

HMDS is a hazardous chemical. Its potential effects
on humans and on the enxironment are barelx' knoxxn
(Material Safety Data Sheet). It is stronglx' adxised to
use the chemical onK under a fume hood and to prexent
inhahng or skin contact.

CONCLUSIONS

HMDS dning is a xen- useful, loxx-cost, easx-to-appK
and time saxing alternatixe to other knoxxii dning tech-
ui(jues for soft tissues. This publication demonstrates the
applicability of this fast dning solxent in sclerochronol-
og\' and for the studx' of organic microstructm-es in some
bivalxe mollusk and barnacle shells. With H.MDS. xxe
aehiexed results ([iialitatixelx superior to the xxidelx' ap-
plied CPD. Further studies should determine if the
HMDS method can be used as a routine teclmicjue in
dning organic stnietures of etched mollnsean and bar-
nacle shells and if the HMDS method can also be ap-
plied to etched accretionan hard parts of other marine
or terrestrial organisms.

,'U:kn{)\\lei:)C.ments

This studx has been made possible bv a postdoctoral
seholarshi]') (Lxnen program) bx the .Alexander-xon-
iiimiboldt foundation (to BHS) and a NSF grant EAR
US().5ifi5 (to Prof. Karl W. Flessa. Department of Ceo-
seiences. Unixt'isilx o( \ii/(iiia. Tucson, AZS5721). We
ari' grateful to Kail W. i-1essa, Daxid L. Dettman, .An-
drew S. Cohen and i^axid II. (Joodxxiu (Department of
CJeosciences, Unixersitx ol Arizona. Tucson. ,\Z S5721 ),
and jose 11. Leal for helphil discussions and to txvo
anonxinons rcniexvers who improxed the manuscript.
This is CT'IA.M ((.'enlio de F.stndios de .Ahnejas Muertast
publication no. 40. The authors herexvilh declare that
the expel iiiieiits are not in xiolaliiin of the i-iirreiit I nit-
ed Stales laws.

LITKHA! I !;!•: CflKD

.Xiidi'isdii, 4". !•'. U).5I . 4'c(iiiiii|iics lor llie ])i('senaticiii ol tiiree-
(limeiisioii.il slriuliirc in piepariiiii spciiiiiiiis lor llir licc-

B. R. Schcine and D. Bentlev. 2002

Pa^e 31

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of Sciences 13: 130-134.

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Davenport, C. B. 193S. CJrowtli lines in l<issil pcctens as in-
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Dekker, N P.. ( .' ]. Lammel and G. F. Brooks. 1991. Scanning
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Heegaard, S., O. A. Jensen and |. U. Prause. 1986. Hexame-
thxldisilazane in preparation of retinal tissue tor scanning
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Hochberg, R. and M.K. Litxaitis. 2000. IIexameth\ldisjlaz;ine
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shell ol .Mercenaria mercenaria iLiniiei. Transactions and
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Leal. j. H. and L. R. L. Simone. 1998. Propiliilium niniiiiiin.
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Nation. J. L. 1983. A new method using hexainethvldisilazane
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THE NAUTILUS 116(l):32-.35. 2002

Paw 32

Gone with the wdnd: a pelagic marine species described as an
endemic land snail from the Bahamas

Philippe Bouchet

Museum national d'Histoiif naturclle

53 nil' Buffon

75()()5 Paris. FKAXCF.

pliouclictC'UiMilin.lr

ABSTRACT

AbaconUi luiiijrui^d (('Icntli, 19oSl. hitliiTto assuiiicd tii rcji-
resent a monotvpical genus and species ot Sul)nlinidae (Gas-
tropoda, Pulmonata) endemic to the Bahamas, is shown to l)e
a SMionvm of the marine pelagic snail Litiofxi inehinostoma
Rang, 1829 (Caenogastropoda, Litiopidae). Instances of genera
of land and freshwater ga,stropods originalK mistakenly de-
scribed a,s marine, and \ice \ersa. are re\iewed. Of the 10
names invoK'ed, four are \alid (Daronia. KiiroJns. Tcivtropoiiui)
or potentialK \alid iDi'^onitixis). one (Bn^iulclhi) has been sup-
pressed In the K'ZN, and the other fi\e iAiiiicoiiiil. Afrocilli-
klea. Aiuiploidiniis. Htilliiiujirrsiliii. I'sciiihililnniiii) are s\no-
nxnis.

iXTHonrcTiox

William |. CJench ( 19.>S) ilcserihcd Lcpllntinii {Almciiii-
ia > iHinfm'^u as a new .subgenus and spciics in the piil-
nionate tauilK' Siihiilinidae, hasetl on s[)ec-inH'ii,s eollei't-
ecl bvJ.C;. and V,. C;reeii\va\- and hinisclfin the Balianias
in April 1936. The clescription was aeeornpanicd li\ hlaek
and white photograph.s. The journal wlieic the deseii[)-
tion \va,s published (Metnorias de la Soeii'dad ('ubaiia
de llistoria Natural) uia\' ha\'e contributed to the lack
ot attention that these new taxa ha\c subsecjuentK le-
ceived in the literature: in fact, the name AlxicDiiia is
not nientioned at all bv Zileli (1959-fiO) in the pulmo-
nate xohirne of the standard llaiidhucli <lcr I'dUiozoolo-
ii,w. Ahacoiiia has receiitK' been elevated to genus lexcl
by .Schilevk., (1999: 513,' fig. 666), :uul ehissified :is :i
member '<t the sublamiK Subulininae.

In the oii',:i!ial publicalion, C^^lench noted that "this
species dil'i'i r:. :iuile uol:ibK from all others known to
[hini| in this genus \1 rptinfiria]. The remarkaliK acute
apex and the :lKinge in the sculpture from a.\ial on the
first four \slioiis ;., ,piral on the remaining whorls is
sharply ihtfen-nt IrMui i,ther species, and the thiekeuiug
of the shell is :ils. " 'i-ual hi this genus". He speculated
that Ij-plinrina (A/.,. <.■,■. u(iufm<ia anil two other en-
cleniic pulmonates mijjl i.nstitute a reumaut of a
much older fauna" with V\ .1 Induni relationships. How-
ever, examination of the type materi;il in WV.'A (Figiues

1-3) re\eals tluit Ahdiouid lunifniiid is not :i snbidiuid.
and not e\en a pulmonate, but :i s\iion\ in ol IJIiopa
iiirhiiuistoiiid. d circumtropic;il pelagic eerithioid (Figs
4-6). (dencli and Schilexko luiil properK described tlie
apical sciilptiue oi Abdcoiiid. but had tailed to recognize
its identibi. The protoconch ol Liliopd incldii(>sl(iiiui con-
sists of four whorls with strong a\ial ribs, aliiittiug adax-
ialK on the preceding whorl anil iiiti'rrupted b\ a smooth
band on the shoulder: the protoconcli/teleoconch ilis-
continuih' is nuirked b\ a sinusigera notch lF,iif|ue et ak.
19SS: l.SiZ. figs 1,4).

Liliopd nu'ldnostddid is :iliiiiidant in (he midst ol ll(i:it-
ing Sdr^d.ssiiDi. and it is eas\ to imagine how tlie\ can
be waslieil ashore and blown inlanil atti'r the se;iweed
Iromls base been sun-dried. The species is known to
occur oft the 15alKunas (Liii|iie et ak, 19SS).

.SYSTEMATIC :.S

Class Ciastropoda ( Jnier. 1797
SiiperlamiK' {Jerithioidea l''leiniii'j:.
FamiK Litiopidae Craw 1S47
(Jeiiiis Litiojid Hang, IS29

1S22

No\\ sxnoiiMii:

Ahiicdiiiii Clench. 19'1S. T\pe species: I.cjUiiuirid I Ah/ifimiri'
iiidifniflii (blench. 1938. b\ original desigiuition.

Hoiiihijxiniis lielanger |iii IjCssonj. 1835 is another ;ilreail\ rec-
(ii.;iii/cd s\ii()n\ni o( l.iliopii

l.ilioj)!! iiicldiii'sloNid l^aiig. 1 S29

\o" sMionvni:

I .(■jiliiiiiha lAhiiioiiiai luiufrii'^ii Clench. 19.'i8: 321. pi. 2-4.

figs 1-2.
Ahdconid iiiinj)<iii<i — Scliile\ko. 1999: 514. lig. (i(i(i.

Type Ma(en:i1: llolnlxpe \IC'/ 1 16705: 34 p:init\pes,
\ic'/ 116706: I p,n,il\pcs NIC/ 116707.

Type L<)c:ilil\: S.uid l^;iiik. ( 'rossing Baw (ireat .Abaco

Isknid. B:ili,iiii:is. I'aralvpes froni the l\pe loc:ilit\ ( MC^Z
Il6706):iiid Mores Island (NIC/. 116707i.

DISCUSSION

Shells of kind anil Ireshwater snails :iie occ:isioiKilK
washed to llic sea :uid ha\e mil ml i i-c |iiiiill\ been de-

p. Houclict. 2002

Paffl" 33

Figures l-(). LiUdjiii iiirhiiioslninii Haiit;, IS29, 1-3. Il(il(it\pr u{ Lipliiuiiiii iAIkicoiiki ' iKiuJni'^ii Clciicli. iy3S. MCZ 116705,
SiUKJ Bank, CJrossing Bav, Great Abaco Isiand. Baliaiiias. Heiijlit 4.2 nun. 4-(J. One (if li\c s\nt\pi's fAlJtiopa iiiclaiwsloma Rang,
1S29, MNHN [uncatalogued], "Atlantic Ocean". Bang coll., 1S34. Height 4.6 nnn.

serihcd as marine gastropiids. Imt the rexcrse lias also
occa.sidiialK happened. The awareness oi tliese sitnations
aeross tlie tratlitional boundaries of marine and non-ma-
rine niollnscan literature is problematic, and it ma\ be
uselnl to take the case of the Abaconia to thaw the at-
tention of taxonomists to names tliat might have eseapeil
their attention. These are presented below in the chro-
nological order tliex" were described, ami snmmari/ed in
Table 1. Two of the genera invoKcd e\en became the
types of new faniilv-group names, Tcri'troponiidae
Hochebrnne, 1S81. and Anaplocamidae I^all. 1921. and
it is probable that, if tliex had not been mistaken abont
their habitat, Rochebnme and Dall would not ha\e
lailetl to recognize them as representatixcs ol the fami-
lies Architectonicidae and Pleiu'oeeridae, res]H'cti\cl\.
Of the 10 genera inxoKed. four are \alid (l^arouui, Kii-
roliis. Tcniwpoina) or potentiali\ \alid (DiiSfutiuxis), one
(Bnmdchn) has been suppressed b\ the K^Z.N, and the

othei' fi\e iAlmci'iiid. Alrocdiiiilcd. Auiiplocdmii'i, Bulli-
uopiTsiliii. I's<-ii(liilil)tniiti! are s\nou\ms.

Daronia A. Adams, 1861. Based on a specimen from
the Canning ( .'olleetion. CUfclostirmti i Daronia': spinila
A. Aiiams, ISttl, was described as a new subgenus and
species of niarine gastropod from the Philippines. Waren
and l^oucliel ' I9SS1 showed that it is a South .American
species of the land snail laniiK l^oteriidae. Daronia \.
.\danis, IS()I. being a senior sxiioinni lA' Bncklcyia Hig-
'iius. 1872.

Bromh'lia iJoiirmiignat, 18(52. l^escribed as a new
genus of \nc\lidae. with two species. B. dronctiana
Bourgnignat. 1S.54. (originalK' described based on ma-
terial from the ( fuming collection from "North Ameri-
ca") and B. ^ihhosa Bourgnignat. 1Sfi2. said to be li\ing
on damp rocks in the forest of Kdough. near Bone in
.Alsii'ria. Brondclia remained in the freshwater fainik' .An-

l^uii' 34

THE NAUTILUS, \'ol. 116, No. 1

Tabic 1. XomiiKil genera of marine sjastropocls iarranged alphaliptically) originally described a.s h

snail

s ana Nice \ersa.

XoMiinal "eniis

I^escrilied as

Revised position

lieierenee

Alxiroiiiii C;lench, 193S
Afrocanidca Connolly, 1929
Anuplocamus Dall, 1S96
Bwndchti lionrgnignat. 1862
Biillinopcrsilia F. N'ordsieck, 1972
Daronia .\. Adams, 1861
Di<loniaxis Jousseaumc, 1889
KiiroUis de Folin, 1870
I'scudolilmnUi Stcf'ani, 1870
Teretropotna Rochebrune, 1881

Subnlinidae

fresliwater Bnccinidae

Trichotropidae

.\nc\lidae

.\cteonidae

Cyclostrematidae

Fenissaciidae

Acteonidae

Daiidebaidiidae

Cvclostomatidae

Litiopidae

Ranellidae

Pleuroceridae

Sipbonariidae

FeiTJssaeiitlae

Poteriidae

P\Ta!nidellidae

Fenissaciidae

Pleunjbrancbidae

.Arcbitectonicidae

this papt'i'

Ben (1998)

Rehder (1942»

Marsball (1981)

Smriglio and Mariottua ,1996j

Waren and Bonchet (1988)

Neubert (1998)

Zilch (1959)

Zilch (1985)

Crosse ( 1882)

cxlidae until Marsiiall (1981) recognizetl it as identical
with tlic marine piilmonate WdUamia Monterosato.
liS.S4 [famiK Siphonariidae]. Incidentallv; the name
Broiuh'lia is the senior s\nion\ni, Init it \va,s sub-seqiientlv
suppressed and placed on the Official Index ot Rejected
and In\a!id C.eneric Names in Zoolog)' h\ the ICZN
(1986. Opinion 1410).

Kciroliis de F"olin, 1<S7(). Kiinilii\ primus de FoHn,
1870, was descrihetl a.s a new genus and species of ma-
rine snail trom off Vera Caiiz on the Pacific coast of
C'entral .Vnierica. Kdrolw-: was not originalK placed in a
hiTuiK; and was omitted Irom standard textbooks and
treatises lor neark a centun. It was treated bv Zilch
(1959) as a \alid subgenus ol Ccrilioidcs Fenissac, 1S14
[laTuily FerussaciidaeJ, and h\ Schileyko (1999) as a lull
genus.

Psc'tidolihduia Slefani, 1S79. Daiidchardia larcidina
•Stelam and I'antanelli. 1879, was described as a land
snail Irom (he hills near Tarantn in southern Italy. It was
made tiu' t\]ie species of a section Psriidoldiania Stefani,
1879, ol Daiidchardia Ilai-tmann. 1821 |lamiK Dande-
bardiidae], later elevated to full genus b\ Forcart (1950).
Zilcii (1985) has shown that D. larcittiiia is a svnonvm
ol Siisania tesliidiuaria Cantraine, 1835, and I'srudoli-
hania a .synonym n\' Sii.saiiia Cray. 1857 |laniil\ Fleuio-
brancliidae].

Jeretropnma Rochebrune, I8S1. Inunediatc l\ :ift( r
die description, based on live-taken specimens fiuui the
Dakar peninsula (Senegal), of Tcirlropoiiw pcrricri
Kocheb-rmii'. 1881, as a new genus and species of 'Cv-
clostomace.i'. and type of a new subfamiK- Terelropom-
idae (sic Cross.' 1SS2) recognized it as an architecton-
icid relalet! to lonuia Cray. 1842. Tercimpoma is cur-
rently treated as a \<i!id subgenus of Heliactis d'Orbignv,
1842, and pi-rricn is rcg-arded as an .Mlantic subspecies
of //. infiiiidiliidif.)ri!ii\ fCmclin, 1791) (Bieler, 1993).

Dificminxin Jousseamnr. 1889. Di<i(>iiiaxis houroui<i-
nati Jousseaumc, is^',), ,\as described as a new genus
and species from vi. n .uid trt\ited a.s a land snail of the
family FenissacliiL;- b\ ifs author and bv Zilch (1959:
342). BiLsed on a re-exaniinalion of the t\pe material.
Neubert (1998) realloialed it to the marine famiK l'\r-

amidellidae. Although Schile\ko (1999), relering to Neu-
bert, suggested that onl\ an anatomical stuck would es-
tablish firmk- the systematic jxxsition of D/gofuV/.v/s, the
heterostrophic protoconch lea\es no doubt tliat it is in-
deed a p\ ramidelloid.

Anaplocamiis Dall, 1896. Anaplocamtis horealis
Dall, 1896, was described as a new genus and species ot
Trichotropidae from .Alaska, and later e\en became the
t\pe of the monotvpic famik -\naplocamiclae D;ill, 1921.
Rehder ( 1942) discoxered that this was in fact a SMionvm
of the freshwater snail Aiiciilosa ddatata Conrad, 1835
[now Lcptoxis dilatata: famiK' Pleuroceridae], from east-
ern United States, which had been erroneously labelled
as from a marine .Alaska localitw

Afrociniidea Connolly, 1929. Afrocanidca ^^cmma
C'onnolk. 1929. was desciibed as a new genus and spe-
cies of freshwater Bnccinidae, reputedk' Irom Shimbi
Hills, Kenya, but the holot\pe was shown to be a lanal
shell of Cipuatiiiin imtriciniuii (Roding. 1798) [family
Ranellidae], with Afrocanidca Connolk, 1929, ending up
in s\"n()u\nn- of Cw(^//mi(»//( Mi'irch, 1852 (Beu, 1998 i.

BullinopersiUa F. Nordsieck, 1972. Based on a sin-
gle empt\ shell, BullinopersiUa sjihacroidcs F. Nord-

.■k, 197:

was itescritied as a new genus and species ot

.\cteonidae from shallow water off the Baleares, in the
westeiii Mediterranean. Smriglio and Mariottini (1996)
recogni/ed it as a jii\iaiik' of the land siuiil Fentssacia
folliculus (Clmelin, 1791' [lamily FeiTissaciidae].

IdTER.ATURK CITFD

Ren, A. 1998. Indo-West Pacific Ranellidae. Bnrsidae and Per-
sonidae (Mollnsca: (Jastropoda). A monograph of the New
(Caledonian l.inn.i and revisions of relatetl t;L\a. Memoires
dn \hisenni national d'llistoire naturelle 178: 1-255.

Bieler. H. 199.'V Aidiiteetonicidae of the Indo-Pacific (Mollns-
ca. (iastropod.i). .Vbbandlungen des Natnnvissenseliaf-
tlielien N'ereins in Ilanibnrg. new series, 30: 1-.377.

C'lencli. W. |. I93S. Land and Iresliwater mollusks ol Ciand
Bahama and the .\baco Islands. Bahama Islands, Memo-
rias de la .Sociedad (.iili.ui.i de llistiiri:i Nalnral 12(4):
.3()3-,3.33. pi. 24-25.

("rosse. 11. ISS2. [Book review ol| Sin mi Ivpe nouvian de l;i

p. Boucliet. 2()(»2

Wm- 35

faniille des C'vclostoniaoea, par le Dr. A.T, dc Koclicliniiir
|ournal de tA)iich\liolosjie 30: 249-250,

F'liriart. 1. 1950. S\steniatique dcs MolliiscjUfs en Idiiiic Ai'
Duudcbardia et re\ision des cspetx-s d'.'Viuitolic ct dc I'ilc
de Crete. Journal de Conchvliologie 90: 1 07-1 17.

Luque, A. A., Templado, ]. and L. P. BiirnaN'. HISS. On tlic
systematic position of the p;enera Litiopii Rans;. 1829 and
.\lilhii II. and .\. ,\danis. 1S.5.). Malarol(ii;i(;il Ri'\ii'\\,
suppl, 4: I SO- 19.3.

Nhnsliali, B. A. 1981. The ij;enns Williaiiiia in llir ucstrni Pa-
rific (.Mollusca; Siphonariidae). New ZciI.likI |()iinial ol
Zoologv- 8: 487^92.

Nenbert, E. 1998. Digoniaxi.s h(>nri;,iiii:,iiiiti |ou.ssfaunie 1S89,
a genus and species of the Pvraiiiidellidac ( tlastropoda;
Heterostropha). Archiv Fiir Moliuskcnknnde 127(1-2):
10.3-106.

KIZX [International Commission on Zoolo'^ical Nomencla-
ture]. 1986. Opinion 1410. W'iUUimUi .Monterosato, 1884
(Mollusca, Gastropoda': consencd Biillrtiii ol Zoolo<4ical
Nomenclature 43: 249.

Rauij, S. 1829. Notice sur le Litiopc, Litiopa. iji'un- ii(iu\cau

(Ic Molhisque gasteropode. .\nnales des .Sciences Natu-

rcllcs Hi: .30.3-.307.
liilidii II \ 1942. .\ note on tin- 'icmis AiKiplocniiiiis l.)all.

I'll.' Naiililus .56: 49-.5().
Schilc\ko. A. .\. 1999. 'IVcalise on Hcccul terrestrial pulinonate

molluscs. Part 4. Kutlicnica suppl. 2: 4.35-564.
Snuiglio. C. and I', .Mariottiui. 1996. Central 'IXTrlieuian Sea

.Mollusca: Xi. Description oiCiillosinicon ti/rrlicnicnin sp.

uo\. (Castropoda, Acteonidae) and remarks on the other

.Mediterranean species ol the lamilv .Vileonldae

d'Orhi^nv. 18.35. Basteria 60: lS.3-193.
W'an'n. \ and 1'. Bouchet. 1988, .\ new species of X'anikoridae

Ircini the western Meiliteiiauean. with remarks on the

northeast .Atlantic species ol the taniik. Bollettino Mahi-

ci)lo<iico 24(.5-8): 7.3-100.
Zilch. A. 19.59-1960. Teil 2: F.uthvncina In: Oil. Schiutlcwoir

led. I. Hiiiullmch tier PiiliiozooUi^c. Band 6, Caslnipudd.

Bomtraeger. Berlin, \ii + 8.35 pp.
Zilch, .\. 19S5. Ddudflxirditi ttiri'iitiua und I'sciuhilibcniid i.Nu-

dihranchi:i: N'otaspidea: Pknirohranchidae. .Archix liir

Mcilluskvnknndc 115i4-6.: 291-.30I).

In Mciiioriaiit

Tiulaslii<2;t' Hahe Rus,sel H. jciiscn

1916-2001 191S-2()01

TIIK NATTILUS 116(l):36-38. 2002

Paw- 36

Note

Biogeogniplu" ol the deep-sea gastropod Palazzia plaiiorbis
(Dall, 1927): an uncommon form of rarity

Michael A. Hex

Departiiient ol Biolog\'
Unix'cisih' of Miissacliiisptts
100 Mon-isse\ Blvd.
Boston, MA ()21iT USA
iriiclia<'l.ro\(a'\iiiil).e(lii

Palazziii plaunrhis iDall, 1927) is a iiiiniitf dri'p-sea gas-
tropod ol uncertain plniogenetic atfinitv, possibly resid-
ing in one ot die groups that were once included in the
Archaeogastropoda (W'aren 1991, 1993), It has a plani-
spiral shell hearing distinctive iixial ribs that branch mid-
\va\^ towai'd the outer whorl margins (Figure 1), Like
many deep-sea species, its natural liiston is unknown,
.Vloore (1971) first noted that P. plaiiorliis occupies an
innisnalK broad geographic distribution in the western
North .Vtlantic, lia\ing been dredged at 2967 ni oft Lali-
rador and at 53S ni oil' Florida, W'aren ( 1991, 1993) sub-
sequently reported the species \mu\ west of Iceland at
241 111 and in the eastern North .Atlantic o IT the British
Isles Iroin 2081-2898 in. Here. 1 proxide additional re-
cords lor tlie western and eastern North Atlantic, tidp-
ical Atlantic, and the western South .Atlantic (Tabli' 1).
These new localitv data come Irom m\' examination of
material pro\id(>d In the Woods Hole Benthic .Sampling
Program (Sanders. 1977) and tlie .\tlantic Continental
Slope and Rise Stuck- (Cirassle and Maciolek, 1992). Pal-
azzia planorbis is revcaleil to have an e.xtraordinariK
broad geographic distribution that spans Southern and
Northern Hemispheres IVoin 43°33'S to (i5"ori'N, and
includes both the eastern and western corridois ol the
deep .Atlantic (Table I, Figure 2), Its bath\ nietrit- range
is similarl) impressive, extending IVoni the uppermost
reaches of the continental slope (241 m) to the deep
alnssal plain 521fi m).

While the shell sculpture of Palazzia planorbis ap-
pears to be niiiijue among [)lauispiral deep-sea micro-
gastropods, tliere is some individual variation in the ex-
pression and nnmbcr of axial ribs and the extent of
branching. Ihr p.iltern of geographic variation throngh-
oiit its ranwe is ditlien'L to quantifV becaii.se of the scar-
city ol material inu wid ■ separation of sampling locali-
ties (Table 1. i'lenn 2 However, the specimen from
the eastern tropica! At! nitu shown in Figure 1 is re-
markably similar in uiik at.i.Kc to those fimired bv War-
en (1991: 77, fig, ih .\-lji iliat were collected in the
we.stern and eastern Nurih Atlanli 913 to 3259 m ,slial-
lower and 45° to 70° t( ihe n.nlh.

Its enormous geograpliic and ballivnietric ranges are

espeeiallv notable because Palazzia jAanorhis is vei"v
rare, and is included among the smallest gastropods.
Specimens from the Woods Hole Benthic Sampling Pro-
gram (Table 1) were all collected bv using epibenthic
sleds (Hessler and Sanders, 1967), which are txpicallv
tovvetl for (jue kilometer on the bottom and vield sam-
ples that represent about HHIO linear nr of seafloor. As
a rough approximation, the data in Table 1 suggest a
median densitv of one individual per 1000 nr, with a
maximum of 17 individuals per 1000 m- in the western
South Atlantic. This is rare, even for deep-sea gastropods
which usuallv exist at low abundance (Rex et al., 1990;
Rex and Etter. 1998). With a shell diameter reaching
onlv between one and txvo inillimcti'rs, it is also an ex-
tremely small gastropod — bolli lor the (Gastropoda as a
vv'hole (Bieler and Mikkelseii, 1998) and tor the dee^p-
sea fauna (Rex and Etter, 1998),

Ri'latioiishijis among geographic range size, abun-
dance and both size are central to macroecologV'. Al-
though a grt'at ileal ol variation has been obseived,
abundance and bodv size are generallv related to one
another negativclv, and related to range size positivelv
(Caston, J994; Brown, 1995; Gaston and Bhickbimi,
2000), Thus the coinbiiKition ol low densitv, large geo-
graphic nmge and small bodv si/e obsciAed in Palazzia
])l(iiu)rl)is is exceptional. Biibinowitz i, I98|l proposed ;i
threc-w;iv (.-kissifieiition ol raritv based on geographic dis-
tiibntiiin, habit;it speeificitx and local population size.
Palazzia planorbis. with its extensive geographic range
in the Atlantic, oceup;ition ol dillerent li;ibitats ideptli
zones) and small inral populations, represents one ol till'
most muonmioii lorms ol raiitv in the Ivabiiiowitz
scheme. Schoenei' (1987' releried to low ii'hilive ;ibnn-
(kmce throughout a s|H'cii"s rangi' as siillusive raritv. and
showed (hat this p;illeni is an extremi' euil-membi'r ol
the r;iritv coiilinimiii.

\\ idelv distributed rare spi'cies are also known among
dei'p-se;i protobranch bivalves. Examples include Pris-
liillouw iiitois and /' alba (Sanders and .Alien, 1973), :ind
Yiildirlla fabula (Allen et al., 1995). Other bivalves are
connnon in one or moi'e deep-sea b:isins and rare else-
where (.Allen .ind Sanders. U)96). a pallcin that Schoe-

M. A. Hc\. 2002

Paec

Fiijiiro I. I'lilm-iii plunoiiiis collected Iidiii station U)5 m
the casttTii tropieal Atlantic (si'l- Talilf 1 Im l(i(alit\ data' Tin-
specimen measures 1.24 mm in ina\niiuni (lianielei.

nei' llVJ.ST) teniR'tl tlittusi^e rafitx. 'Hie iiiciili'iiee ol lare
deep-sea gastropods oceHp\iii'4 ludad geographic ranges
is still unclear since most taxononiic s\ntlieses, siicli as
Boncliet and W'arens (19S0) excellent monograph, haxc
been regional in scope.

Studies of raiitx' and I'ange delineation are liaiiglit
with imt'ertaiiit\ and methodolosrical diificnlties (Ciaston

U)U I It should he eaiitioned that the apparent aliun-
danee ol I'dldz-Jd j}ltiiiorl}i\ iTahle 1) nia\" be paitK" due
to sampling error. The mesh size of the collecting net
used on epibenthic sleds deploxed in the Woods Mole
Beiithie ,Sam|)ling Program is 1,(1 mm. Hiis is fine
enough to e.iplure adults. Smaller jn\enil<'s could pass
through the mesh, but some do in lact appear in sam-
pk's. The siexc si/e used to sort the niacroiauna is 420
fjLm. whii'h should retain .ill but the smallest post-lanal
iudi\idiials ol this spi'cies. Karlier sampling. lio\ve\'er,
used larger mesh and sii'Xi' sizes, suggesting that rec'0\'-
er\ of /' pliiiiorhis was happenstance. .Since the samples
aie widi'K separated and lepiesent onl\ fi\e regions of
the Atlantic I'lable 1, Figur(> 2), we cannot doeiiinent
the full geographic range or know relatisc abundance
th.roughout the range. It is possible that R pUiiiorhis has
an unrecognized ceiilei' of high ileiisitx' and shows dif-
fusi\c lather than suffusixc raiitx" (sensw Schoener.
19S7). l'"iuall\. it should be pointed out that, while V.
phiiioiiiis appears tii be a coherent morjiliospecies. some
pheuot\]iicalI\ unifoim deep-sea mollusks show strong
population structure at the genetic level (Etter et ah,
19991, /' jiliiiinrhis mas be a coiuplex of geneticallv di-
\ergent populations or species. E\cn considering these
potential probleiiis, the combination of a Pan-.Atlantic
distribution, consencd moiplioiogw low abundance and
siriall size would seeiu to represent an extremeK' unusual
form of raritx lor lu.niue mollusks.

Table 1. Localitv' data for Palazzia plaii(>ii)i\ in the ,\tlantic. Where the source of data is not indicated, it refers to this study.
.XCSAK refers to the Atlantic Continental Slope and Hise StuiK (Grassle and .Maeiolek. f9!)2i. WliOl refers to the Woods Hole
Benthie Sampling Program (Sanders. 19771. and is followed In the name of the \essel and iruisi- immher. Please see references
listed under source for information on other ((illcelm^ exprditions

Kegion

Source

lv\pe(fitiou

Station

l)e[)thilll

F.lUdule

Fuiigitude

Specimens

West,

111 North .\tlaiitie

Wareii.

]m.\

|(ni lidgason. ( .'oil.

—

241

fi.5°0fi' N

26°42' W

Moore.

1971

\U 1 1,1)1 ic

—

20(i7

55°3fi' N

54°33' W

WllOI/CllMN fOf-i

334

4400

4()°43.3' \

46°14.2' W

6

ACSAK

13

12.50

39°48.35' N

70° .54.94' W

WIIOl/Aii.WTis II

24

12(S

:5S0(i

39°37.25' N

66°45.55' W

\\ll()l,/K\oHH .3.5

.34(1

.3310

.3S°l(i.O' N

70°21.55 W

WIlOf/AirwTis 11

to

17.5

KiSO

.36°.36.()' N

fiS°3()' W

Moore.

197f

.\l,H\rHciss

26riS

.5:5S

.3(f5S' N

79°3S' W

Faste

rn North ,\tlaiitie

Waren.

f99.3

INCAI,

DSOl

2001

.57°59.5' N

1()°40.5' W

Waren.

199.3

INCAF

I)S()2

20S1

.57°58.7' N

10°48.9' W

\\"aren.

1993

INCAF

nS(l5

2.503

.5(i°27.9' N

ini.s' W

Waren,

1993

INCAI.

DSOti

2494

.56°26.3' N

irio.fi' w

Waren.

1991

INCAF

CPOo

2SS4

.5.5°0().4' N

12°30.2' W

Waren.

1993

Cn,\i.i,i:N(a n II
Wll01/f:ii\i\ 10(1

190
:52,'5

2 SOS
.3:547

.54°4F N
.5()°()S.3' N

12°1S' W
|.3°.52.3' W

W.'sti

irii lYopieal ,\tlaiitie

W ll()i,K\oKH 15

295

1011

S°()4.2' N

.54=2 F3' W

W IIOl/.Vri.WTis II

31

1.50

SS7

7"5S.()' S

.34°22.0' W

Fast(

•rii Tropital .\tfiiitic

WIIOI/.Vri.WTis II

42

lOS

45(s:5

I(r26.5' S

9°()6.5' F

\\II(M/.\tl\\tis II

42

197

459tt

10"29' S

9°04' E

WllOI/.Vri WHS II

42

195

3797

14^45' S

9°55' E

West.

erii South .\tlaiitic

WIlOI/Ari.WTis II fit)

239

l(i70

.36°49.()' S

.5.3° 15.4' W

WII()l/.\Ti WHS II (ill

247

.52 Ki

4.3°33.()' S

4S°5S.l' W

17

Pa<ie 3S

THE NAUTILUS, Nol. 116, \,

Figure 2. I'lic (listrilnitiiin ol P(il<r.-Jii jihnutrhis in llic .'\t-
liUitif (.sec 'l"al)le 1 lor stiitioii data). Oirclfd nuiiibeis represe-nt
the niinihcr ol samples xieldinij P pldiiorhis in that region.
( .'ircles wiliioul nuiiiliers represent iiiilividual samples.

.'\fK\0\\I,Kn(:\IF,\TS

I thank .\iulrea iiex, Hon Kttcr, .\nders W'aivn. Pliiiippe
iioiichi't, and ('arol Stuart lor llieir eoniiiicnts on tiiis
paper, jack ('ook lor the map of the Xtlantie and Maria
r'apiisja lor iielpintj to prepare the manuscript. The ma-
terial from the Woods Hole Bcnthic Sampliuii Pronrani
wa.s provided by Howard Sanders, and From the .-\tlantii-
Continental Slope and Hise Stud\ In James Blake. This
research \va.s .supported 1)\ the National Seienee l-'oun-
dalion CIrant C)( :i':-9:3()l(iS7 and In llie Uni\eisit\ ol'
.Vlas.saclinsetts.

iJTEIl.VrUHE CITED

•\1!( n. J. ,\ and II. L. Sanders. 1996. The zootjeoijraphv. di-
M'liii'. liici orinin ol' the deep-sea protobraneh hixakes ol

the Atlantic: The epilo^ne. Progress in Oceanography .38:
95-15.3.

Allen. J, A., H. L. Sanders and F. J. Hannah. 1995. Studies on
deep-sea Protohranchia (BixaKiaJ: the subfamily Yoldiel-
linae. Bulletin of the British .Museum (Natural Histor\),
Zoolog;) 61: 11-90.

Bieler, R. and P. M. Mikkelsen. 199S. Aiiiinoniccni in Florida:
notes on the smallest li\ing gastropotl in the United States
and comments on other species of Omalogxridae i Het-
erobranchia). The Nautilus 111: 1-12.

Bouchet, P. and A. VV'aren. 19S(). Rexision of the north-east
,'^tlantic bathyal and abxssal Turridae (Mollusca, Gastro-
poda). The |()\n"nal ol Mollusean Studies. Supplement S:
1-119.

Brown, |. H. 1995. Macroi'cologN. Uni\(Msit\()f Chicago Press,
( 'hicago, 269pp.

Ftter R. ]!, M. A. Rex, M. C. Chase and J. W. Quattro. 1999.
A genetic dimension to deep-sea biodi\ersitx. Deep-Sea
Research I 46: 1095-1099.

{".uston. K. |. 1994. Raritv. C:liapman and Hall, Foiulon. 205

PP
Caston. K. J. and T. .\1. Blackburn. 2000. Pattern and process

in macroecolog\. Blaek^well Science. Oxiord. 377 pp.

(irassle, J. F. and N. J. Maciolek. 1992. Deep-sea species rich-
ness: regional and local di\ersitx' estimates Irom quanti-
tative bottom samples. American Natundist 139: 313-341.

Hessler, R. R. and H. L. Sanders. 1967. Faunal di\ersit\' in the
deep-sea. Deep-Sea Research 14: 65-7S.

Moore, D. R. 1971. .\ deep water Omtiltnj^ijid in the wt'stern
.'Vtlantic. The Nautilus 84: 113-117.

Rabinowitz, D. 1981. Se\eu forms of raritv. hi: Svnge, H. (ed.)
The biological aspects ol rare plant consenation. pp. 205-
217. Wiley New Y<n-k.

Rex, M. A. and R. J. Etter 1998. Bathsnietric patterns ol bod\
.size: implications for deep-sea biodi\ersit\'. Deep-Sea Re-
search II 45: 10.3-127.

Rex, M. A., R. J. Etter and P W, \nneskern. jr 1990. Densitx
estimates lor deep-sea giistnipotl ,issenibl,i<4es. Deep-Sea
Research 37: .5.55-569,

Sanders, II. L. 1977. K\c)lntion:in eeolog\ :md the deep-sea
benthos. l\i (ionlden, (.'.E. led. i The changing scenes in
natural scienics 1776-1976. pp. 22.3-243. .-Kcadenn ol
N:itnral Sciences Special I'nblication, Philadelphia.

Sautlers. II. L. and |. .A. .Vllen. 1973. Studies on the deep-sea
Protobrancliia (Bixakia); Prologue and the Pristiglomitlac.
Bulletin ol the Museum ol (^omparatixc /oologs 145:
237-262.

Sthoenei'. T. W. 1987. The geogniphical distribution ol rarit\.
Oecologia 74: 161-173.

Wan'ii, A. 1991. New and little known Mollusca Ironi Irel.md
.md Scandinavia. Sarsia 76: 5.3-124.

\\:in II \. 1993. New and little known Mollusca liciiii Icej.md
.md Scandina\i;i. P:irl 2. Sansui 78: 1.59-201.

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T H E€9N AU T I L U S

CONTENTS

\'(>hiiiif IK). Xmnhcr 2

June IH. 2002

IS'SX 002H-1344

Russell L. Million A cladistic anaKsis ol LUha\'ui i (lastnipoila: I'li-uiDtrridae)

iisiiiy; inoipliold^ical c'liaraftci's 39

Kic-ardo Ne«a i'itrtilliojilida krinfilonniL a new species l( iasti'opoda:

Hodollo ^<-'l^a ( 'oialliopliilitlae) Iniin tlie (Jaiian' Islamls li\iii'4 (in

Ajigel A. Luque Aulipallics in>ll(i\li>iii (( 'nidaria: Antlio/na: \iili]iatliaiia! 50

Martin A\er\ Sn%der I'ushiiis iloi jxh-di. a new spi'cies ((iastrnpoda:

I'aseidlaiiitl.ie) liiiin tin- Ri'd Sea. and lan^e e\lensi(in lor

two odier species 56

Helena Fortunato Tlie systematic position ol Sln>iiiliiiiu \('(il<>ii(ij)sisi liiidiic

Petncli. I9SS (Castropotla: f.'olniiiliellidae) 59

H. Lee Fairbanks The repiddncli\e aiiatonix, t;L\ononiic slatns, and raii^e ol

Oivolu'lix (iljiiiKi (I'drotl. 19011 iCiastropoda: I'ninionala:
Oreolielicidae! 62

Notes

Bruce A. Marshall Antliorsliip and ilate of |inlilication nf (hirca r7///rii\/,s

I'liilippi /■)( Kiister. 1S44 (Bi\aKia: Ostreidae) 66

Paul Callomon Tadaslii'^e ilal.e 1 1916-2001 ) 67

Book re\ iew 6J

THE NAUTILUS 116(2):39-49. 2002

Paee 39

A cladistic anal\ sis of Lithasiii (Gastropoda: Pleuroceridae) using
moiphological characters

Russell L. Minion'

IJcparlMiciit ol Bu)l()j;ii'al Siiciict's
Uiii\ersih- ol' Alahanui
Tuscaloosa. AI.354S7 USA

ABSTRACT

Till' fla.ssificatinii ol plciiroiinil snails ami otlirr Ircsliwatcr
iiiolhisks has liistoricalK liccii hascd on inoipholo<;ic'al char-
acters. Dcspiti- \cars ol ta\oiioiiiic work on plenroccrids. no
sinijlc work inchides all rccm^ni/ed ta\a Ironi a u;i\cn tjronp
anil onl\ a tew systematic treatments ol the lamiK' or indixidnal
genera exist. Modi-rn methods ol ph\ loijenetic s\stematics ha\c
shown that some moipholoi;ical traits do not support liistori-
calK accepted mollusk classihcations. II anaKses ol nioqiho-
logical characters do snppoil ciuix-nt taxononiic li\potheses,
then the classification ol tliesi- e;roups can lie tonsidered as
stable. It not. our approach retjarding diagnostic cli.u'acters tor
these groups must change. This paper uses 25 shell and radular
characters in a cladistic anaKsis ol the pleurocerid genus Ijth-
asiu, and compares flie findings to prexionsK snggestt'd clas-
sifications ol the group. Cladistic anai\ses do not support an\
current or historical classification ol Litliiisiii. However, these
moiphological characters are found to rongliK delineate IJth-
iisid and other extant pleurocerid genera, challenging pre\ious
works that suggest such characters ha\e limited ntilitx.

IXTKODUCTIOX

HistoricalK. the i-lassific-atinii ot tresliwater iiKilhisks has
relied lieaxiK on iiidiphdldgical features such as shell,
soft anatoiiu', and reproiluetive .structures (e.g.. Tnou.
LS73: Heard and Cuekert. 1970: Davis and Fuiler. 19S1;
Burcli and Tottenham. 19S()). Thks is partieiilarK cxident
for pleurocerid snaiks, win-re sliell features aec-ount tor
the inajorit\' of diagnostii- characters used in the taxon-
omy and clas.sifieation ot the group. One grouji ol ph-ii-
roeerids that lias !)eeii clas.sified on the liasis of shell
eharacti-rs is Lilliusid llaldeiiian. IS40. a genus ol large
ri\'er snails loiind tlirongliout tlic ( 'nnilierland, Ohio.
Mi.ssissippi, and Tennessee River drainages.

Species of Lilliasia pos.sess eonie to o\ate-conic shells
with tusiforin apertures, a posterior eallns on the parietal
wall, and fre(|neiitl\ some degree ot sculpture on the
hoiK whorl (linrcli. 19'S2). .Most species were tlescrihed
initialK as Mcltinia. and classified according to shell eliar-
acteristics such as sliape and sculpture, llaldeiiian

'Current address: Department of /oologx. Iii\ei1el)rate l)i\i-
sioii. Field Museum of Xatural Histon, 14(10 .S. Lake Sliore
Dri\e. Chicago. IL 60605 US.\: rmintonta fieldiimseinii.org.

(1S40I eri'cteil Lilliiisin .ind dcsignalcd /,. ociiii-ulald
llaldeiiian. 1.S40. .is llie t\p<' tor tlie genus and lati'r
erected AniHlniiid (1S41 ). I'resi'Uce of posterior and an-
terior calluses united Aiiiiitniud and IJlhasia. The pri-
inan characters separating the genera were that .\/(g(-
hviiui shells were spinous and had apertures with an
anterior sinus, while l.itliasiti shells lacked senljiture and
the aperture was not as distinctly channeled in front a.s
the t\pical Aniiilri'iiKir iTnon. 1873). Goodrich (1921)
supported Filshn and Rhoad's ( 1S96) reduction ol A/i-
'^itrcuui under LilluisUi. and IJlliiisia snhseciuentK has
stood as a single genus. Cloodricli ( 19401 recognized lour
si'parate groups wilhin lAtluisin. Iiased priinarlK on |)e-
culiarities ot shell siulpture. In tlu" 1970s, authors sug-
gested taxouoniie revisions ol the genus, placing all
nieiiiliers in /<) (Da\is. 1974) or Flfiiroccni (Stansl)en',
1971; Stein. 197S). Bnreli and Tottenham (19S0) rec-
ognized Lithasia scnsii stiirtc and Aiiiiitrciita as subgen-
era of Lithasia based on position ol sculpture on the
body whorl (Burcli, 19S2), and not according to the orig-
inal diagnosis, while Tnrgeon et al. (1998) followed
Hurcli in recognizing iJlhasia as one gemis.

.Vutliors lia\"e liistoricalK assembled pleurocerid geii-
ir.i based on grouping taxa with shared shell characters.
Such is the pri'Nalent ajijiroach found in the literature
published oxer a |)eriod ol 150 \ears, and no aualxses ot
these characters exists lor IJlhasia or an\ other pleu-
rocerid genus. This max be a result ol the lack ot nni-
lorni iiilormation louiid in llie literature. Diflereiit aii-
lliiiis rarcK proxided eomjiarable levels ol (|nalitati\(' or
(|iuuititatixe data in their original descriptions, and sel-
dom used terms and expressions that max or max not
liaxe the same ilescriptixe counotations (e.g.. tapering
xersus broadly conic shells), making it dilficiilt lor read-
ers to draxx comparisons betxxei'U xxorks. Manx descrip-
tions were based on one or a lex\' shells, jiixeniles, or
partial shells gixeii to the author, and radulae xxere not
included in these descriptions. FinalK', descri|)tions were
subjectixe based ou the experience ol the author the
amoiiut ol xariation tliex accepted, ami their imderstand-
lug ol the other taxa in the literature. (Jixeii the current
state ot Ireshxxati'r mollusk tayoiiomx in general and
pleurocerids specihcalK, and the fact that moiphological
characters are still usi'd to coufinn taxon identity, anal-

?:v'e 40

THE NAUTILUS. \ol. 116. NO. 2

Figures 1-8. Illiistration.s ol '.selected iharaeters anil charaetei slates iisid m the eladistic aiuiKsis niljflmsin. 1. Lithasia annigcra
sluming lusilonn aperture (2:2), and presence oT aiiteriiir (7:1 ' and posteiKii i(i:l ' calluses (in the cehnnelhi. 2. Litliasiii vcrniri'sii
.showing nodulose brxK' whorl (5:1 ) and poslenor lenij;llienni<; el ;ipeitiiic larmw: III: I ' .'?. Alisenee el cusp ne\t In lalenil tooth
(13:0). 4. I'lcseiiee ol cusp next to lateral tooth iarrow: l.'5:li. 5. Lamellar main lateral cusp i 19:11'. 6. Kectan^iilar iii:ini Literal
cusp (19:1 I. 7. Trapezoidal main lateral cusp (19:2). 8. Trian<;nlar main lateral cusp (19:3).

y.ses ol tliese cliaraet<'rs cuiploNiny; modern tccliniqiies
is pnulciil. Moileni methods orpiivlogeuetic ssstematies
ha\e shown that some moqiliological trails do not sup-
port historicalK aeeepled iiiolhisk ciassifiealioiis KJral,
2()()(); Lydeardet al.. 2000). 11 analv.ses of moipho!o<j;ieal
characters do sup])ort emreiif tuNonomie h\|K>llieses.
tluMi the classification oi these groups can he sta!)i!i/ed.
If not. our approacli to iisinsi diaiinostic cliaractcis lor
those groups unist change, j'lnlogenetic taxonomies of

this kind Iiunc iieen advocated in several studies (e.g..
de (,)uein)/ and Caulhier. 1090. 1092, 1994; Biyant,
1996; .Sereno, 1999; l,\de;n-d et ;il.. 20001.

l''urther complic;iting this |irol)leni is th;it pleiirocerids.
Ill a simihu' hishioii to wIkiI h.ippens to nuun other Iresh-
watei inxertehratc groups, ;ue experiencing (k'clines in
llieii iiumher ol species iuid iiidividuiils eiuised In river
impiiimilmenl. haliit;il di'gi";id,iti(iii, and poor laiid-nse
practices I Stein. i9T(i; Hogiiii et al, 1995; l,V(k';u-d et ;l]..

R. L. Minton, 2002

IVc 41

1997). For example, in tlie Moliile Basin, one ticiiiis (Cw/-
roto})ia) iuul approximately 31 other species an' prcsnniei!
extinct (Stein, 1976; Bogan et al., 1995; L\(leanl and Max-
den, 1995; Lydeard et al., 1997). Even with the loss ol'
di\'ersit\' in the tamiK', onK' five of 156 ri'cogni/.ed pk-n-
rocerid species (Turgeon et al., 199.S) are listed as either
end;mgered or threatened as of Ma\. 2001 (U.S Msh and
Wildlife Senice). It nioiphologN- ;ilone is to he used in
iilentifieation ol these imperiled taxa, then anaKses ol these
characteis gain e\en more importance. .\ lack of sncli anal-
yses can hinder efforts to recognize, manage, and consciAe
distinct taxa (\\aples, 1991; Mawlen and Wood. 1995)
within these affected groups.

Tile goal ol this stndx is to compile shell and radula
characters from Lilluishi. anaKze them using cladistic
techniques in ordei' to test historical and modern clas-
sifications of the genus antl its species composition, aiul
to determine possible relationships of the genus ;uul its
taxa to other pleurocerids based on those anaKses. Po-
tential changes to the taxonom\- of Lifh/isin based on
these analyses and the utilit\' of using these characters
in pleurocerid classification are discussed.

MATERIALS AND METHODS

Specimens for die study were either collected li\e oi' bor-
rowed from mu.seum collections (Appendix 1 ). Slu'll ch;u-
acters were taken direcd\ from specimens. Radnlae wt-re
extracted, cleaned, and prepared according to the method
described bv Holznagel (1998), viewed using a ilitaclii S-
2500 .scanning electron microscope, pliotographed, and an-
iilvzed. For Liflw.siii. at li-ast one represeiitatix e from each
recognized species and subspecies (Bnrcli and 'lollcnliani.
19S0) was includeil. Spetiuiens of selected taxa iipii'scnt-
ing five odier extant pleuidcerid genera (EUuiul /c /"g".
Leptnxi.s, Plcuroccm) were al.so inclntled (.'\ppendix 1 ).
Data consisted of a matrix of 25 cliaracters (.\ppendix 2,
Figures 1-S) coded as either binan or multi-state (Appen-
dix 3), and an;il\y,ed plnlogeneticalK under maximum par-
.sinionvwitli NONA 2.0 (C;olob()ff', 1998) n.sing the follow-
ing .settings: nnonk'red data, 100 replic;ites, with /;/gf; s'd-
iciila and Mrlaiuiidcs luhcrculald Miillen 1774. as out-
groups. //(g(; is basal to the rest ol the Xortli ,\nieriean
Plenroceridae (Holznagel and L\'deard, 2000) and M. Iiih-
crcnlata was chosen as a more distant outgrou|). jackknile
analysis (37% deletion, 1000 iterations of 10 i-eplicates
each) was performed in XAC (Fanis, unpublished: F;uris
et al., 1996) to test the stabilitx- of the data A strict con-
sensus tree mapped with characters was prodnc-ed with
Winclada 0.9.99m24 (Xixou. 1999). The anaKsis was run
twice, once using shell cliaracters alone and once with iill
ch;u-acters combineil. Because most pre\ious classifiealions
(e.g., Tnon, 1873; Cioodiicli, 1940) were based on sliell
characters onK; tlie\' were anaK'zed sepaniteK and com-
bined witli radnki data.

Once the classification li\potliesis was establislied. it
was compared to fixe diflereut classification selicnies
proposed b\- prexious authors (Appendix 4':

:\. bitliasia lepresenls a single genus. This assumption
lolioxxs the cuiR'ut iTurgeon et ;il.. 1998) xiexxofthe
genus.

B. Ijtiuisia represents a single genus comprised of txxo
subgenera, Lilliiisia scif-.ii strirlo anil Aw^itrruui.
sciisii Hiurh d\n\ Tottenham il980). This classifica-
tion is conimoiilx used as ;i st:uting point in pleu-
rocerid studies leg., i.xdcard et al,. 1997: Holznagel
and Lxxk'iu-d, 2000). Bnrch and Tottenham's (I980»
genera and subgenera differ in species composition
Ironi those ol TiAon ( 1873).

O. Lithasia repicsents a single genus comprised of four
species groups based on peculiarities of the nodulous
sculpture ((ioodrich. 1940). Lithasia luihriclili
()lench. 1965. had xet to be described and is includ-
ed in (irou|i 3 based on (dench i 1965' allxing it to
Lithasia ti'micasa i jiafiuesque, 1820'.

D. Taxa presentlx included in Lithasia bi'long to one of
tluei' genera: Lithasia, Ani^itrciiia. or Annihisa. scii-
sii Trxon (1873). In this case, the original descriji-
tions <il Lithasia ;ind .\ii<iitrciiia ;n'e used to group
taxa based on shell characters. Tnon failed to in-
clude Lifliasia ciiiia (Lea, 1868), and Lithasia hii-
hiichti luid xet to bi- described. Both species are in-
cluded in Ti-xons Aiiiiilrciiui based on their nodulous
shell sculplure. Lithasia i^ciiiciilata jiiirj,iiis i Lea.
1852) = Aiiciilosa piu^uis.

E. Taxa presentlx- included in Lithasia should be con-
sidei'ed speiies ol lo based on dexelopmeutal char-
acters, SCIISII Daxis (1974'. lo Lea, 18.31, has prece-
dence oxer Lithasia Haldeman, 1840, uml Aufiilrcma
Haldeman, 1841.

.\ sixth scheme. Steins (1978b' recommendation that all
Lithasia be considered Piciiroccra. is not treated here as
it IS nomiiicl.iliiial. not taxoiioinic. and has since been re-
solxed b\ the liilernational (>)iiiiiiission on Zoological No-
menclature's decision to make Flciiroccra anitiis tlie tx]^'
species ol the genus i Melxille. 1981: see discussion in Bo-
gan ;ind I'armalee. 1983i. (Characters supporting relation-
ships in the paisiinoux analxsis xxcre com|)ared to charac-
ters that groiijied spei ies in the other ckLssific'atioiis.

RESULTS

Maximuni p;irsiiiioiix analxsis of shell cliaracters alone
xielded 372 trees of 27 steps (Figure 9). Lithasia was
rendered non-inonophxktic b\ the placcMiient of Lith-
asia 'Uiiiiiilata jiiii'^iiis in the clacie ol Lcploxis species
;iiid the piac-emeiit ol Lithasia ohoiata (Sa\\ 1829) in a
polxtomx of I /(I ) some Lhiiiia + {pin<iiiis -^ l^cptoxis)
+ remaining Lithasia'. .\nalxsis ol all characters xielded
20 trees ol 107 steps that rendered Lithasia non-mono-
plixlclic i Figure 10'. Lithasia <iciiiciihita ])iiiiiiiis speci-
iiieiis were basal to a clade ol Lcptoxis s|)ecies supported
bx a te:n'drop-slKiped aperture, and Lithasia oboiata xxas
iiesled belwecn ckules cA Ph'iiroccra and LUiiiia species
near llie base ol the tree. The remaining Lithasia taxa
loiined a clade xxitli In supported bx three cliaracters:

Paw 42

THE XAUTILl'S. \oI, IKS. Xo, 2

Melanoides tuberculata
— — E. caelatura
— ^ E. hydei
^^ Juga silicula

81

1

P. canaliculatum
P. prasinatum
P. walkeri

Lithasia obovata
^^ lo fluvialis
E. olivula
E. alabamensis

65

Lithasia geniculata pinguis Duck
Lithasia geniculata pinguis Collins

Leptoxis ampla

- Leptoxis crassa anthonyi

- Leptoxis plicata
Leptoxis praerosa
Leptoxis taeniata
Leptoxis virgata

— — - Lithasia armigera Ohio
— i^^— Lithasia armigera Stones
^^^— Lithasia armigera Harpeth
-^^^ Lithasia armigera Wabash
^^^— Lithasia duttoniana Duck
^^ Lithasia duttoniana Harpeth

69

Lithasia geniculata fuliginosa Buffalo 1
Lithasia geniculata fuliginosa Buffalo 2
Lithasia geniculata fuliginosa Harpeth
Lithasia geniculata fuliginosa Duck
Lithasia geniculata fuliginosa Red
^^ Lithasia jayana
Lithasia salebrosa florentiana
Lithasia salebrosa subglobosa

Lithasia geniculata geniculata

- Lithasia hubrichti

- Lithasia salebrosa salebrosa

- Lithasia curta
^— Lithasia lima Bear Creek
^— Lithasia lima Elk

T^

Lithasia verrucosa Ohio
Lithasia verrucosa TN
Lithasia verrucosa Wabash
Lithasia verrucosa White

Figure 9. Chuiistic- iiiuilysis ol' Lithasia. Strict consensus of tiie 372 most parsimonious troi's (27 steps: CI = 0.74) ijencratecl
usiTii; siiell clianic-ters aloia-. Jackkiiile \alnes > 63% aliove nodes, number of imamhisiuous synapomoipiiios below nodes.

Mintnn. 2(102

Patie 43

^- Melanoides tuberculata
— Juga silicula

81 T~~ P. canaliculatum

2 I I P- prasinatum

* — P. walkeri
t— Lithasia obovata

^^ E. olivula

77

E. alabamensis
E. caelatura
E. hydei
— Lithasia geniculata pinguis Duck

Lithasia geniculata pinguis Collins
Leptoxis plicata

Leptoxis virgata

Leptoxis crassa anthonyi
Leptoxis praerosa
Leptoxis ampla
Leptoxis taeniata
— Lithasia armigera Ohio
Lithasia armigera Stones
Lithasia armigera Wabash
Lithasia hubrichti

Lithasia armigera Harpeth

Lithasia duttoniana Harpeth

- Lithasia curta

- Lithasia lima Elk

- Lithasia lima Bear Creek
Lithasia verrucosa Ohio

- Lithasia verrucosa TN

- Lithasia verrucosa Wabash

- Lithasia verrucosa White

— Lithasia geniculata fuliginosa Buffalo 1

Lithasia geniculata fuliginosa Harpeth

— Lithasia salebrosa florentiana

— Lithasia geniculata fuliginosa Duck

— Lithasia salebrosa subglobosa
— Lithasia duttoniana Duck

— Lithasia geniculata fuliginosa Buffalo 2

Lithasia salebrosa salebrosa

I — Lithasia geniculata fuliginosa Red
^H r^ Lithasia geniculata geniculata

I I Lithasia jayana

' lo fluvialis

Figure 10. (,'lailistic an.il\sis of Lilhasia Strict consciiMis of llu- 20 most p.irsiiiionioiis trees i lOT steps. CI = ()..374) <;eneratecl using
shell aiul radula cliaraeters conililiied. j.iekknite \alues =r (i:5'< alune iKides. miiiil)er "I uiiaiMhi'.ni()iis .sviiajToiiioiphies below node.s.

Page 44

THE NAUTILUS. \\>\. Ilfi. No. 2

tusifbnii aperture (c-liaract(>r 2: state 2), posterior i-allus
on aperture l6:l). and slij^ht anterior eanali/.ation ol the
aperture (S:l). Despite being well resolved, little snppoit
for an\' elade existed as evideneed 1)\ low jaekknili' val-
ues. Three other plein"oeei"id genera. Eliiiiiti. Ij'jitoxis.
and Vliiiroccru. \w\v all I'eeovfri'd as ni(iiiiipli\ letic.

DISCU.SSlOX

The c'ln'rent taxonomv ol pleuroeerids is based on shell
cliaracteristies. and most work on the faniilv has foeuseil
on these characters. An extensive literatnn' exists lor the
laniilv, with most works being either wholK tlescriptive
or taxonomic shullling taxa among groups. This studv
stantls as the first cladistic treatment ot all cnrrentiv rec-
ogni/eil l.itliiisiti species and their relationships to other
pleuroeerids based on shell and radula characters.

Neither anaKsis completely recovered the five classi-
fications being comparetl. In both [ilivlogenetic treat-
ments Lilha.sia taxa did not tnrm a single group, which
refutes the taxonomies of Bnrcli and Tottenham (f9S()),
Goodrich (1940). and Tnrgeon et al. (199S). Tiyou's
(ISTo) groupings ol the cnrrentiv recognized species of
Litluisia was also not supported, as sculptured and
smooth taxa did not group separatelv. Davis's contention
that LUhasia species should be considered as members
of /() was partiallv supported in the parsimonv aualvsis
of all characters, where lo was nested deep in a clatle
containing most Lithtisiii taxa.

The const'nsns ti'ees suggest tliat sliell characters alone
do not recover cnrrentiv or historicallv recognized groups.
However, shell and radula characters combined can be
u.sed to recover jilenrocerid genera, but do not resolve spe-
cies level identilv well. In ihc lota! chaiattei an;Jv,ses, EJi-
m'ui. lA'j)l(>xis. and licuioccra taxa all grouped in their re-
spective genus. Onlv hvo IJthasid taxa grouped awav Ironi
the others, suggesting that these two species mav be mis-
placed. lAtliaski >!fnicnlalii piii'^iiis was placed iu h-jiloxis
( = Anailosd) by Tnon (1S64I, and subse(|uentl\ [ilaced
in LilJias'm, vvliere it has icpreseuted the headwaters lorni
of the ^ciiinildld '^ctnaddld-fnliiiiiHKd-p'uiiiiiis complex.
Based on the moiphological cliaracters examined here. /,
i^eniculutd piiiiiiiLs should be classified as a species (i\' Ia'j)-
toxis. lAtlui.sid ohovdid is the onlv species in the genus thai
(X'cnrs in the Creen River drainage of Kenlnekv and has
included njanv nominal forms that are (jneslionablv l.illi-
asia. L. ohoiaUi shells lack the calluses on liic apci-tmc.
fiisifonri a])ertures. and have radulae most similar to Elimiii
species. Though the plivlogenetie aualvsis suggests alloca-
tion o( /.. iilnnald to a separate genus, I belii've the ad-
dition ol more EliinUt and Fleiirorcrd species or more mor-
plioiogicid chaiaclers In ihe aualvsis would icsolve its ge-
neric designation. The remaining Litluisia species would
lie cousiilered h>. as hi is the oldest name for that clade.
As such, lo would be dia'gnosed bv having a fusiform ap-
erture (2:2), posterior i alius on columella (6:1), and for-
mation of canal of the aperture (S:l-2). This change would
reflect the opinions ol fJavis ' 1974) regarding the two gen-
era. Within Litha-sia, liowevei', individual species weic not

recovered, and continued aualvsis is re(]uired to elucidate
diagnostic characters al the species level. None ol the five
current and historical tlassifications ol Eitluisid evaluated
iu this studv are completi'lv cousisti'iit with the aualvsis.
OuK the diagnosis ol Bnrc-li (19S2) is partiallv supported.
A posterior cdlus on the columella (6:1 ) and the fonnation
of anterior canal of die aperture (S:l-2) unite all Litlwsia
(minus ff'nicuUitd pin^iiis and ahot nla plus la i in the corn-
binetl character aualvsis.

This studv offers evidence refuting pri'vious notions
that shell and radula characters have limited utilitv in
recognizing pleinocerid groups and supports the use of
these characters in defining pleurocerid genera, in the
most inclusive treatment of pleuroeerids to date, Tnon
(IS73) offered an extensive discussion on tlie use and
validitv of shell characters iu separating t\\v various gen-
eia and species in the group. Tnon recognized that shell
characters can van' greatlv and looked towards the use
ol other anatomical characters to separate "natural gen-
era" and discover corroborativ e sliell characters for these
groupings, (a)oilrich (1940: 1) noted that shell charac-
ters "once. . .considered inunutable have proved to be
secondan' and more or less evanescent'. Tnon (1873:
liii-lv ) figured some ol Troschel's illustrations ol radulae
and conunentetl on Stimpson's obsenatious that sliell
and ladnla characters seem to unite pleuroeerids but do
little lo sejiarate constituent genera and species. It is
triH- that gastropod radulae (Padilla. 199Si and tresh-
vvatei' mollusk shell characters can be plastic, olten ex-
hibiting clinal variations (e.g., .Adams, 1900, 1915; Ort-
maun, 1920), ami therefore potentiallv contrilinting ho-
moplasv to phvlogenetic aualvses. Adtting solt anatomv
characters to a studv such as this might tlieoreticallv im-
piovc the resolution ol the analvses, lint manv eharactei's
lre(|ueutlv used to delineati' taxa van little among pleu-
locerids (Dazo, 1965) and in phvlogi'iietic reeonstna-
tious shell chaiacters often are less homoplastic than an-
aloniic;il cluirai'ters i Schander and Sundberg, 2001).

The results given here are consistent with studies em-
plovnig molecukn' methods (Lvtleard et al.. 1997: Lv-
di';n-d el :il., I99S; llolznagel' and Uvdeard. 2000) to
idenlilv [)Ieuiocerid genera. In tlii'se works. Eliiiiid and
I'lcuroccni represent natural groups, and the uioqiho-
iogic:il characters siippoi't both genera. However. Lv-
(leard (I al. (1997) and llolznagel and Lydeard (200())
showed lli:il Lilhdsid :md Eiploxis are uon-mono]iliv let-
ic, ihoiigh the eombmed iiiorjiliological analvsis given
here supports the reeoguitioii ol Ecj)l(>xis as a natural
giou|). \ (horough moleenlai' sindv ol IJllidsid. including
all ol its nomin:il species and lornis. \\\\\ pro\ ide \ ;ilu,il)ie
inloiiualiou on the species composition ol Lillidsia. anil
subse(|n('iil ehaiacler :iiialvsis will lielp driiui' ihe diag-
nostic le:ilin'es ol tlii' genus.

.ACKNow i,i;i)(:\ii:nts

I would like to Ihank ,S, Ahlsledl. |. ( ;,uner, and I), \eelv
lor s|)((iiiirns. \I. (ilaubreeht. I'. I lariis. ('.. lAtk'ard, and
v.. Strong provided heljilul comments on the manuscript.

R. L. Miiitnii. 2002

Paiif 45

W. Hol/iuiiifl ami |. NiinlcN' lussisted witli radula prepara-
tion and illustration, and \'. Albert pnnided access and in-
stniction to NONA, W'inclada. and -V\(". A Resi'arcli ( aant
from Concholc joists of Anienca Inndetl this project.

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R. L. Miiitoii. 2002

raui' 47

Appendix 1. Ssstciiiatic list dl la\a used iii iIk i I.kIisIic analxsis nl tin- minis LUliasia i ii = I lor rarli t.iMiii '. ( lissification follows

Turgeoii v\. al. (UWS): la\a aic iiaiiicd as m Hiinli (liJSOi, C pldc ImalitN iiitoniiatioii is axailalilc Ikhii iIic aiillior, I'MNII —

Field Miisi-iiiii nl Xatmal llislon I\1IS -llliiiiiis Natural ilistorx Siirvi'V: NCSM — Nmlli Carnliiia Stair Miisiiiin iil Xaliiral
Sciences; UAti — Lluivcrsitx ol Alaliaiiia Castropinl ( .'ollectioii; I'MM/ — l)iii\cisil\ of \licliii;aii Miisciiiii nl /iiolii'j;\

Taviii

Localit\

( A)llrctiiiii
niinilxM"

FaiiiiK Plriirocrridae
(aMiiis Eliinid

F. (llilhdliuil.sis (Lea, ISdll))

E. ciicliitnrd iinliiliini iKnAe. ISIidi

K hi/ilci (Cloiirad. IS:54lii

K. (tliiiilii (Cimrad, IS34a)

Cleniis Id

I,> fliiiialis (Sa\. 1S25)

Ca-iiiis jiii^ii
I silinilii iC.nild. 1S47)

Cieniis Lfj)loxi\

L. init])l(i (AntlmiiN. 1.S55)

L. cnissii (iiitlioin/i ( l^edfield, IS54)

/., /)//(■(//(/ (Conrad. l.S34li)

/. pnicrosii (Sa\, IS2I i

/,. Iiiiiiiiitd ((Jinratl. 1S341))

/„ tinj^atd (Lea. lS41al

(ieniis Lilluisiii
L. iirini''! Ill I Sa\. 1S21 )

/.. aiiia (Leu. 1868)

L. iliittimiiniii (Lea. 1841a)

I, •j^i'iiiniliilii ■j^iniiiiliilii {\\Mv\iidt\. 1S4I)
E >^ciiiciiliitii fiili^iiKKii (Lea. 18421

/.,

IciiifuJiitii jiiiipii'i

(Lea, 1852)

E- hnhiirhti CIcikIi, 1^65
E. jaijdHil (Lea, 184 llil
E liiiui I ( 'niirail, 18(34a)

E oh, It, ilii iSa\. 182V)t
E MilcliiiKii sidchi'iKii (Conrad. 1834a)
L. salrhnisii fh>nuli/iiui (Lea. 1841a)
L. salfhntsa siih^Lihosa (Lea. LSfila)
L. verrucosa (Hafiuesque, 1820)

Genus Plrurocrii
P rinudiiidaiuiii filuin (Lea. 1845)
/' pnisiiiiiliiiii i(!oiir.id, lS34a)
/' uidki'ii C.H.drieli. UJ28

FauiiK Tlilarldae
( a'lllls M, 1,11101, lis

SI liihintiliilii i.Mullcr. 17741

( !uosa l\i\er, ('oosa ( 'o.. M ,
(111 Kii linen 1 ( reck, ( 'all HI 11 II ( (i,, .\L
Black Warrior Hiver. Jericisi.ii Co,, AL
Alabama Ri\er. .Monroe (!o,. .\L

llolstoii Rl\ir Siilli\aii Co T\

Oak Creek. Hciitmi ( o OK

Liltic Cahalu Uimi, Kilili ( ii \L
Sci|natcliie Ki\er. Marion ( !o, AL
Black Warrior Hi\cr, |ellcrsoii ( o , \L
llaipetli Hi\ri, OaMclsoii ( ji '!'\
( lioccnioceo (deck, I'alladi'^a ( (o., .\L
(diiicli l!i\ei, I laiirock ( 'o,. IN

Wal.asli l;i\ci Wliilr Co 11,

Oliio Hixcr. Massac Co,, IL

h'.asi I'oik SloiHs i;i\(i Uiitlicrlord Co TX

llarpitli ItiMi ( :lii-,itliaiii ( n lA

lennessee l\i\ci. Lauderdale (ai.. .\1,

Duck Hi\er. Maun Co.. T\

llarpctli Ki\er, Davidson ('o,, T\

Caiicx Fork. Dekalli Co,. TN

Biillalo Hi\er. Iluiiiplire\s Co., 4"N (1)

Red River. Rolicrtsou Co,. I'N

Duck River. Maun (;<)., TN

RulTaki River. Pern Co,. TN (2)

llaipeth River. Davidson ('o., T.N'

Oillins River, Warren Co., TN

Duck River. Coffee Co,. 1\

Ht'i Black River. Hinds Co., MS

Duck River, lliiinplircvs C.n.. TN

I'dk Hiver. Limestone ( 'o M.

Bear Creek. Collierl Co,, ,\l.

Careen River, Mel .r.in ( n K\

Tennessee River. Lauderdale Co.. .\L

Tennessee River. Hardin Co.. TN

Tennessee River, ILirdiii Co,, TN

Waliasli River. Wliile ( " il.

Olio River, Massac Co,, II.

Tennessee River, Hardin ('o,, IN

Tennessee River, L.iiiderd.ile ( (o , \l.

Duck Ruei M.iiiiA Co IN
Vellovvleal Creek. Slielliv Co,. AL
Slioal Creek, Lauderdale Co,. AL

Draina'^e ditcli. Cainesville. ,\l,icliua Co,, |-'L

N(:SM-R-4658
NCSM-l'-465y
NCS\LI'-46(i3
NCS.M-l'-4(i(i4

NCSM

IfifiT

NCSM-l'-4(i7()

NCSM 1' 4671
NCSM-l'-4(i72
NCSM-l'-4fi74
NCS\LR-4(i75
NCSNM'-4fi7fi
NCSNM'-4(S77

INI IS 2.3628
INHS 236.32
UA(; 397
UAC 572
UMMZ 2122()()

vm: 402

UAC 405
U. MM 7.51363
VAC. 406
UAC; 398
UAC 403
UAC 395
UMMZ 53233
UAC 407
UAC; 392
FMNH 137751
l^AC 573
UAC 571
UAC 570
FMNH 46219
UAC; 565
UAC; 425
UA(; 416
INHS 23629
INHS 23631
UA(; 427
LAC 568

NCSM-I'-46S6
NCSM-I'-4689
NCSM-l'-4692

N(:SM-P-46S2

ra<re 4S

THE NAUTILI'S. \(.l. 116. N<

Appendix 2. M„n,l,..lo,ic.l elunacto a,ul cluuactc,- states used in tlu- clad.stic anaKs.s of the genus LUl...

1 Shell shape, l"! t;li>bose. 1 1 » conic, (2) ovatel) c v

0 Aperture shape. \()) teardrop, i 1 1 oNate, Cl) lusih>nn.

^' Sculpture on posterior hodv whorl. lO) none, (1) carinate, (2) tu hercles.

t f n,e!lialK on hodv whorl. ,0) none. U) sha.p angle on InxK f'^^^^^^^^;^^

5 Sculptu'v on enfre l,o,h uhorl. (01 n,>ue, (1) even lateral rows «i snudi nodules, (2) costate. ^

6 Posterior callus ..n cohunc-lla. (()> absent. (1) present.
7. Anterior callus on cohnnella. lO) absent. (1) present

S Lencnh of anterior eanal of aperture. (0) none, (1) slight, (2) elongate.
9 TwisHna of aperture anteriorly. (0) absent. (1) present.
10 Lencnheuingof aperture posteriorly along bodv whorl. (0) absent. (1) present.

;:■ Sa::f 'i;:;i:^ S: ti tS^l ^-^^^^o..-..^ ^H^H l'->.tb, ^.) ...ore than one-han shell length,
1.3 Cusp next to lateral tooth exteriorly. (0) absent, (1) present.
14 Shane of upper rachidian margin. (0) convex, (1) straight.

5 nhAvic th raHo of rachidian. (0) tooth as long as uide (1) tooth ongerhan v. le

,6 Lenath/width ratio of central rachidian denticle. (0) eiigth and uukh equal, (1) longu than u.ck.

1;: ?±:;: ^f^;^! .^tl? oifrlnttl^lC^^ - u.edia, quarten (1) edge restncted to medial h.. >2, edge

s- EHSU'io '- - 1:^^^^^^^^^ - =i->- - '• -■ '--- "'■ ---

01 Sgt£:r;:;io:n;:::nS C5.-fe.i. ^^.." ; '-^ --!■ <; > ^-f ; ---■ ^"^ "'"■ ' " ^^^'" "■"■'' '"" '"*'

22: Shaix" of leading edge of mmn lateral cusp. (0) pointed. ( 1 ) rounded, (2) stiaight,

0.3 Shapeof marginal teedi. (0) pointed, (1) round.

04 \uniber of inner niar0nal teeth. (0) 1^, (1 ) 5-.S, (2) more than S.

25'. Number of outer marginal teeth. (0) 1^, 1 1 ) 5-S, (2) more than S.

Appendix 3. CMadistic anal
ta\(iii names where needeil

\sis

IJtlu.sia Data matrix of t^ixa and character states. Uixm' names loll.m approi

iriate

~ ~ -202010100011011002101112

I.ilhasia armfj,,,;, OInu 2201010100011011012102112

Lilhasia armi'^rra Stones 2202 010100011011001012101

IMluisw anm<icra Haipe li 2202010100011011001102112

iJllmsUi (inniiH-ra W'abas

2200111101011011010112112

Lilhasui nuiri 2201010100011010020212112

Lilhasu, dnllouuma Dn.'k 2200010100011011013020102

/.,•/ m.sm </„»..MJ.,m 1 aipetl, 2200011100011011020211112

IMhasm ^■n,rnlala juhiimos,, Kullalo I 2200011100011011021212112

2200011100011010021212112
2200011100010011021212112
2200011100011010120211112
2220011100011010110111112

alo

Lilhasia iif'nindata fnliainosa Bullal
Lithasin oniiniliilti fiili<I,iiu>sa Duck
IMluisia \Sfuinilat(i ftdiilinosd llarpetl
Ijlhasiii iLcuinilala fidifliniKd Hi-d

lilhasia i.-nindala grmn,/«J. 0000000000001011011102112

.„W, ^.nndala ,nu^u. Duck 0000000000011010111202112

/,i//w.vm -cmr„/r;^/ ,.mt;,».s Colhus ooonnn nnnm 1 Oil 0111121 1 2

IJlliasia huhiichli

2220011000011011011112112
22 02010100011110011111111

lUlmsu, jaynnn 2200110101011011011112112

Ullwsia hnw iear (a.'ck 2200110101011011011112112

Ulhasm h,„a I- Ik 2100000000011011001021112

.,( m.s,« <.to/ 0220011100011010020211112

I,lha.sw saleh.u, salrhnsn 2200011100011011021212112

.,...,« salehn '' tlon.'ann 2200011100011010121212112

./ ,«.v,« .v«/.Ws« .,/,,/,./..« 2200111101011011011112112

'"""'" '^'■'"'•'"•'"■" -V^.'" 2200111101011011011212112

Ulhasu, .crn.rosa IN 2200111101011010011212112

lilhasu. vrrnuosa abash 2200111101011011001102112

./.;.v,.a.rn,o..s.«Uh>.e 2200000200011010110100111

/() jiuiiali'i .

R. L. Mintoii, 2002

Pam- 49

Appondiv 3. ( cmti

.1

Li'ptoxix 1 1 mill 1 1

Lcptoxis cyiisMi iiiillitniiii

Leptoxis pliciitii

Lcptoxis jinicntsii

Leptoxis liiiiiidlii

Lcptoxis lir^dtd

Etimiti cilabtnucusis

Elirnia caclatnra

Eliiuia hijdci

F.Iiiiiiti olimld

juffi silicuhi

Pleurocera cdiitiliriiltiliiin liliiiu

Plcurorrni jtiyisiiiiiliiin

rlrui'iHCi'd It iilki'i'i

Mrliiiioidrs liiliiiridiilil

00000
01000
00100
00000
00000
00000
21000
21002

? 1 n 0 -•

00000
00000
00000
00000
00000
00000
00000
00000
noooo
tOOO

02010102
02111012
02111101
02111012
02010102
02111002
01101100
11101100
11101100
01101010

1 r\ 1 n 1 1 n f)

LU02000UU10111110

0212101
0212111
1002101
0212102
0212101
1212101
1001111
1001111
1002111
1001101
3020011
1011122
' " : 0122
.1122

joionj:

Appendix 4. (;i'iiiis l.ilhiisiii Classification scliciiirs used in comparison to claciistic li\potlics(>s. Taxa marked until a (*) were not
treated li\ tlie i>ri<j;mal .iiitlior lint are included in the primps liased mi tlnir works isee te\( lor explanation'.

Tnr<jeon et al.

Bnrel

loltl

I9S()

Coodneh, 1940

Tnoii. I ST.)

I)a\is, 1U74

rein IS Litluisid
L driiii^ciyi
L. riiiid
L. (Iiittonidiui
L iicniciilatd
L. hidirichti
L. jnijana
L. lima
L. oboidtd
L. sdlchnisd
L. icrmcosd

(icnns Lididsid
.Snlit;eniis Lilluisid

L. flcniciddtd •j,ciiiciddtd

L i^ciiicidiita fidiiiiiiosd

L iicniculiild piii'j^iiis

L. ohoidtd

L. snicbvosd sdlcbrosd

L s(drbnisd florcntidiid

L \idrhni\d sidnjohosd
Snligenns Aii<j,i1n-iud

L dnni'^crd

L rnrtd

L diillniudiid

L huhnrlili

L jdi/dUd

L liiiid

L nrniciisd

',r\\\is l.ilhdsid

( ironp 1

/, iiriiii'J^ri'd
I. dllllnilidlld
I. Idljaild
I. liiiid

( ironp 2
/, 'J^i iiiciildid <^cniciddtd
I, •J,riiiculdld fidi0nosd
L. iifiiicnldld pin<itiis
L. sdlcbrosd s(dcbrosd
L. Sdlcbrosd jiorcntidiui
L. Sdlcbrosd sul)<slobosd

Cronp •■!
/, nirld
L liiibriiliU*
I. irirutosd

(iroiip I
/, obonilii

Genus Ancidosd

L '^ciiiciildld piiiiliiis
( iemis Aicj^ilrciiid
dnni\2.crd
rnrtd ■

.. llllllolliillld

.. '^ciiiciildld '^cuiculdtd

, hiilirirlili'-

. \d[\dna

. liiiid

,, sdlcbrosd sulchrosii

,. rcrnicosd

nns Lithdsiti

. <^cniciddld jidii^iiiosd

.. oboidid

,. sdli'brosd fliirciilidiid

, sdli'brosd sidr'loliosd

(a'nns lo

fliiridlis

iinniiH'rd

curld

duttonidud

'jfuiculatd ^cniculdtd

<S.ciiicul(ild fulifiinosd

•iciiiciddtd piii<^iiis

liiibriihli

jdt/diid

liiiiii

obordia

sdlcbrosd

Sdlcbrosd florcntidiid

Sdlcbrosd siib^lobosa

icrnicosa

THE XAITILUS 116(2):50-55. 2002

Pasrr 50

CoraIIi()j)Juhi kaofifonoii, a new species (Gastropoda:
Coralliophilidae) from the Canaiy Islands Ihing on AiitipatJies
wollastoni (Cnidaria: Anthozoa: Antipatharia)

Ricardo N'ega

Aliapanoii, 65
29(1 IS Malaga
Sl'\l\

Rodolfo Vega

A\da. dt' los Meiiceses. 38
3820.5 La Laguiia (Tenerife)
SPAIN

Angel A. Luqiie'

Lahoratorio de Bioiogfa Marina.

DfpartanK-iiti) dc Bioloiii'a
Facultad de ( Jcncias. Universidad

Autoiioiiia
28049 Madrid
SPAIN
angel. lii(jue(S I lain. cs

AHSJKU.T

.A iK'w spt'cius ol Cdralliophild Iroiii tlie (,'anan' Islands iiviiisi
on the antipatiiariaii Antijidthcs udlln.stoni is desciilied. The
new speeics diliers Iroin its closest Eastern ,\tlantic relatiNc
Coralliopliila lirciis iBlaiiuille. 1832) h\ its smaller si/e and
spiral cords ol similar widtli. and Irom the Western Atlantic
Covdlliopliila ciirihdcii \l>liiitt. IV).58. li\ liavint; a thinner shell
and narrower a[)ertun' and In the scnlptiirc and smaller size
ol the protoconch. .Additional remarks on the t;L\onom\ and
feetlin<r of some Easleni .Atlantic and Mediterranean Corallio-
jihilidae are incliiileil.

IXTKODrcTlOX

Tlic laiiiiK ( '(irallii)pliiliilac ((iiiipriscs tropical tci tciii-
pcrate, siiallow- to deep-water niariiii' species tiiat li\t'
and feed on cnidarians. The coiicliolo^ical characters
may \ai-\- <freatly wilhin the rainiK. and there ari" dilTer-
ent grades of interspecific and iiitraspeeifie \ariatioii
Shell \arial)ilit\-, togetlier willi llii' raril\ ol most of the
species due to their restricted haliitat (specialK those
li\ing in deep water), the absence of radiila, tlie absence
ol protoconch. nsnaliv eroded in adults and c\eii in
yoiiiig specimens, and the poor kiio\\leilij;e ol anafoniv
and hiol()g\' have led to a conliised taxononix of the fam-
ily tliat inaki's it tlie generic allocation of specii's diffi-
cnlt. The (airalliophilidae lias been IradilionalK consid-
ered a .separali family closeK ri-laled lo Muricidac. but
Ponder and Waivii i 19SS), witlioiil linlhei' discussion.
proposed Corallioplnlinac as a sublamiK of Mmieidae.
This taxonomic si ihis has been adopted b\ some siib-
se(|iicnl authors i Poiidei l!J9S: liiedel. 2()()()). ilowe\er.
Kantor ( 1995, 199(Vi Idniul enough differences in foregnl
anatomy to jiistif\ tin ;: separation into two different
families. Since the taxonomic status of the Coralliophil-
idae is cnrrentK nuclei- diM nssiou, uc prefer to use the

traditioualK acceptetl ranking ol tlii' taxnn at the l.iniiK
le\'el.

No critical re\isions ot the taniilx' haw been published
except for tliat of Vlassin (1982) for the genera -A/(/g(7(/.s
and Li'ptocoiichus. but two catalogues (D'.Attilio, 197S;
Kosuge and Suzuki, 1985) consider about 200 Recent

species. At least

<iei

lera ha\e been d

lescnhed

but the

Author for torrespoTKleiici

actual number is probabK' smaller (DWtfiliu and Beitseh.
1979: Kosuge and Su/nki. 1985'; a largi' number ol s[ie-
cies (about 80) are inclucled in tlie genus CordU'wplnld
II. and .\, ,\dams. 18.53. but this genus still needs to be
better dehiietl. .Most coralliophilids Iced on scleractinian
corals, but there are also spi'cies that ti'i'il on (ioigonacea.
C.'orallimoipharia. Zoaiithidea (Miller, 1981; Robertson,
1970. 1981; Wicksti'ii and Wright, 1993). Actiniaria (Spa-
d;i, 1979; ()li\erio, I989;ii. and .-Ucvonacea (Lorenz.
I99(il OiiK h\() species ol Rhizochiius iRliizocliiliis an-
tijuilliinii Steenstnip. 1850. and Rhhi>chihis sp.' and ('or-
ullii'hid iiiiiiiii'j^ii (II. and .\. .\il;ims. 1S6.3) are known to
Iced nil \iitip;illiaria in the Inilo-PaciHc ( Iva\'. 1979; Poor-
iiuin. 1981: Kosuge ;ind Su/uki. 1985; I^Wttilio and Ko-
suge. 19881 The t\pe ol U'l'diiig and the degree ol as-
soci;ition willi :intlio/o;ins \:u'\ Irom boring, eiidobiotic,
almost p;ii':isilic. liigliK hosl-selecti\e spi'cies (.\/r/g(7(/.v.
LcptdCiiiH-liiis. Rcl'HjituuTdid) (Nhissin. 1982. 1983. 1987.
1988. 1990). to less selective species with a relati\'el\- high
mobiiih i(',i)v////<7)/i/7r/> iMiller. 1981).

Kelerenees lo I'liistern .Atlantic coralliophilids are scat-
teied in ])ublic;itioiis Irom dillerenl ;ire;is: Meiliterni-
iiiMii ;md I'.iirope.in ;\ll.iiilie iS;ibelli :in(l Sp:iila. U)8():
()li\eiio. I989;i. li; Poppe and (ioto. 1991', cnnliiieiihil
WesI Alric;i iKmidseii, 1956; Bernard. 1981; laiaxeni,
1975; (;ol:is. Piiito-.AIonso and Brandao. 19S5: Kosugi-
.111(1 Fenumdes. 198S; Smriglio :ind Maiiotliui. 20001.
deep w;ilers ol the iioiihtMslerii All.inlie i Honchet ;iiid
Waren. 1985). (.ape \erde Iskmds (Cosel, 1982), Sao
Tome Island (Kosuge :ind Kernandes, 1989: Rohin and
PeriKiiides. 1990), Saiiil llcleii.i Isi.ind (Smith, 1890),
;iiid iVislaii (!:i('niili;i IsLuul Walsoii. 1886). Four spi'-

R. Wiia ct al., 20(12

Paei' 5!

cies were recorded from the Canan' Islands In- Nord-
sieck and Garci'a-Talaxera (1979): Coralliojiliila luci/fii-
cloijfii (Caleara, 1.S451. C. hiviis (Blainxillc. \H32}.' Bti-
bclomiiivx cahnifcnis (G. B. Sowerbx; 1834) las Coral-
liophihi bahclis (Requien, 1848)), and ConiUiophila
riclwrdi (P. Fiselier, 1882) (as C. lacfuca (Dall, 1889)).
In addition to C. richardi. hvo additional deep-sea spe-
cies are known from tlie Canan- Islands: C. sqiuimosa
(Bivona, 1838) and C Ixisilcus (Dantzenbersj and Fi-
scher, 1896) (Boucliet and W'aren, 1985). FinalK", Cor-
(lUiophila foutanaii^ioi/i Smriglio and Mariottini. 2()()().
has been recenth' described from Tenerife.

During SCUBA-cli\ing samphng around the island of
Tenerife (Canan- Islands), a coralliopliilid living on the
black coral Aiitipatlws wolla.stoni (Cra\ ) has been found.
This third species ol the famil\- knou-n to li\e on anti-
patharians is described here as a new- species, since it
shows clear differences from the pre\iousl\- known cor-
alliophilids iroTU the .\tlantic Ocean.

Institutional abbresiations used in this work are: BAU,
Dipartimento di Biologia .'\nimale e deU'LJomo, "La
Sapienza" Uni\-ersit\-, Rome, ltal\-; MNCN, Mnseo Na-
cional de Ciencias Naturales, Madrid, Spain: MCNT,
Museo de Ciencias Naturales de Tenerife, Tenerife,
Spain; DBU.\, Departamento de Biologia, Unixersidad
Autonoma, Madrid. Spain.

SYSTEM.\TICS

SuperlamiK Muricoidea Rafinesi|ue. 191.5
FamiK- Coralliophilidae Chenu, 1859
Genus C(>nilliiij)liilii II. and A. .\dams. 1853

Type species: Fiisiis niiitoUlcns Lamarck, iSKi, In
subsequent designation (Iredale, 1912). Recent. Indo-
Pacific.

Coralliiipliilii kiuililiumn new species
(Figiu-es 1-7)

Description: Shell i Figures 1-4' up to 24. H nun of
length. 15.5 mm width, solid but ratlier thin, ovoid-iu-
siifirm: spiri' high, conical, with up to 7 con\e\ whorls,
shoulder slightK angulate: suture well defined with un-
dulate narrow groove. Slu'Il color uuilorniK vellowish-
or pale-brown, milk-Ti -white when cleaned. Protocouch
(Figures 5-7) of about 3y2-4 w-horls, 1 uuu length. 940
fj-Hi width. :ukI diametei- ol first whoil 280 |jLni. Pioto-
conch indicative ol [ilanktotrophic lai-val development.
Protocouch nucleus (Figui-e 7i smooth except lor small,
scattered pustules, subseciuent whorls with tw-o strong
spiral keels, crossed b\- axial ribs tormiug nodules at in-
tersections and scattered, small pustules (Figure fS).
Thick xarLx inilicates limit between protocouch :m(l te-
leoconch. Protocouch color pink to reddish- or pale-
brown. Protocouch lacking or usuall\- eroded iu adult
specimens, but when present is frequeutlv somewhat lilt-
ed in relation to shell axis. Teleoconch with 5-7 convex
whorls, somewhat angulate at shoulder on eai'lier w liorls.
Last whorl larsie, convex, more or less angulate at shoul-

der, then uarrowiug toward base, representing almost 3/
4 ol total shell length. .Aperture large, oval, lirate and
white within: but one specimen with inner side of ap-
erture pinkish. Outer li]i thin, with fiuelv- serrated outer
niargin (margin rippled internallv): inner lip slightlv an-
gulate in middle. Siphonal canal broad, motlerateK- elon-
gate, open, slightlv recuned. Umbilical aperture absent.
,A\ial sculpture of 9-11 ion specimens lougei' than 10
mm) or 8-9 (on smaller specimens) broad, elevated axial
ribs on last wliorl, and usuallv- 10 axial ribs on penulti-
mate whorl. .Axial ribs less marketl towai'd end of bodv
whorl in longest specimens, but clearlv- mai-ked in small-
er (younger) specimens. Spiral sculpture on last whorl
of up to 18-19 cords of rounded cross-section, all of
similar width, and bearing prominent denselv packed,
fine and long, scale-like lamellae. Spiral cords 8-9 on
the penultimate whorl. Shell usuallv covered bv encrust-
ing organisms [Miiiuicind. bn-ozoans. seqiulid polv-
chaetes, two different small species of bivalves, sponges
and calcareous algae were obsened on a single speci-
men). Living animal white oi- whitish, inc-luding tentacles
and siphon, with deuselv-packed, vellovv speckles, which
form a ring on distal part of the siphon. Operculum
brown, mahogauv-, or reddish-brown.

Type Material: llolotvpe (Figures 1-2 . .\L\CN"

15.05/32583. 20.0 mm length X 13.5 mm width: five
paratxpes, .MNCN 15.05/32583: txvo paratxpes, .VICNT
TFMCBMMO/000277 and TF.\1CBMM6/()00278: txvo
paratxpes, DBUA. 15.05/18. All from tvpe localitx-, at-
tached to black coral Aiilijiallics wcillasloiii (Clrav). Xov.-
Dec. 1994,

Type Localitx: Puula de Tt-no, Tenerife. C^anan Is-
lands, 28°20' N, 17°55' W. depth 22-24 m.

Other Material Exaniined: Twelve specimens Irom
the txpe loi'alitv. Ricardo and Hodolfo \ega Collection,
txpe localitx. R'. and R. Wga coll. Nov.-bec. 1994: 18
specimens, B.VU, Punta de Teno: 12 specimens, BAU,
Diente de .\jo: 21 specimeus. BAU. N'eril de Masca, all
Tenerife, Canan Islands. \l. Oliverio coll.: one speci-
men, 20.9 mm length x 13.fi nun width, R. and R. X'ega
Collection, La Bombilla. Palma Island. Canan Islands,
collected aliv-e on Aniipatlic.s udlla^lciii. 40 ni depth. 1).
R. Gallowav coll.. 24 Aug. 1996.

Distribution: known onlv Irom the tvpe localitv, Pun-
ta de Teuo. two olhei- uearliv points ou the Tenerife
Island (Diente de .\jo and N'eril de Masca', and from La
Palma Island (La Bombilla. David Roig (ialloway. leg.).
The planktotro|)hic tvpe ol development indicated bv-
tlie protocouch of CondliopJiila kaojUonun suggests a
wider distribution than that currentiv known, but the
sjiecies has not vet been lound in other Maearoncsian
Islands, the West .\fricau coast or the Mediterranean.

Habitat: .\I1 collected and obsened specimens vvero"
found attached near the base of colonies of the black
coral Aulipiilhi\ uolldsloiii (Crav) or near the major
branch nodi's in the case oi ven large colonies, between

Patle 52

THE XAUTH.US. \nl. 11(1 No. 2

Fifjures 1-7. i'oidlluiiihdn kdiifittiniiit new sprcics. I, 2. llolotxpe, MNCN, 15.05/32583. 24. (i X 15.5 nun. I'linta (le Tciio.
TfiKTife, Caiian Islaiul.s. 3, 4. l\iia(\pc 1. Mi\t;N, I5.()5/325S3. from the hpc Icicalih. 14.9 x i).fi mm. 5, (i, 7. I'lotouoncli ol
paratope I. 5. (Iciieral \i('\\. Scale liar: 200 ijliii. 6. Dt'tail of the sciilptmf of the last wliorf Scale liar; 10(1 jinr 7. Drtail of tlit"
milieus ami tlic firsi wliorl. St-alc liar: 10(1 \y\n.

18-30 Ml iFiinta dc Tcmio ami Dicnlc ilr .\|()) ami 42-4S
m (\'eril de Masca). An averagt' iiiinilicr ol 5-6 speci-
mens per antipathariaii colonv were ohsencd in Pnnta
de Teno, with a nuLxiniuin of 10-12 specimens, and np
t(j 20 on a single antipatliarian colonx in \eril de Ma.sca.
Of 27 .specimens checked for their posilinn on three col-
onies, 20 were with the si])hon directed upwards and 7
do-Anwards (()|i\erio, pers. eonim.). No ohservatiuns of
(ceding on I'ik I'laek coral have been made.

Etymology: "flic specific name comliines to the nick-
names ol the soii'^ ,i! tin first two authors: Kao (Hicardo)
and Ofito i Rodollo

Remarks: Comllini'hila kaofilonini is tenlalixcK as-
signed to the genus C'^uallinplula liecan,se it resemliles
otiier species (C. cunhara. (,' brcvAs) that are presentK
included in that genus, jiemiiue a generic revision of the
laniiK.

Coralliopltila kaojilnniin differs from its closest rela-

ti\(' ('ordllicjiliilii lii'ciis iBlaiusille, lS.-)2i. which is also
pi'escnt in the ( ^auaiA Islands and has lu't'U collected at
the liase of the \-ellow (lj>j)ln>i!,or^ia riiiiiiidlis^ and red
(/,. nilnnimd^ gorgouiaus, oil Puerto de la Cnv/.. at 20-
.'52 m depth. ;uid oil the northeiii side ol the Teno Pen-
insula, in identical en\ ii( inmental conditions, Coriillic-
jiliild lirciis is ,1 longer species mp to .'lO-tO mm'. \cr\
variable, espeeialK in spire length and shape ol the aji-
erturc and the siphoual canal: the spiral sculpture con-
sists ol 15 25 (illen uiarkedK imbricate, alternating
bioad and thin cords: lrec|ueulK one or two more swob
len coi'(b on the upper p;irl ol the spue acceutu:ile the
carene ol the w liorl. In the .Mediterranean, ('oi'dlliopliilii
hici /s lives also on iioi'gonians {Pcirtniiiirircii cliiimiic-
Iciiii. I'.iniiicllii •■ilricliL l,(ijilii)^<>r<iia saniiciilosti) (Olive-
rio, l«S9b: Po|)pc and Coto, 1991).

('ortillidpltila riirihricd .•\bl)()tt, 195S. from tin' West-
ern .'Mlantic is a similar species, but it is somewii.it solid
and has a wider ajierlnre usuallv purple on the inside.

R. \V>'a ft al.. 2002

^ai'e 53

It lives on a varieh' of antliozoaiis intlmliu'j; sclcractiiii-
ans. gorgonaceaiis. z.oantliiileans and (x)ralliiii()ipliaria
(Wells and Lalli, 1977: Miller, 19S1: De Jong and Con-
mans, 1988). The enibnonie shell ol C.onilliopliihi r/ir-
ihaca was illustrated hv Bandel (1975), and the piotu-
eoneh and lai"\'al shell h\ De |iing and (,'ooinans ( 1988)
and Leal (1991). The prottieuneh of f.'. rnrihrica has its
nucleus dcnscK eowred 1>\ small piistnles. is somewhat
longer (4.5 wiiorls, 1.2 mm ieiigtii), and has mon- pi'om-
inent nodules than tliat of (', kdnjilnnnit PrestMitK mo-
lecular insestigation is in progress 1)\ Marco C)li\'erio to
assay the relationships of the ni-w species with Condlio-
phila hrcvis and Coralliophila airihucd.

The new species also differs from C.nniUiojihilii mctj-
endoi-ffii (Calcara, 1845), another littoral species of the
Canai^v Islands wideK distributed along the .Meiliterra-
nean and Eastern .\tlantic. wliich has a longer (nji to 40
nun) and more soliti shell, with 5-fi teleoconch wlioiis
with about 13-15 spiral cords antl 8-10 strong axial ribs,
canal rather short, and umbilical aperture. In the ('anan
Islands, C incijvmloi-ffti prt-xs on Aiuinon'ui sidtdhi
(Perez-Sanchez and Moreno-Batet. 1991), and in the
Mediterranean the species is known to \nr\ on \hi-
monia suIcatiL Chidocora ciicspilosa. anil BiddHiiplnjllui
airopaca (Sabelli and Spada. 1980; C)li\erio, 19S9b),

The Atlantic-Mediterranean ('(irdlliopliilii sipininosd
(Bivona, 1838) has a longer (up to 40-50 mm) and more
soliil shell, with 8-10 iLxial ribs and 12-20 major spiral
cords, alternating with the same number of lamellose
cordlets, the siplional canal is of medium lengtli and tlie
umbilical aperture evident (Bouchet .md Waren, 1985;
Oli\erio, 1989b). Corallioplnld s(iiidiiiiisd pre\s on flor-
gonacea or deep-sea scleractinians (Oliserio. 1989b).
CoraUiophild luontcrosatoi (Locard, 1897). from Spain,
and CordUiophila pwfundicohi Haas, 1949, from Ber-
nnula, should be considered junior s\non\ins ol (', stjiui-
niosa, according, respectixeh, to Bouchet and Waren
(1985), and Kosuge and Suzuki (1985).

Cordlliiiphild hdsilciis (Dautzenberg and II. Fischer,
1896) reaches up to 27 nun, has :i thicker shell and
broader siplional canal. It is found on the upper p;iit nl
the insular slopes around Azores and the (^anan Islantls
(Bouchet and Waren, 1985); its prev/host is unknown.

C.nndliophild riclidrdi (P. Fischer, 1882) is a deep-sea
(batlnal) species tliat li\es (and probabK pre\s) on the
scleractinians Madrcponi ocidatd and l.iiplulid prrtiisd
in other Eiastern Atlantic localities (Bouchet and Waren.
1985; ()li\erio. 1989b. and obsenatious of the tliirtl au-
thor). The shell of this species is \en dillerent from
t\'pical C'ondlidphild. with 8-10 axial roli;ite \arices in-
sti'ad ol solid ribs and with relati\el\ lew spnal cords
without tlu' characteristic scaK' sculpture. (\indlio}>Uild
rii-lidrdi is included b\' Kosuge and Suzuki ( 1985) in the
genus Eiiiczdiiiid Iredale, 1929, along with the similar
Ciinillidjiliild jdrli Knudsen. 195(S. lonnd hniii Sierra Le-
one to Angola.

Comlliophila foiitdudngioiji Snuiglio and .Mariottini.
2000, describeil from Teno, Tenerife Island, is snuiller
(up to 7.5 nun) and has a solid biconical shiil. It li\es

on the scleractiniau Madracis rispcndd Milue-Ldw;n(ls
and llaime. 1850 (Perez-Sanchez and .Moreuo-Balet.
1991; Smriglio and Mariottini. 2000; authors obsena-
tioirs in Teneiile). which pi'ob;ibl\ is a MacaroTiesian eu-
tli'mism (Zibiowins, 1980^. .According to Smriglio ;uid
.Mariottini (2000). ('.oidlliopliila l(indldiidii<i,i(>i/i issimihu'
to CaraUioplidd dlliodii'^iddla ( !•',. .\. Smith, 1890) li'om
St. Helena Island. Comlliopliild ididniuiiddliis Kosuge
anti Fernandes. 1989. from S;io i'om(\ ('oriilliopliild pd-

rci Petneh. 1987. h l'l(irid;i, ;uid I'ordlliopliild iiirld

Sowerln; 1894. from .Mauritius Island; all of them clearK
diffi'r from f kdofilnniin b\ the smaller size and dif-
ferent shell sluipc.

The remaining; corallio|)hilid species found in the (."a-
nan Islands, Bdhcloiiiiii'cx ((iiiniffnis [C. B. Sowerbx,
1834) is included in a different genus on the basis of
conchological and anatomical li'atnres (Kosuge and Su-
zuki. i985; Hichtei' and Luijue, in press). This sprcies
pre\s on the scleractiniau Plii/lldiiiiid tnoiifliczi (authors'
obsenatious). whereas in the Mediterranean it pre\s on
('Iddiirnni iiicspiliKii. Asli'ciidcs itdijiiildrts and I'oli/ci/-
dtlius iniiillcrdc (()li\c-rio. 1989b. and authors' obsena-
ti(jns); ol these three kilter species, only PcdljOjiltluis
iiiiu'llcrdc might possihK be lonnd in the ( 'anar\ Islands
(Zibrowius. I98()i.

One other, [)robabl\ nntlescribed littoral species of
coralliophiliti is found in the (.'anaiA' Islands li\ingon the
scleractini;in Driidroplii/llid rdiiicd (from Lanzarote. 40-
50 in, C;usta\() Perez Dionis and Marco ()li\erio, pers.
comm.). It is a large (u]) to 40 mm) and solid shell. \ri\
similar to that iliusti;ited as ComUiophdd (i. jinii Irom
C;abon b\ Bernard (1984: pi. 29, fig. 120).

Tlie remaining .Mediterranean species of Condliophdd
are all \"en' dilTerent. Q'.ondliopUdd pinicniiHaiid i.Mon-
ti-rosato. lSfi9) is solid, with 10 :ixi:il ribs anil 22-24 spi-
ral cords with sm;ill sc;iles. This species lives at the base
of Fdrdiiuiiiccd cluniidlrdii :ind associated to ('ondliuiu
nihniin (Oliveiio. 19891)' or Epizoantlins arcuacciis
(Templado et ak, 1993). Cortdllnpliila saphiac (.Arailas
and Benoit. I87fi) has a solid, more or less globose shell,
similar to Ciiiidliiipliild sijiuniKisn in size, and li\es in
bottoms with strong conilligeuons eomponeut: its diet is
unknown (Olixerio, 1989b).

.\mong the easti'rn .Atlantic species, (.ondliopliilii dc-
(hiiiiiis (Watson. 1880) from Niiihtin^ale Island (Tristan
iki (!unha' li;is a stroni4 shell, with a high, scalariform
spire. relativi'K small aperture and umbilicus. C.ordllio-
pliild g//())i (Dautzenberg. 1891 ) from the (."ape \'erde
Islands and Sao Tomi> is a smaller species (up to 20 nun),
with a thick elongate fusiform shell usnalK eroded and
iucnisted, 7-9 broad axi:il ribs ;ind :ibont 27 s]iiral cords
on the last whorl (Kosuge aTiil Fernandes, 1989; Holan
and [•'ernaudes, 1990). Ccndlioplidd alldiitica (F. A.
Smith. 1890) from St. ilelen:i IsLiud is :i solid, small
species (17.2 nun), with low axial ribs and s])iral cords
with weak sculpture, ('ondliopli'da rn/llirost())>ui F. A.
Smith. 1890. also from St. Helena is solid, sub-rhom-
boidak and has a reddish ajK-rture and evident umbili-
cus. C'lirdlliiipliild pdlnnlis i Iv .\. Smith, IS90i, from St.

Paire 54

THF. \Al TILLS. Nol. 1 IH. NO 2

Helena, is small ( 10.5 inni). As far as we know, the three
latter species are onI\- known Irom t\pe material, illus-
trated and briefl\' described h\ Kosuge and Siiziild
(1985).

Coralliopliila kraemmeri Knudsen. 1956. from Nige-
ria, and CoraHioj)liihi marrati Knudsen. 1956. from Li-
beria, ha\e sohd, umbilicated shells; the first one has 16
axial ribs and 14 spiral ridges on tlie body whorl, whereas
C. marrati has rounded ribs and onl\ 8 prominent spiral
ridges (Knudsen. 1956).

Coralliophila occidentale Kosuge and Fernandes.
1988, from .\ngola is a small (up to 9.4 mm) and rather
solid species. Coralliophila adansoni (Kosuge and Fer-
nandes, 1989) from Sao Tome is also solid and has the
inner side of the aperture denticulate. FinalK; Corallio-
phila knudseni Smrigho and Mariottini, 2000, from I\or\'
Coast and Coralliophila schiottei Smriglio and Mariot-
tini, 2000, from Sierra Leone, are smaller and liaxe bi-
conical and more sohd shells than Coralliophila kaofi-
toruin.

The remaining western Atlantic species of Corallio-
phila are also different. Coralliophila aberrant (C. B.
.Adams. 1850) has a thick, globose shell with strong spiral
cords. Coralliophila galea (Ree\e. 1846) also has a more
solid shell and wider aperture. Coralliophila salcbrosa
H. and A. Adams, 1863. has an o\"ate-fnsiform shell, with
angiilate spire, and Coralliophila scalarifonnis (La-
marck, 1822) has a scalarilorm profile.

RECOMMENDATION

Tiie host of Coralliopliila kaofitonim. the anthipatarian
Antipalhc.s wolhi^toiii, forms in the Canarv Islands a
characteristic communit)- on walls below 50 m of depth,
but it can be found at sh;illower depths (18-24 m) in
ca\es or overhangs at certain sites (Perez-Sanchez and
Moreno-Batet, 1991 ). A similar itssemblage occurs in the
Caribbean islands with other antipathaiian species (Au-
tipathcs spp., Humann, 1993). Antipadies uollasloni is
only known with certaint) from Madeira. Selvageni and
Canan Islands, but it is probabK' also present in the
Cape \'erde Lslands (A. Brito, pers. comm.). It is pro-
posed for protection in the Canar\ Islands and included
in tiie Prcliniinarv- Red List (Bacaliado et al., 1989) and
the Threatened Marine Fauna List (Boimet-Femandez-
Tnijiiio and Rodriguez-Fernandez, 1992). We strongK
suggest to the environmental authorities of the Canan'
Islands C()\eniinent to include Coralliophila kaofitonim
in the li',! (li jiriiletttd species, since it is up to now onK
kn(jwn horn ili. ( aiiarv Islantls and is strictK associated
wiih Antij)iitUi-. :loII(i.sI<iiu.

ACKNOW LF.Dr;MK\TS

We are indebted l<i Marco Olixerio for information
about the habitat of C. knofitomm. and to him and Al-
exandra Ricliter for in Iplni comments on the manu-
.script: to Gu.stavo Pen/ Dionis for data on coralliopliii-
ids from the Canarx IshnKK; to Jose Bedova uMNC.N)

for SEM photographs, and to Rogelio Sanchez and Jesiis
Mufioz iMNCN) lor photographs: to Alberto Brito and
Rogeho Herrera for information about conserxation.
This work was supported h\ Fauna Iberica W project
(DC1C\T PB95-0235>.

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with Indo-Pacific rtih/thixi iCoelenterata: Zoanthidae).
Pacific Science -34: 1-17.

Rokin. E. and F Feniandes. 1990. Coralliophila adiinsoni i Ko-
suge and Feniandes, 1989) new name for Ocinebrina
adansoni. Bollettino Malacologico 26 (5-9): 14:3-1-14.

Sabelli, B. and C:. Spada. 1980. Guida illustrata airidentificazione
delle conchiglie del Mediterraneo. Fam. CoraUiophilidae.
Bollettino Malacologico 16 (7—8. suppl.': 3 pp.

Smith. E. \. 1890, Report on the marine niolluscaji fauna of
the island of St. Helena. Proceedings of the Zoologicid
Societi of Londcm 1890: 247-,317.

Smiiglio. C. and P. .Maiiottini. 2000. Description of three new
Coridliopliilidae from the Eastern Atlantic ocean. La Con-
chiglia 296: 11-18,

Spada, G. 1979. (Uiida prattica alia fbrmazione di una raccolta
di coiichiglie marine. Bollettino Malacologico 15 (1-2,
suppl.): 7:3-80.

Talavera. F. G. 1975. Molnscos de sedimentos de la plataforma
continental de Mauritania. Boletiu del Institute) Espafiol
de Oceaiioijrali'a 192: 1-18.

Teinplado. | . .\. Guerra. J. Bedma, D. Moreno. J M. Remon.
M. Maldonado and M. .\. Ramos. 1993. Fauna iiuuina
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la Campaua Oceanognilica "Fauna I ". .Museo Nacional de
Ciencias Naturales, Consc^jo Superior de Investigaciones
Cientfficas, Madrid. 138 pp.. 45 pis.

Watson. R. B. 1886. Rejiort on the Scaphopoda and Ciitstro-
poda collected In H. M. S. Challenger during the \t^ars
187:3-76. Report of the Ciialleiiger Expedition, Zoology
15(42): 756 pp.. 50 pis.

Wells. F. E. and C. M Lalli. 1977. Reproduction and brood
protection in the Caribbean gastropods Coralliophila ah-
breviata and C. raribaea. )ounial of Moiluscan Studies 43:
79-87.

Wicksteu, M. K and K T Wright. 1993. Predation h\ Latiaxis
oldroifdi (Gastrojioda: Coralliopliilidae) on Con/nactis cal-
ifoniira (.\ntlio/.oa: Corallimoqihidae). The \'eliger:36: 92.

Zibrowius. H. 1980. I^>s Sderactiuiaires de la Mediterranee et
de r.Mlantiijne nord-oriental. Memoires de I'lnstitut
()ceanos;raplii(iue. Monaco II: 1-227.

THE NAUTILUS 116(2):56-5.S, 2(){)2

Patre 56

Fusi}ms dovpch'di, a new species (Gastropoda: Fasciolariidae)
from the Red Sea, and range extension for two other species

Martin Avei-)' Snyder'

Departiin'ut ol Midacologx
Academy of Natural Sciences
19''' and Beiijaniiii Franklin Parkway
Piiiladeiphia.' I'A 19103 USA

ABSTRACT

Fusinu.s dovpcledi new species from the Red Sea is descrihed.
The new species is distinguished hv shell characters from the
similar Red Sea species F. leptorht/iicliua (Tapparone-fJanetri,
IS75) and from F. coins lon^caiida (Lamarck, ISOl) from
southeastern Africa. New range records are provided for F.
hifmns iSUudm. 1900) and F aralnnis (MeK-ilf 1898).

INTKODUCTIOX

tni

A new species in the li;cihis I'lishius is ilcscrilied ticiiii
the Red Sea aiici tiLxoiioinic affinities are cliscussed. Most
Fiisiniis are subtidal burrowers, although some species
are tound in deep water. Tlie discoNciv and description
ol this new species is part of a trend in recent wars in
which a nnniber of new fasciolariids have been nanietl
(e.g., Gofas, 2000; Hadom and Rogers, 2000; Snyder,
2000; Snyder and Snvder, 1999; and others). Specimens
ol the new species described herein ha\e been collected
by fishermen at a depth of .300 m. AdditionaJK. the re-
discoverv- of Fusiiiii\ hifrciis iStunuiv. 1900). another
Red Sea species, with a ran<j;e e\lension lo the Indian
Ocean, is reported. ,\ probable range extension is also
reported for Ftisiniis anihicus I VleKill, 1S9S). Tin- ab-
breviation USNM, National Mu.seum of Natural lliston.
Smitlisonian institution, W'asliiii'^lon DC. is used here.

SYSTEMATICS

Family Fasciolariidae Gray, 1853
Snbfaniil) Fusininae Wriglev, 1927
(^i-uns Fusinns Rafines(|ue. IS15

'Ivpe .species: Murex coins Linnaeus, 17.5S. Hecent.
hidu \'v. ~: Hacific. by monot\-py.

Ftisiiiiis ildijtclcdi new species
(Figures 1 2'

Descriplion: Sh H elongate, .si/e moderate lor genus,
length to 107 imn. Mplional canal long, slightK nndnlal

' Research Associat*

ed, apex subtending an an<j;le ol ;ippro.\iniateK- 29\
Sculptiu'e of iLxi;il libs and spir;il cords, ribs lorming
prominent angular knobs on later whorls, with knobs
hilling approximateh at the centi'r ol whorl. V>ud\ whorl
with 11-13 ri])s, penultimate and earl\ wliorls with 12-
14 ribs. Earlv whorls ofparatype (onlv) lightly sculptured
with ;Lxial ribs, becoming evanescent. Axial ribs crossed
1)\ unuierous spiral cords, some lirowii on later whorls;
strong brown cords alternate with weaker unctjlored
cords on bodv whorl. Penultimate whorl witli 12 strong
cords, becoming evanescent toward the suture, luid un-
uierous weak, fine, axial lines, visible onlv under mag-
iiificiition, between axial libs. Aperture tvpicallv ovate
;iud elongate; parietal shield shiny, waxy white, extending
onto siphonal canal. Spiral cords bene;ith shield raised
to lorin lirate sculpture. Canal long. thin, somewhat sin-
uous. Operculum claw-like, light brown. Piotocoucli.
periostnicum. and nulula unknown.

Type male-rial: llololxpe USNM 90.3651. length 107
mm, width 24.4 mm, spire height 40 mm, live-collected.
P;u'at\pe, IVled ( Collection (d;un;igi'd), length 72. S nun,
width 22. S unn, spire ;iiid siphon. il can.il broken, dead-
collected. Ii'oui t\pe loc;ilit\.

Tvpe loealilv: f-led Se;i oil Neviot (Egvpt), 75 km
soutli ol F.hit (Israel), .300 m depth on s;nidv bottom.

Etvinologv: The species is named ;ilter l)o\ P<'le(l
(Tivon, Israt'l), a student, colleetoi' and dealei' in the
shells ol the lU'd Se;i ;u('a.

Distriliulion and liabilat:

c'alilv.

Know u onl\ li'om t\pe lo

Diseussion: I'nsiiiiis (hnpclcdi is most closely rekited
lo i hplnihijuchus (T;ipp;u'one-( 'auelri. 1875'). which is
well illustrated ni Shanib;iti (19S1: pf 23, fig. 7). The
knobs on F. Icplorln/iirliiis. loi'med b\ the ;Lxial ribs, are
lewei', immbeiing just 9, ;ind ;ire uioi'e S(|uare th;m those
on I (liii jHlcdi. The opi'iculnni ol /' Icplorhtjnchns is
distinetivek reddish brown. ,uid the shell is sm;iller. tvp-
ic;dl\ r;iiiging between 7.5 and SO unii. with ;i propor-
tioiKitek shortei' siphon;il c;mal. These dillerences ;ire
snllicient to iide out F. dovpvlrdi Ixing a deeper-water

M. A. Sinck-r, 2002

Page 57

Figures 1-4. Species of /•//,s(i///,s, 1, 2. /■//s;//((.s doipchdt neu species. I. Ihilohpe. I'SNM U()3(i51. 107 iniii leii<;tli. tioiii saiuK
hottnni off Neviot (Eg\pt). 75 km sontli of Elat (Israel), Ued Sea. deptli 300 in. 2. I>arahpe. Peled Collection. 72.8 mm leiisitli.
from hpe localiU. 3. Fusinufi bifrons (Sturam-. 1900), Peled Collection. 99.9 mm length, from Elat, Red Sea (Israel), depth .3.50-
400 m. 4. Fnsinm arabirus (MeKill, 1898). Snyder Collection. 96.6 mm length, trawled hy fishermen oil Kl Tnr, Gulf ol Suez
(Egvpt), depth 32^0 in

Paee 58

tup: XAITILUS. \ol. llfi. \<

lonii of F 1c))lorhi/nrlnis. Comparison can also l)t' made
with /'" coins hmgicnuda (Lamarck, 1801) troni the east
coast of southern Africa (Cernoliorsk)', 1972: pi. 4S, fig.
la); that species has 10 axial ribs on the penultimate
whorl. Ho\ve\er, whereas the luunber of axial ribs in-
creases toward the posterior end of the spire of F. clo-
ipeh'di. the opposite is tnie in F. rolus hmisicauda. The
axial ribs are not neark so pronounced on tlie bodv
whorl in F. coins hmiijcaiida as in F. dorpclcdi; the for-
mer grows to appro.ximateh' 200 nun.

Fiisimis hifrons (Sturanx; 1900)
(Figure 3)

.Sturan\' (1900:197) introduced a deep-water species, Fn-
siis hifrons, dredged in depths of 490-900 m in die Red
Sea. He compared this axialK' ribbed species to F. for-
ceps (Perr\', 1811), F. multicarinatns (Lamarck, 1822), F.
torenma (Desha\'es, 1843) (a sviionvm of F coins (Lin-
naeus, 1758)), and F. leptorhijncns (Tapparone-Canefri,
1875), and he named a smooth varietv' of his new species
"form pancicostata." Later, Sturan\' (1903: pi. 1) illus-
trated F. bifrons and erroneoush^ referred to the taxon
pancicostata at specific rank. According to R. Janssen of
the Senckenberg Museum, Frankfurt (reported prixatelv
by R. Hadorn). the t\pe material of this "form" differs
from that of the nominate species onK' in sculpture and
this "form" should be considered a synonym of F. bij-
rons.

Two IreshK-collected specimens of the smooth form
of F. bifrons were examined, one 99.9 mm long, 26.fi
mm width, with broken siphonal canal (Peled Collection,
from the Red Sea), the other 106.5 nun long, 26.9 nuu
width (Sn\der (Collection, from the Indian Ocean). The
smaller specimen was dredged on a sand\^ bottom at
.35()-4()() m off Neviot (Eg\pt), 75 km soutli of Flat (Is-
rael), Red Sea. The larger specimen was collected by
Russian research expeditions to the Sava de Malha Bank
in the Indian Ocean (Bondare\ and Roeckel, 1992; Sir-
(Miko, 1995). The larger specimen was illustrated b\' Sir-
enko (1995: fig. 7). This represents a significant range
extension for this species from the lied Se:i lo the indi;ui
Ocean.

Fnsinns anihicns

(Figure 4i

McK

1898)

.■\ siL'iii'icMiii r inge extension of Fnsinns arabicns (Mel-
sill 1S9S- IS ;ilsu reported. This species, previously

known! from the Culf of .Arabia, is figured in Bosch et
al. (1995: 136, species 578). Two specimens were trawled
by fishermen in the 19S0s off El Tur, Gulf of Suez
(Eg\pt), in depths of 32—40 m. Both specimens were
trawled as dead shells and hence this range e.xtension is
stated without absolute certaintx'. E\en though taken In
commercial trawlers, with notoriousK' xasjue collectinsi
data, it is \irtuall\- certain that the specimens were col-
lected in the Red Sea rather than the Arabian Sea. These
two dead-collected specimens are identical in ;ilmost all
respects to Arabian Sea specimens of F. arabicns (Mel-
vill, 1898).

LITERATURE CITED

Bondarew I. and D. Roeckel. f992. The shells of Sava de Mal-
ha Bank. La Conchicrlia 262: 21-38.

Bosch. D. T. S. P. Dance, R. G. .Moolenbeek and P G. 01i\en
1995. Seashells of eastern Arabia. -Motivate Press. Dub;u,
296 pp., col. pis.

Cernohorsk). W. O. 1972. .Marine Shells of the Pacific, II. Pa-
cific Publications, Sydney: 411 pp., 68 pis.

Gofas, S. 2000. Four species of the familv Fasciolariidae (Gas-
tropoda) from the north .\tlantic seamounts. loumal of
ConchologN' 37: 7-16. figs. 1 — 3.

Hadorn. R. and B. Rogers. 2000. Re\ision of recent Fusinus
(Gastropoda: Fasciolariidae) from tropical western .Atlan-
tic, with description of six new species. .Ar^onauta 14(1):
5-57, pis. 1-16.

Sharabati. D. 1984. Red Sea Shells. Routlcdsie 6c Kegan Paul.
London: 12S pp., 49 pis.

Sirenko, B. I. 1995. On the fauna oi shell-bearinij molluscs in
the Sa\a de Malha Bank. Indian Ocean (part 2i. La C^on-
chiglia 276: 20-24, figs. 1-15.

Sn\der. .\1. .A. 20()(). Latiiiis licckt/cc. a new species ol F"ascio-
lariidae (Gastropoda: NeoCTastropoda' Irom Bra/il. Tin-
Nautilus 114: 161-163. figs'; 1-3.

Snxder, M. A. and N. C. Snyder. 1999. A new species from
Columbia. South America: Fiisiiuis cohunhicnsis n.sp.
(Ga.stropoda: Fasciolariidae I La Conchiglia 291: 21-22,
figs. 1-1.

Sturans, H. 19110. Diagiioscn iieuer Ciastropotlcn aiis dem
Rotlien ,\leere. .\n/.eiger der Kaiserlichen .Akadeniie der
Wissenschaiten. Matlieniatisch-NatunNisscnsciiaftliciu'
Classe 37(17): 197-201.

Sturaii\, H. 1903. Gastropoden des Rotlien Meeres. Bericlite
der (Commission fiir Oceanographisclie Forschungen. Fx-
peditionen S. M. Sciiill. Pola' in das Rotlie Meer ncirdli-
clie nnd siidliche lliilltc l.S9.5/9r>-I,S97/9S. Zoologisclie
Ergebnisst" 23. Denkschriften der K:iiserli<iien Akadeniic
der Wissenschaiten. .Matheinatiscli-N'atni'wissensiiialdi-
che Cla.s,se 74: 209-2S3. pK. 1-7, 1 text fig, (.Mso issued
as a separate, pp. 1-75).

TIIK NAUTILUS 1 l(S(2);59-fil. 2002

Viv'v 59

The systematic position of Str()]nbi)ia (Cotonopsis) J'nidae
Petuch, 19SS (Gastropoda: Columbellidae)

Helena Fortunato

('niter tor Tropical I'lileot'ColoaA anil

Arclu'i)I()ti;\
SinitliSDiiian Tropical Rcscarcli

Institute
r, (). H(i\ \m
Balliiia. PANAMA
lortiinae(S'ancon. si.edu

ABSTRACT

The s\steniatic position of Stroiiilnim limliic I'clneli I9SS, is
discussed and revised. The species sliould lie allocated in the
colunihellid genus Cotonopsis Olsson. 1^)12 The shell char-
acteristics that define the two known Cotoiiojisis subgenera. (.'.
(Cotonopsis) and C. {Tiirrind). are iteiui/ed and contrasted.
The holot\pe of S. liiulac is re-illnstrated and the pmtocouch
Hiiured tor tlu' first time.

INTRODUCTION

111 his rcNlsion of the Stroniliiiia o;r()up, Jtiny; (19S9) gaxe
Cotonopsis Olsson. 1942. t^eneric status ami recogiii/ed
two subgenera, Cotonojjsis scusii stricio Olsson. 1942,
and Cotonopsis {Tiiniua) Jung, 19S9. Cotonopsis is ivp-
fesented bv 18 species, two of them known only as fos-
sils, and 16 extant. The majoritx of the living species (12)
are found in the eastern Pacific. Two species are loinid
in the Caribbean Sea (Houbrick. I9S3: Fetncli. I9.S8):
one species in West Africa (Emerson, 1993); and a
fourth species in the Andaman Sea (Kosnge, Rous.sy and
Mnaiiginaii. 1998: Kronenberg and Dekker. 1998, 1999).
.\s noted b\ Kronenberg and Dekker (1998). this distri-
bution miglit indicate an earlier origin ol this geims. i.i'.
prior to eaiK Pliocene as proposed by Jung ( 19S9). On
the other hand, bodi the West African anil the Asian
species ina\" be part of separate lineages, the laiik ol
which can onl\ be addresseil pcniliiig aiiatoiiiical and
molecular data.

The aims ol this paper are to correct the svstematic
]iositioii ni Stroinhina iColonojisis^ limlur Petuch. 19SS.
to discuss the subgeneric position ol tins species, and to
rc-illn.stratc the holotvpe deposited in llie National .Mu-
seum of Natural Historv. Smitlisoiiian Institution
(USNM 859942). with special attentimi to llir proto-
conch. This latter task was prompted b\ bntli die poor
ijualitv ol the original illustrations and the lack ol iiiior-

iiiatinii nil the 1 1 lorpl lologv ill the pi otoci inch III tile orig-
inal description.

syst1':m.\tk:s

FaiiiiK ( Jiiiiiiilii'llidae Swaiiisoii. I S4()
Cienus Cotoiiojisi\ ( )lssi)ii. 1942

Coloiiiipsi^ oKsoii. Ii)l2: 227 i7o'. T\pe sjiecics (In- original
desi^iiatiuii 1; SI ro nihil III Coloiiojisis • iiiiiuicostiirii-cnsis
OKsoii. 1042 lOlsson. I!)t2: 7."x pi, 1(1. fig. 10'.

Deseriplioii: Shell small to large > 1 l-5(S iimiL general
shell shape \ar\ing Iroiii lairK stout to slender. Most
species lightlv sculptured with axial ribs predominantly
on earK teleneniiili whorls. Outer lip usnallv wcaklv
thickeueil. both cohiiucllar and parietal callus usually
present, weakK to well developed. Parietal ridge and
posterior canal present, prominent in several species.
Reclined and well-developed anterior canal. Proto-
coucli smooth with I'i to :5 whorls, i according to Jung,
19SS.)

Siibgeiiiis Colonopsi\ sinsn sirioto

Description: Sti.iit shells, inllateil bodv vvlioH. axial ribs
on earlv teleoioiieh. well developed parietal and colu-
mellar callus, weaklv developed ixirietal ridge and pos-
terior canal, and smooth protocoiieli with I ': to 3
w liorls.

Cotinnijisis (Cotonopsis^ lindiir (I'etiieh. I9SS1 new

eombination

(Figures 1-6)

SlioiiibiiKi tCotonopsisi lindfir Petuch. 199S: 161-lfi2. pi. .3S.
figs. 15-16 (Off St. James, Badiados. trawled TO in

depth, 1

Descriplion: Shell small, stout, up to 22.9 mm in
length, mllated bodv whorl, axial ribs on the eariv teleo-
com-li. relativelv well developed colimiellar and parietal

Page HO

TUF, NAUTILUS, Yd. 116, \(

1^

Figures 1—4

l,atenil \ic\\.

:>'^

('(>li>U(>ii\is {('(tidiiojisi.',! liiiddc lioldhpc. I'SWI S.i!)iM2, lieiij;lit 22. SS iiiiii, wultli VJ.dO mm. 1. .\pcrtuml \ic-\\. 2.
ri^lil siili' 3. Miapcitiiia! \ic\\. 4. I'mtdciiiuii aiul caiK Iclcdrniii-li wlidils.

calluses. I'lolocDucli siikioIIi, witli 154 wliorl.s and .ilicnil
TOO fiiii iliaiiiclcr.

Discussion: 'I'lic hvo siil)y;cncra rccocriiizcd In' Jmii!;
! HISS I (lillcr ill several inoipliological Iratiircs. In con-
trast to the ciiaractcrs altoxc dcscrilicd lor ('dtdiinjisis
sciisu slrirto, species ol Cotoiidiisis {'runiiia) have
smooth. niostl\- slender sliells, with well- and sometimes
heavily (le\(>l()ped pari(Hal ridge and posleiioi laiiai; pa-
rietal and colnniellar callnses missing or vcia weak, and
a smooth protoconeh with 2 % to 3 whorls. Based on
shell eiiaracteristics. C. liudac is host placed ni die snh-
genus C(>li)iii>j).sis scnstt slrirfo.

Colotiopfiis {Titniua) seems lo he reslncled to (lie
eastern I'aeific aTid has no known fossil reconl. ('dlnii-
(tpsis sciisu stricio has a much wider disdilmlion, in-
eliuling llie eastern Pacific, the Carihlx-an .Sea. the West
■\tri( an coast and the .\ndainan .Sea. StratigraphiealK.
the latter ranges from the earK Pliocene ol' l'",smeraldas,
Keuador. .ind ( "harco \vw\. Peninsula Burica, Costa Hiea,
through (he Rei ent. As mentioned ahoxc, the recent dis-
tril)utio;i ■)! th.is snljgeims seems to indicate an origin
older than (he I'iinr cni'.
This is the sni.illi St of all lour species repoited li-oni

outside (he eas(ern Pacific, ('(iloiinjisis liiiil/ii' resemhles
(,'. i>liukclc)isis (Kosnge, Houss\' and .\Inangman. 1998)
Ironi the \ndanian Sea in its reduced sculptural ele-
ments, hut the la((ei" is .i mueli kirger .species, with an
incised suture and a protoconeh with a larger number
ol whorls. Cotoiiopsis fin^ciitcfi llouhrick. 19S.3, from the
Dcinnuican Hepulilic and (.'ii/iiiio/jxi.s monfilsi Emerson,
lOO.i. Ironi Si'uegal are ,ilso unich larger, with hea\ier
sculptured teleocoucli whoils. Besides. ( '. inoujilsi lacks
axial rihs, hearing insteatl sjiiral lines. Its protoconeh has
(hree whorls. The ])rotoconch ol C. ar<irnlc(i is similar
(o (hat ol ( ', liiidiic in si/.e. lin( the lornier has a weaker
colnmell.ii (■.liins.

.\mon'4 till' eastern I'aeilie species, (he diiiiensioiis ol
(' lindiic are similai' (o Coloiiiipsis iitcndozaiKl iSli,isk\.
1970^ from the ( aiH' of h'onseea, Kl SaKador. lint (he
i.ittei h.is .1 iiineh less reciiiAed siplioiial ean.il. more
liea\ll\ senlplnred e.ilK teleocoucli wliiirls. \\ell-de\el-
oped eiilnniell.ii callus, and ineiinspicuDiis liiae on ils
ou(er lip. SimilaiK (o (' liiidiic. ('()li»uipsis isincraldcit-
sis (Olsson. 19(iU Ironi the I'.smeraldas pro\ince, I'x--
iiador. also lacks lieaw seiilptme and li.is alinost (he
same dimensions: |iii( i(s siphonal canal is lnn'j;ei ,nid it
has a iiioic strongK dexeloped eoliiinellai" callus

11, Fortiiiiatd. 2(102

Fauc 61

Figures 5-6. Ci>tono))sis (Cotonopsis) lindac. liolotNpc,
USXM S59942. 5. SEM of protoconeh, sliowins; a slijj;litl\ sig-
moid !ip. and the sculptuie of tlu' carK' tt'lcocoiit-li wluirls. 6.
Same, apical \ie\v. Sade lines = 200 (xni.

ACKXOWI.KDCNIKXTS

Special tliaiiks arc due to |ern- llarasewvcli (USWl)
who kiii<ll\ photoiiraphed the liol<)t\pe and helped in
main other \\a\s. Tlianks also to two anoininous re\ ■Jew-
els who kindl\' read the pajicr and made helpful sug-
"estioiis.

LITERATI RE CITEU

Emerson. \\. K, KW'v .\ new species of coluinhcllid gastropod
from the old world tropics. The Nautilus 106: 147-151.

Houhrick. f-i. S. 19S.">. .\ new Stromhiiia species (Gastropoda:
I'rosohranchiai from The tropical Western .Atlantic. I'ro-
ceedinsis of the Biological Societ\ of \\'ashin<iton 96: 3-19-
354.

Jung, P. 19S9. Re\isioii of the Slnmihinti-(.',nmp (Gastropoda:
Columbellidae), fossil and li\ing. Distribution, biostratig-
raphv and svstematics. Mcinoires Suisses de Paleontology

in:'l-29S.'

IvoMiiie. ,S,, P. il, Hoiiss\, and I'-i^, Mnangman. 199S. f^eport
on the fauna of Thailand ( 1 i with the description of a new
species ((Columbellidae and Ikiccinidae). Bulletin of the
Institute of Malacolog\ ol Tok\o 3(5): 75-76.

fCronenbcrg, G. C. and II. Dekker. 1998. A new .species of
Colonops!.\ Olsson, 1942, from an unexpected l()caljt\
(Gastropoda Prosohr.iiK hi.i: (Columbellidae'. X'ita .Marina
45(3-4): 11-16.

Kronenberg. G. G, and II, Dekker 1999. ('Dloiiiipsis viiiintil-
Ic'^liciiii Kronenberg i^ Dekker 199S. a junior s\iion\in of
Siroiiihiiiii phukclcii\i.s Kosuge. Kousss 6: Mnangman.
I99S, with som(> notes on the generic position and colour
pattern i GiLstropoda: Pro.sobranchia: Golumbdhdae). \'ita
Manila 46(1-2): 69-72.

(Olsson. .v. .\. 1942. Tertian and Quaternary fossils from the
Burica Peninsula of Panama and (.'osta Hica. Bulletins ol
American Paleoutologx 27 if06): f53-25S if-I()6!. pis.
14-25(1-121.

Petiich. E, ]. 19SS. Neoneuc hislon oi tropical ,\merican niol-
Insks. The Coastal Education 6c Research Foundation
(GERE), Charlottesville, 1-217 pp.

THE NAUTILUS 1 16(2):62-65. 2002

Page 62

The reproducthe anatomy, taxonomic status, and range of
Orcohclix (ilpi)i(i (Elrod, 1901) (Gastropoda: Puhnonata:
Oreohelicidae)

H. Lee Fairbanks

Pcnn State L'ni\ersit\'
100 l)niver.sit\ Dri\p
MdiKK-a, PA 15001 I'SA
lill lCap.su.edu

ABSTRACT

rlic icpii)ilucti\i' aiialiiiiix (it spcciiiii-iis cil (h'iiiliili\ iiliiiiiii
Iroiii the t^pe loeaiitx in llic Mi.s.siciii Mountains ol western
Montana is fij;in"ed ani-l eonipaied with that ol speeiiin'us col-
lected in the Swan Mountains, approxiniately 20 kilometers to
the east. It appears that O. ctlpina is found in hotli the Mission
Mountains and the Swan Mountains, which ecmstitutes a range
extension lor the species. In addition, the reproductive anato-
niv ofO. (ilpiiiii is compared with that olO, suhnidis iipaiium.
Orcoliclix (ilpiiia lias been incorrectly placed in Pilslins ()
.suhnidis liroup ol species and should he included in the O.
4-/rii^().v« <j;roup

INTHOUUCTION

Tlie lieiius ()ri-()licli\ (Pilslm. UJ()4) is cduiposcd cil a
larije iniinlu'r ol s]iccies loniid throimlioiit tlu' Hock\"
Mountains Ironi southern ("anada to soutlieru Arizona
and Xcw .Mexico. .\ hin^v anionul of iutraspecific \aria-
tioii in shell shapt' an<! color is present in tlie i;;('nus, and
it is olten dilficult lo dclevniiiie wlicllicr the eairse of
the \ariation is cnvironniental or nenetie (Filslnx. 1939).
There is, however, little xariation in re|>rodneti\e anat-
oiTiy: Piishrs- (1939l round no iclexanl inleispeeilie var-
iation in the niorphologv ol the nproduetixe s\st( in
Nonetheless, that author, based in pari upon llie ralio
ol tlie plicate (ridijed) portion to the pusliilose portion
ol tlie internal surlaee of the penis, allocated s|)ecies of
Orcolirlix into one of three groups, llie () s/r/ijosY/
group, tlie O. i:hn((lis group, or the () i/iniipal group.

Elrod (1901 i d.scnlied Orro/zc/a' alpiiia from (lie Mis-
sion Mounlams ■ .1 western Montana on the basis of the
shell only. With no I now ledge of the internal aiiatoni\ of
that species, l'ilsbi\ !,S9,) tentatively placed O iilpiiiii in
the (). siihniilis group becmse O. sithnidi.s occurs at lower
elevations in t]ie saiai- inountain range and because no
members ol the other .'roups occur in the area.

Speciuiens of Orcohclix (dpiuc from the txpe localits
had not been collected lor neark 50 \-ears. lieceiitK,
however, Hendricks (199S^ collected and reporied on
several specimens from the t)pe localit\, some of which

he sent to rue lor dissection. The goals ol the stuck' were

(1) to figure the reproductive anatonn ol O, olpiiid and

(2) to determine its taxonomic position relative to I'ils-
bn's three Orcohclix groups. A third goal was to com-
pare additional specimens similar to O. alpiiia Irom a
localitx' in the Swan Mountains to those Irom the t\pe
localit\ ill the Mission Mountains, which is located ap-
pidxiniateK 20 kilometers to the west.

MATERIALS AND METHODS

Specimens ol Orcohclix iilpiiui were collei'ted i Hen-
dricks, 199S) from the southeast ritlge of St. Maiys Peak
which is near the t\pe localitx on the southwest ridge of
St. Mans Peak, Tl'SN H1S\\''S21 (Township, Range, and
Section Irom USGS topo map). Mission Mountains,
Pake (JoinitN, Montana, elexation approximatek 2S00
meters, on 27 Aug. 1997 (Figure 1 ). .\dditioiial speci-
mens of Orcohclix. similar to O. alpiiui. were collected
b\ Bill llamiiier. on 7 -\ug. 1974 aboNC Rumble Lake.
,Swaii Mountains {T2()N RlfiW S2), Mis.soiila Comitx,
Montana, elevation approximatek' 2400 meters (Figure
1). Two topotxpes and three siieeimeiis from the Swan
.Mount, liiis vvei'c dissected .iiid llieii reproductive svs-
lenis removed, for coniparalive purposes, the repro-
ductive s\ stems ol three specimens ol ( ), suhnidis (ipdr-
imii Bern. 1919. collected on 2 Aug. fy75, at Vellow
Bav, Flatiiead Lake (T24N RI9W S4). Lake CJoimtx,
Montana, elevation ap[iroxiiu,itelv S9() meters), were
iisi'd. The reproductive svstein ol each specimen was
li.ieed Irom the moiiitoi' ol a dissecdiiLi inieroscopeAid-
eo svstem, llie penis ol eaeli speeiiuen was then dis-
seeled lo .illovv loi tracing ol theli inlernal snrlace. Mea-
surements ol the plicate and the pusliilose portions ol
(he inlernal peiiial surlaee wer<' made using an ocular
iiiici'omeler. llie di.iwin^s ol one s|')eciineii were used
as represeiilaliv<' ol llie popiilalion, I he diameler, height
ol shell, and numlier ol vvlioiis ol the speeinu'iis Irom
llie Ivpe loeaiilv and those Irom llie Swan Mdiinl.iins
were measured. N'oiielier specimens ol O. ulpinii Irom
the tvpe Iocalit\' and from thi' Swan .Mountains are de-

H. I.. F'airhanks, 2002

Pasre 63

1 MISSION MTS

Figure 1. Wcvstiiii Montana sliimin'j; tlic Imatinns nl llic
Mission and Swan nionntains. Stale l)ar = 2(1 km

posited ill tin- collt-ction of niollusks at tin- National Mu-
seum of Natural lliston-, Sniitlisoniau lustitutidii. W'asli-

inatou, D.C, USA (USNM 100233:
1()02353).

1.1 rsN\i

RESULTS

Siiolis lidiii till' Mission Moiilltains ami lioni llic Swan
Mountains arc shown in Figures 2 ami 3; their inea-
siiri'iuents are given in Tahle 1. The gross reproclueti\t'

%^ ^b

Table 1. Siicll rncasnreincnts lor Orcolii'lix alpiiui. Measure-
nienls in millimeters.

IJkimelei ol \nnil)er ol

shell Shell heiijht whorls

Mission Mountains

Speeinien A 8.0

Specimen B 7.9

Specimen C 7.5

Specimen U 8.4

Mean 7.95

Swan .Mountains

Specimen .\ 11,1

Specimen B 8.5

Specimen C 9.0

Specimen D 8.6

Mean 9.:'5

4.5
4.6

4.6
4.57

6.4
5.0
5.0
5.2
5.4

3.3
4.0
3.5
4.0
3.70

4.3

4.25

4.0

4.0

4.15

* Shell was [lamat^ed dnriiiii the takint; of njeasurements.
Hi'i'Jit could not lie olilaini'd.

anatonix III individuals troin all I luce localities are shown
in Figures 4-0, and the inteiiial penial aiiatoiiiies are
shown in Figures 7-9. Tahle 2 suniniarizes the iiiea-
sureuients ol the internal penial aiiatoun'. The plicate
poi'tion of the intt'rnal penial surl.iee ol the Mission
.Vhnnitains (Figure 7i and the Swan Mnuntains (Figure
iS) specimens was less than 50', (il the total length in
contrast to that nl (I suhriidi's (iiuiriiiin which was great-
er than oO'/r.

DISCUSSION

The reprochietive organs of Orcoliclix alpina from tlie
tvpe locaiih' (Figures 4 and 7) are t\]')ical ol the genus,
and no diagnostic eharai'lers at the specific level were
detected.

Pilshn' (19.')0i noted thai ni the (). s/r/go.sY/ group the
plicate poiiion dI the p<'ins was ". . . decidedlv less than

Table 2. Me.isiiremenis ol llie plicate porti<ins and imstnlose
portions ol llic mleni.il smi.ices ol the ])eni.scs ol Oivohclix
tilpiiKi .uid () siilinidis iijiiininii. Measmemcnts in millimeters.

Total I'lisliilose I'licate Percent
len^tli ])ortioii portion jilicate

O (ilpiiia. Mission .Mountains

Figures 2-3. Shells ol Onolulix ulpiiui 2. From the tvpe
localitv in the Mission Mninitanis 3. Kiom llie Sw.ui Moiin-
tains. Scale har ~ 5 miii

Speilnieii .\

7..T

4.1

:5.4

45

Specimen H

10.0

5.6

4.4

44

( ) iiljiiiKi. Swan

Mountains

Specimen \

7.5

4.4

3. 1

41

Specmieii B

7.5

4.6

2.0

.39

Speeimeii ( '

6.5

4.0

2.5

38

O. sulinidis (ipdiiinn

HLF.59-1

17

7.5

9.5

56

HLF.39-2

15.5

7.0

8.5

5.5

11LF39-3

14,5

6,5

S.u

.5.7

Pa"e 64

THr: NAUTILUS. \o!. llfi. \(). 2

(iP

F'igures 4-(). IvipiiKliu-tiM^ ssstcrns ol spctics ol Oirnlulix.
4. O. (ilpitta lioiii llic Mission Mountains. 5. (), aljiiiiti Ironi
tlic Swan Mountains, (j. (). siihriiilis apdriiiiii Iroui tlif Mi.s.sion
Mts. A(; = albumin ^land; K = epipliallns; Gl' = genital poR";
OV = ovotestis; PH = [icnial ivtiactor muscle; V = penis; S
= spermatheea; U = uterus. Se;ile liars; l^'ignics 4 ;uul 5 = 1
mm; Figure (i = 5 mm.

half..." the total pcnial Iciimli. liiil ^rcMlcr ilian luill
tllc total peiiial lciii;tli in hotli the () siihntilis aronp
and the O. i/tiiajxii groii[), thi'se two gidiips then lieing
separated on die iia.sis of whether the ;inlerinr hall ol
the penis \v;is " . . . decidedly .swollen . .' or not.

Fiishn originaliv placed O. aljiiiid in the () Mihnulis
group hecause no species from the other groups ((). stri-

Figures 7-9. Internal penial analomx' of species oi' Orcolic-
lix. 7. () alpinu from the .Mission Mountains. 8. O. suhnulls
(ipai-ium from tlie .Mission Mountains. 9. O. nlpinii from the
Swan .Mountains. E = epipliallns; C,V = genital pore; PH =
penial retractor mnscle; P = penis; VL = plicate surface; PS
= pnstulosc surface: \' = vagina. Scale bars = 1 mm.

H. L. F'airliaiiks. 2002

Faire fio

oosa and O. i/avnpai) are fbiiiul in tlic Missinn Moun-
tains. Ho\ve\er, tlic lensrtli of tlie jilicate pciition ol tlic
penis of (). iilpuui was less than 1/2 tot.il [leiiis length.
whereas tlie [iliivite poitioii ol (.). siihnuli.s (ijKiiiiiin. also
from the Mission \h)untains. was <j;reatei' than 1/2 total
pi'nial lentjth (Table 2). In the O. tjUKquii <j;r()n]) tlie
plicate portion ol the penis is also greater than 1/2 total
penial length, lint in addition the pro.xinial portion of tin'
penis is enlargeil. which is not the case for O. alpiiia
(Figinv 4). Therefore, if the groupins; s\'steni of Pilsiin'
(19.39) is to lie lollowfd, I suggest plai'emi'nt of (), al-
pina in the (), .sV/'igosy/ group.

The shells ol Orcniiclix alpiiKi lioni the .Mission
Mountains anti the shells Ironi the Swan .Mountains
(Figiu'e 1) could not he separated niorphologicalK (Fig-
ures 2 and .3). C.oinparisons of reproducti\e svsteni mea-
surements for these two populations (Table 2) did not
appear signifieantK different. Therefore, it appears that

these two populations belong to the same species, and
tliis represents a range extension lor (). tilpiiici.

LITERATURE CITi:!)

licrn. .S. S. 19H). \Ii)lliisca ol Clacicr National Park. .Montana.
Procccilint^s ol the .Vcailciiiv of Natural Sciences ol Phil-
adelphia 71: 19.5-21)9.

Elrod, M. |. 1901. Montana slulls. Ho(k\ .Moiuilaiii Ma<;a/.iiie
2: 691-697

Hendricks. P. 199S Krilisco\crv ol Discua hndii.stini Bern',
19.5.5 and OiciiIkIix alpiiui (Elrod, 1901) In the .Vllsslon
Mountains, Montana, with comments on Orcniiclix cirotii
(PIlshiT. 191)01. The Nautilus 112: .5S-62.

Pllshn, H. A. 191)4. (icnenil Notes. The Nautilus 17: 1.31.

Pllshr), H. A. 1939. Laud .Mollusca of North .America (North
of Mexico), N'olunie 1. I\u-t 1. The .-Vcadcuu' of Natural
Sciences ol Philadelphia. .Monographs Niuuhcr 3, Phila-
delphia, wli + 573 -I- l\ IItuIcnI pp.

THE XAITII.US 116(2):6r>-ft7. 2(){)2

Pasie 66

Notes

Authorship and date of pubHcation of Ostrea chilensis Philippi
in Kiister, 1844 (BivaKda: Ostreidae)

Bruce A. Marshall

Museum of New Zealand Tc Pupa

Tongarewa
P(). Box"467
Wellington
NEW ZEALAND
BruceM@tepapa.go\-t.nz

Ostrea ch'dcnsis. a comnierfial cnster of New Zealand
and (>'liile iBuroker ci a]., 1983; Chanle\- and Dinamani.
1980; Jeffs and Creese, 1996; 6 Foighil et al., 1999; 6
Foighi! and Ta\lor. 2000), eonsistentK has been dated
from 1845 nnder anthorship of Philippi in all references
seen. The few autliors wlio cite the original reference
for O. chilensis (Dall, 1909; Hanley, 1856; Lamy. 1929)
refer to Kiister's edition of Si/stematisches Conchi/Iien-
Cahincl ion Maiiini unci Chemnitz, p. 74. pi. 13, fig. 7,
8. in N'ohniie 7. part 1. The text describing O. chilensis
("Philipjii in Lit.") is part of Lieferung 186, issueil in
1868 (Kiister. 1868) (John.son, 1968: 366; Smith and
Englami. 1937: 97; Welter-Schultes. 1999: 185). The tax-
on Ostrea chilensis (as "O. [strea] chilensis Philippi") in
fact originates from the list ol species depicted on the
plates inclnded in Lieferung 45 printed on the wrapper
thereof (Kiister. 1844). which bears the imprinted date
1844 and is assumed to ha\(^ been issued then in the
absence olCxidence to the contnuA i K]ZN. 1999: Article
21.3). Here Ostrea chilensis .st;intls technically as a no-
men nudum, for at the foot of the list is the statement
"W'egen Krankheit des Kuplcrstechers k;um Taf. ,N111
erst niit der niiciisten Lielernug ;insgegebeu wcrden"
("Because of tlie engraver's illness, Plate 1) will !)e dis-
triliuted with the next delixcn-.") In the iibsencc olexi-
dence to tiie contran it is ;ippropii:ilc to :iccept dial
plate 13 was indeed issued in the next Lieferung (which
is also imprinted 1844 and assumed to haxc been issued
theni. and that the name (). chilensis became available
fnjm the nomcnclatural standjioiut at tlu> time of issue
ol that Licfenm,;. I Note that pi. 13 was erroneonsK at-
tributed to Lielciuug4l. and Lieferung 45 erroueonsK'
dated 1843 by Welter S.hultes ( 1999: 185), but the latter
correctly dated elsewliere in the compilation.] Although
wrappers for this work were intended to be discarded,
and most indeed were, tax;i introduced on ihem with
reference to illustrations (uKill criteria for availabilit\-
(ICZN Articles 8.1, 12). Tivxa introduced on these wrap-
pers were acceptetl as published b\ C. D. Sherborn (in
Smith and Engkind, 1937: 89. ' nomen el /igifw"). who
failed, howexer, to include Ostrea chilensis in Index An-

iinaliuin (Sherborn. 1922-1933). The authorship and
date of publication of O. chdensis are thus Philippi in
Kiister, 1844 (Kiister, 1844).

.ACKNOW LEDC;-\1ENTS

For checking accession records tor parts of Si/steina-
tisclies Conchi/Uen-Cabinet von Moiiini und Chemnitz
and other works. I am grateful to R. Janssen (Sencken-
berg Museum, Frankfurt). .-K.L. Kiibat (Washington,
D.C.), D.G. Reid (The Nattual Histon- Museum, Lon-
don), A.H. Swann (Wheldon and \\esle\- Ltd., Leighton
Buzzard), A.. Waren (Swedish Museum of Natunil His-
ton-. Stockholm), and F. W. \\'elter-Schultes (Unixersitiit
Gotiugen). Special thanks to S. Jordan (La Habra
Heights, California), who supplied a photocopv of a
wrapper of Lieferung 45, and to P. Bouchet (Mnsenm
Niitional ilHistoire Naturelle, Paris) and A. L. Kabat for
extensive comments on the subject.

LlTER.JiTURF CITED

iiiu.ikcr. N K,. W Chanlev, If | Cnmlidd, 1", Diiuini.ini. 19S.3.
Svsteniatic status of hvo instcr populations oi the ijenns
Tiosirrd honi New Ze;ilancl and Ohilc. Marine Biologv 77:
l()l-:2()(),

C.'lianliv. V. and T. Dinamani. 19S(). ( .'onqianitive descriptions
oi some ovster lanae irom New Zealand and (,'liile. and
a description ol ;i new i^cnus ol ovster, Tiostrcii New Zea-
land |onrnal ol \1. nine' and Freshwater Heseauli It: 103-
120 ■

D.ill, W II U)09. Report on :i collection of shells Irom I'c'ni,
Willi .1 snmniai'V ol the littoral marine Molliise:i ol the
l'rriivi:m /oologieal province. I'loceedings ol th<' I'nited
SI. lies National .Museum .')7: 147-294.

Ihiiilc-v. S. lS12-lS.5fi. .\n illnstiMlcd ,ind dcsiriplive citaloune
ol Kecc'iit bivalve shells Willi, mis and Nor<;:ite. London.
W'iii + .'W2 pp.

UrZN (Inteni.itional ('omimssioii on Zooloijical Xonniiilie
lure). 1999. Inteinalioiuil (lode ol Zoolos^ical Nomencla-
ture. Fourth Edition. International Trust lor Zoological
Nomcnrhiliirc. London. .306 pp.

Jells, \. C. and IL C. Ca'cese. 1996. Ovcimcw and hihliogra-

p. Cal

ZOO

'd''v 67

pli\ (it research on the Chilean cnstcr Titistrcd rhilviisis
il'hilippi, 1S45) from New Zealand waters. |i)urnal ol
Shellfish Research 15: 305-311.

|iiliiiMiii. H. I. 196(S. Martini and ( 'heninit/ iKiKslers cilitionl
S\steniatisches Conchvlien-Calmiet. 1S37-192(). a com-
plete collation. Jonmal ot the Societ\ lor the Bil)lio^r.ipli\
of Natnral Histiin- 4: 363-367.

Kiistcr. H. C. 1844. Inhalt [list ot species depicted on plates
inclnded in Liefennig 45: pi. 13, howexcr, exidi'utK issned
ni l.icrcrun'.i 46. 1844]. S\steniatisches ( jiiicli\ lii'ii-( .'ahi-
nrt \oH .Martini nnd Chenniitz (Kiister, II. (^ cd.^ 7 i I ) 45
wrapper Baner and Raspe. Niirnherg. (Phot(K-op\ in Mol-
liisca Section libran'. Mnsenni ot New Zealand Te I'apa
Tonsjarewa).

Kiister, H. C. 1868. Conchifera. Zweischaline Mollnskt'ir ( 'mi-
chitera inononi\aria. .Mnschelfhieri' niit eineni Sclilii'ss-
nniskeln. Svsteniatisches C'oncli\lien-CJabinit \i>ii Martini
nnd Chenniitz (Kiister. H.C. ed.) 7(1) 186: 5(1-S4. Baner
anil Raspe. Niiniberg.

Lani\. Iv U)29. Revision des Ostna \i\;tnts dn Mnsruni N:i-

tional d'llistoirc Natnnllc dv Paris. |oin'nal de (.'onch\-

liolot^ie 73; 1— 4().
6 Foighil. I) . B, .\. .Marshall, i j, llill.ish and M. .\. I'ino.

1999, Transd'acihe ranije c\ti-nsion 1)\ raltini; is iiilerred

for the Hat osster ()\lrici c/ii/r/is/s. IMologiciil Bnllctin 196:

122-126.
(J Foighil, I). :Liid I). |. I'.ivlor 2lll)(l. I'Aoliitioii ol parent. il

care and oxnlation lielia\i(>r in ovslcrs. .Molei iifii' l'li\lo-

genetics and K\ohition 15: .301-313.
.Sheriiorn. C O 1922-1933. Inde.\ .-Vninialinni 1801 IS50,

British Miiscuni (Natnral lliston), London; Longmans,

Green ami Co.. B. OM;iritch. Dnlan, ()\lord l'ni\crsit\'

Press. Wheklon and Weslew and Oliver and Bo\il.
Smith, E. ,\, and 11. W. Kngland. 1937. NLulini and Chemnit/

( tinester's edition' Sxstematisches Conch\lien-Cal)inet.

1837-1918. Jonrnal of the Si)ciet\ lor the Bililionraphx of

N:itin:il IlistoiA 1: 89-99.
Wclter-Sdiiillcs I" \\ 1999. S\ stenKitisches Conch\lien-Cal)-

inct \on M.iitmi mid ( 'licnmit/ i 1 S37-I920'. !iil)liograph\

of the Milnmcs m (Kiltingin Archives ol N:itnral lliston

26: 157-21)3.

Tadashicre Habe (1916-2001

Paul Callonion

Department ol Malacolo'j;\
The .\cadem\ ol Natmal Sciences
Philadelphia.' P,\ 19103-1195 US,\
callomon(£!'acnatsci.org

Tadashige Halie, who died at S5 on Dccciiilier 29. 2001,
was one of the most prodnctixe and iiilhieiitial /odlogists
of the twentieth tentun'. He plaxed a nia|o|- pait in tlie
development of malacologA' in [apan anil the western
Pacific, and authored a large mimlier ol hooks that set
the stanikuiis \nv the stniK ol the swstematies and dis-
trihiition ol niciliiisks Ironi that area ol the wcirlil,

l)i. lialie was horn March 31. 191fi, in the village of
Hioki, part of what is now Sasavama in Hyogo Prefec-
tnre. He attended Ikeda High School in neighhoring
Osaka Prefecture, sta\ing on to wmk as an assistant
teaclier until 1939, Electing to stiidv zoologx at Kvoto
Imperial Universit^■ from 1939 to 1941, he then trans-
ferred to Seto .Marine Lahoraton in Sliiiali;inia. return-
ing to Kvoto in 1945. Dr. Hahe had joined the stall :it
K\'oto as a junior researcli assistant in 1942 and liecanie
a full research assfstant in 1949. From 194fi to 1950 he
also served as an adviser to the Natural Resources .Sec-
tion ol .\llied (U'lieral Heaili|uarters in Tokvo.

.\lter receiving his doctor ol science degree ni 195i.
Dr. Haiie was appointed assistant professor at Kviisini
Universit\' and moved to the Maiine Biological Laho-
raton' on .Amakusa Isl.ind in westi'rn Kvushu, lie re-
tained his position at Kviishn liniversitv lor five vears
after moving ni 1962 to the National Science Museum
in Tokvo (NSMT).

Dr. Hahes career at tlie NS.MT lasted from his ap-
pointment as researcher in 1962 to his retirement as
head of the /oologv depiirtnient m 19S0. I le then seneil
for four vears as a professor at the Ocean Institute ol
Tokai Liniversitv. where he remained |irofessor emeritus
until 19S9. From 1985 to 19SS he w:is also director of
tlie Tokai L'niveisitv Museum of N.itnial llistor\. In
19S6 Dr. llahe was awanled the Order of the Sacred
Treasure iZuilio-shoi |,\ the |apanese government lor
his contributions to /oological reseaicii in Japan.

Dr. ilalie lirst puhlished formally as co-author ol a
1935 report on shell collecting in .Manchuria and Korea.
For more than hftv veais he wrote |)roi!igiously. alone
.ind in cd-.inthorshiji with all the other senior Japani'se
inakicologists of the post-v\ai- period. Several ol iiis
hooks are well known outside japan, including ,S7(('//.s of
lite Wcslcni Facijic in Color il964i and S/(c//.s of tlw
World in Colour (\ol. 1. 19(i5. with Kivoshi Ito; \ol. 2,
1966, with Sadao Kosnge),

Pearlier in his career, Dr, llahe worked closelv with
Toknliei Kuroda ( 1SS6- 19S7 . whii had arrived at Seto
M;Mine Lahoralorv in 1940 and was to have a formative
inllnenee on tlie vounger man. The i\\o published their
first joint book, a svnopsis of the Japanese Helicac(>a, in
1949, This woik was followed In the exten.sive Illustrat-
ed Catidoiiuc of }a])(iiu'sv Shells (1949-1955) and the

I'.iilL' bb

TIIK NAUTILUS, \ol. IKS. No. 2

landiiiark Checklist find Bihlionriipluj of the Remit Ma-
rine Molliisca of Japan i 1952). Tlic latter was piililishetl
ill part as a result ol its Dr. llalie's work tor .Vllii'd Gen-
eral IIeacl(|uarters. and pro\ided the first thorough col-
lation oi Western and local literature on Japanese Mol-
lusca. In 1971 Kuroda and Habe were joined In Katsura
0\ania (1917-1995) in producing the lavish The Sea
Shells of Sa'fiami Bat/. This extensive work owed its ex-
istence in part to the favorable relationships all three
atithors enjoved with the Imperial household, a connec-
tion further reflected in a number o( subsequent papers
lo-authored b\ Dr. Habe and Prince Hitachi (Masahito),
brotluM' of the current emperor.

During the four decades that followed tiie .Second
World War, [apanese malacologv was to a considerabk'
extent sustained by the sponsorship antl participation of
a inunber of keen amateur collectors and authors. Dr.
Habe was an enthusiastic advocate of collaboration with
amateur collectors, and skillfulK helped chamiel the en-
ergv and resources of benefactors such as Tetsuaki Kira
and Rvosnke Kawamura into tlie production of useful
publications and scientificalK' significant collections. He
was instrumental in obtaining the vast Kawaunu'a collec-
tion lor the NSMT, and co-authored an impressive iio-
nographv based on it. World Seashelh of Raritij and
Beantij (1991).

Dr. Habes specialitv was taxonomy, and he was the
first Japanese malacologist to sene on the council of the
Internationd Trust for Zoological Nomenclature. In
1977. he published Si/stcmafics of Mollusea in japan:
Bivalvia and Seapliopoda. a complete iliustratetl taxo-
noniical treatment of both groups.

.Another important but less well-known lacet of Dr.
Habes career was his work in translating and sunnnariz-
ing developments in Western malacologv for the benefit
ol his Japanese colleagues. This took the form of a long
series of well-researched biograjihies of Western mala-
cologists and zoologists as well as reviews of books and
monographs aTuI rejiorts on Japanese l\pc spi^cimens in
overseas nmsenni collections.

He was v\'ell known among malacologists in the West,
and traveled overseas more olten tlian mauv of his con-
ti'inporaries. During his peiiod with Allied (General
Headquarters, he worked both with Dr. .\lv-in ("ahn and
Dr. Mvra Kec'u of Stanford Universitv. In later years, he
was a ri'gulai' visitor to the United States and a frienii
in particular of Di'. H. Tucker Abbott, with whom he
shared a broad outlook concerning amateur scientists
and popular publications. \'isitors to the NSMT during
Dr. Habes tenure alv\avs bi-nefited troni his renowned
hospitalitv.

A member from its early davs of the Malacological
Societv of [apan. Dr. Habe served as vice president from
I9fi:3 to 1979, as president from 1979 to 1995 and as
emeritus president until his death. In recent vcars his
failing evesight began to restrict his activities, but lie
remaineil an active contributor to conlerences and pe-
riodicals until the lati' 199()s.

In the course of his career. Dr. Habe introehiced more
than a thousand new names to science. Collations of his
new taxa wt-re published bv Inaba and Ovarna (1977)
and Okamoto (2001). The latter work is complete, in-
cludes a full bibliograpln and cites 976 new species
names anil .'ilo new genera. .-\ partial bibliographv was
publishedlnDr llal>e himself in 19S().

IJTKH.VniRE CITED

llahc. T, t9S(). Bibliographv of Tadashigc Halic. T llahe. To-
k-vo, 47 pp. [Partlv in Japanese. |

Inaba. T. and K. Ovania (Eds.) 1977. ( Catalogue ol nmlliiscaii
ta\u described In Tadasliige Hahe during 1939-1975.
vritli illustrations ol hitherto uiifi<;nre(l species. Okiiiaehi-
su-k;u. Chiba, 185 pp.. 7 jils.

Okamoto, M. (ed.) 2001, .Molliiscan ta\a tlcsinhetl li\ Tada-
shige Hahe. \'i>l. 1. (Gastropoda (excluding lleterohran-
chia): i-wiii + 1-S91. \'ol. 2, (lastropocki (Heterohran-
chia). Cephalopoda, Bivalvia & Scapliopo(hi): 892-1630 -1-
xix-vxxix + index 1—42 + xl-xli. Connnittee lor celehrat-
itig Dr, T. Habes eightieth birthdav, Tokvo. [Partly in Jap-
anese, I

THE XALTIIA'S 1 lfi(2):fi9-7(). 2002

Paa- 69

Book Review

Panamic Proiiuce Molhiscdti Literature:
Additions and Chauges from 1971 through
2001. 111. Gastropoda.

Sko^liind. C\ 2002. Fananiic Pnnincf Molliiscaii Littr-
attire: Aiklitioiis and C'liautics from 1971 tlin)ii!j;ii 2001.
III. Gastropoda. Tlie Festi\ii.s 33 i snpplcinent): i-\i +
1-286 pp.

PossibK' inspired h\ the ininiensits' oi A. M\Ta Keen's
Sea Shells of Tropical West Aiiicrica (Keen. 1971) and
elearK' based on a liietime of stncK ol tlie subject, lor
\ears C^arol Skoiilnnd has been updatin'j; the in\enton'
of Panamic niollnsks as ori<j;inalK jiublished in Keen's
wiirk.

In the couchision ol a recent ie\ie\\ ol Skoglunds \ol-
unie on bivaKes and chitons I Leal, 2001, re\ie\\' ol Sko'j;-
knid, 2001 >' 1 had wished that the autlior eonld provide
an equixalent npilate on tlie gastropods. That work, also
pnblished as a supplement to The Fi\linis_ is now avail-
able. The exhaustive compilation results from vears ol
record keeping, as attested to b\ the long list ol perti-
nent publicati<ins In the author in that periochcal. autl
SkogluiKl's nnqnestionable grasp ol the subject of Pan-
amic mollusks. The general lorniat follows tint adopli'<l
for the previous parts. Although most ol the updates
consists of the inclusion of new species. Skoglnnd also
presents newlv adopted use of svnonvnis, new combi-
nations (these latter t^vo categories indicated bv Keens
original species number), as well as thi' occurrence of
supraspecific taxa new tor the region (indicated in bold-
lace).

In a similar lashion to die volnme on bivalves and
chitons, the new publication is spiral-bonntl and [iro-
tected bv a plastic cover Although the compilation is not
illustrated, reference to original works and existing fiti-
ures is abrmdant and seems to be complete.

Like the previous volume, the gastropod update cov-
ers the coast, shell, and slope between 25° N and fi° S.
Most welcome bonuses are the extensive treatments of
mollusks from oceanic islands in the tropical easterii Pa-
cific, (e.g., Clipperton. Cocos, and Ciorgona) and ol those
associated with geologicallv active features, sucli as hv-
drothermal vents and cold seeps, of the adjacent deep-
sea areas. The onlv oversight I noticed in a brief rea(hng
of the suprageneric taxa were tiie misspellings ol "(Joc-
cnlilormia" [Cocculiniformia] (pages ii, 23i, as a subor-
der under the subclass Neritimoipha. Skoglnnd used the

' luadverteiitlv, in the title ol iiiv review I citeil the liivalve anil
chiton volume as ", . .from 197i to 2(1(11 vvIkmi it slioiild have
been ". . .from 1971 to 2()(»(l," M\ apoloLjii's to the autlior , mil
editor.

correct s[)elhng for the subclass (^occnliniiormia (same
pages), and 1 wonder whether the misspi'lli-d woril was
meant to be "( .'oci iiliiioida oi" a re|)etition ol "('occii-
liniformia. "

The hierarchical lanked anangeiiicnl ol lainilies and
genera generallv lollows that adopted bv Keen. In manv
cases, however, those were modified through the atlop-
tion of more recent taxonomic rearrangements, inelnd-
ing some ol the groupings nltimatelv compiled b\'
N'anght (19SS) and the inclusion of additional, mostiv
deep-sea, lainilies anil geiier.i Mi.il h.ive been intro-
duced in the last 30 vcirs. iiiainlv through the ellorts
ol |. 11, Met, can anil others. 'I'lie ailvaiK'cs in our nn-
derstaiRling ol gastropod phvlogenv over the past lew
decades (e.g.. Ponder and Lindberg, 1996. 1997) have
resulted in a higher classification of the Class Gastro-
poda that differs significantiv bom that incoiporated in
earlier works. While this new classification is being
adopted in manv recent works on gastropods (e.g.,
Beeslev et ak. 1998; Higo et ak, 1999; Okutani, 2()()())
the opportiinitv to update the taxonomic arrangement
of Panamic gastropoils has been missed in Skoghmd's
work. Tliis. however, shall not be a major handicap; tiie
author's chief objective is to give a consistent update
on the inventon and nomenclature of Panamic gastro-
pods, providing links both to Keen's book and the orig-
inal references, lii mv opinion, she has achieved what
she set out to do with great success. If voii use Mvra
Keen's work, voii must have these updates.

The cost in the US is S35 postpaid, overseas S5() (air-
mail onlv 1. Contact the San l)ie',io Shell Club. % 3883
Mt. Blackburn .Ave.. San Diego, CA 92111, USA, or e-
mail Carole Hertz (cmhertzC«pacbell,nel) for further in-
lorm.ition.

LITERATUHK CATED

Beeslev. P. 1... C. |. ti Ross and A, Wells <e(ls. i 199S. Molliisca:
the Soiilhern Svnihesis. Fauna of .\iistralia. \'oI. 5. CSIKO
Puhlislmii;. .Melliourne. .5fi'5 pp. Part \l pp. 565-1234
pp (Part B'

Higo. S.. P Callomon and V. (ioto. 1999. Catalogue and Bih-
hographv of the Marine Shell-hearing Moliusca of Japan.
F.lie Scicniihc I'nlilications. Osaka. 749 pp.

Keen. A. \1 I97F Sea Shells of Tropical W'e.st America: Ma-
rine Moflusks from Baja California to Peni, 2'"' edition.
Stanford L'niversitv I'rcss, Stanford, \iv + 1064 pp.

Leal, J, II. 2001. liook review: Panamic Province Molluscan
Literature: .\dditions and Chani;es from 1971 throui;li
2001 12000]. The Nautilus 115: .37.

Okutani. 'I", leifi 2000. .Marine Mollusks in Japan. Tokai Lni-
versitv Press, Tokvo, xlviii + 1173 |-H2] [)p.

Ponder. W. F .mil 1), Lindberg. 1996. Gastropod phvloiienv:
challenges lor the 9()s. in: J. I). Tiylor (ed.) Origin and
Evohitionai-v K;idiation of tin- Moliusca. Oxford University
Press, Oxford, pp. 135-151

Pagr 70 THE NAUTILUS. \ol 1 Ifi, No. 2

Ponder, W. F. and D. Lindbcrg. 1997. Towards a plixlogcnvol N'anglit, K. (,'. 1988. .\ (;Ia.ssification of tlic U\intT Mollnsca.
gastropod niollu.scs: an anaKsis using niorpliological cliar- .Kniciican Malafologists, Inc., .Vlelbonnie [Florida], xii +

acters. Zoological |(iurnal nl llu- Liiuican Socii't\ 119: 189 pp.

•'^^-seo. j^^^. ^^ Leal

Skoglund, C. 2001. Fanaiuic l'i(.\MKv \l.,llnscan Literature: -[-|,,. Hail,.v-.\l:ittli,.\\s Shell .Musnnn

Additions and (,'lianges troni 1971 throngh 2000. 1. Bi\- p(^ gox 1.580

alvia. 2. PoKplacophora. The Fcstixiis 32 (supplement): Sanibel, FL .>59.57 US.\

i-v + 1-119 iBi\al\ia): 1-20 i PoKplacopliora). jleal@.slicllnins(inn.org

This piihlication sponsored in part 1)\

till' State of Florida. Department ol Stale

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THE NAUTILUS

Volume 1 16, Number 3
September 30, 2002
ISSN 0028-1344

A quarterhj devoted
to malacologij.

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OCT 0 9 2002

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EDITOR-IN-CHIEF

Dr. Jose II. Leal

The Bailey-Matthews Shell Museum
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MANAGING EDITOR

Christina Petrikas

The Bailey-Matthews Shell Museum
3075 Sanihcl-Captiva Road
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EDITOR EMERITUS

Dr. M. G. Harasewych
Department of Invertebrate Zoology
National Museum of

Natural Histoiy
Smithsonian Institution
\\'ashington, DC 20560

CONSULTING EDITORS

Dr Riidiger Bieler
Department ot Invertebrates
Field Museum of
Natural Iliston*
Chicago, I L 60605

Dr. Arthur E. Bogan

North Carolina State Museum of

Natural Sciences
Raleigh, NC 27626

Dr. Philippe Bouehet
Laboratoire de Biologic des

Invertebres Marins et Malacologie
Museum National d'Histoire Naturelle
55, rue Buifon
Paris, 75005 France

Dr. Robert H. Cowie

Center forConserx'ation Research

and Training
Uni\ersit\' of Hawaii
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Dr. Robert T.Dillon, Jr.
Department of Biolog)'
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Florida Museum of Natural History
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Dr. Cvharles Lydeard
Biodiversity and Ssstematics
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Department of Malacologv'
Natural Histon Museum
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Department of Living Invertebrates
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Histoiy
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Dr. DiarmaidO Foighil

Museum of Zoology and Departnu^nt

ol Biology
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Dr. Gustav Paulay
Florida Museum of Natural Histon
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Mr. Richard E. Petit

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Dr. C;aiy Rosenberg
Department of Mollusks
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Natural History Museum
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TH E€7NAUTI LUS

Voltimc 116. Siimhcr 3

Srpfcinhcr 30. 2002

ISSX 002H-1344

CONTENTS

S\en \. Nielsen Tertian Xt'iioplidriilae (Castropixla^ ol western South

Tlionias J. DeN'nes America 71

Jose VVillibaldo Tlioine Hedescriptioii ol tlie (j;emis and species llvtcrovainniiui

Suzele Kodrigues Cioiiies liiiuit/aua (Lesson, 1830) (Gastropoda: Soleoliiera:

Rosanc Souza da Silva Wronicellidae) 79

Kristiina 0^"aska Aiiatoiiix of the droiiiedai-\ juinpin'^-shii^. livinplidlUi

Lvie C^liichesler ilromcdinius Bi'aiisou, 1972 (CJastropoda; St\loiiiiii:iliipliora:

Heike Reise Arionidae), with new distributional records S9

William P. Leonard
Jim Haui^h

Philippe Boiiehet New species ot deep-water Caneelhiriidae ( (laslropoda^

Richard Petit from the soutliwestern Pacific 95

Guido Pastorino Spawn ol' the Pata^oniaii nastropotl I'liriiilliria phunhcd

Pable E. Penchaszadeh (Pliilippi, 1S44) (Bnccinidae) 105

Sponsored in part by the State of Florida, Department
of State, Division of Cultural Affairs, tfie Florida Arts
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FOR THE ARTS

THE NAUTILUS 116(3):71-7.S, 2002

Paije 71

Tertiaiy Xenophoridae (Gastropoda) of western South America

Sven N. Nielsen

(ii'ciIdt^iscli-Palaontiilot^isrlics liislitiil

and Must'iini
Universitat Hanilmrg
Bundesstrasse 55
20146 Haiiibuis;
GEKMANV
iiielsenS'geowiss.iiin-luiiiihiin^.tlc

Tlioinas J. I3e\'ric's

liiirkc Miisiiiiii III \aliiial llislon .iiid

Ciilliiiv
I ni\ciMl\ (il Wasliiii'^tdii.
Sraltic, W \ 9.SU)5 USA'

Al^STHACT

Tlii'cf spcrifs ol Xi'iiDplimular aic ri'|)(irlc'il Imiii die TcrtLiiA
ol western Soutli Amenca: Xiiiopliord iiiniitiiS.i'rii new species
from the Oligoeeiie of Peru. Xciiophnni piniliiiHf new species
from the Miocene Na\idad Formation of central Chile, and
Stclliiiifi kririicrhiiiihdhli new species from the Tertiar\ ol Pen-
insnla .\ianeo. sonth-eential ( 'hile. .\11 ol tliese species are first
recortls ul .Xenophoiidae lor tlic respe(.ti\e conntrii's 'iTie rc-
lationsliip ul Sunlli Aniernan \i iiiijilioni species wjlli 'I'ertian
New Zealand tiL\a and tlie tlillicult\ in identilxTng a species
closelv related to the other .South American Siclltiria are dis-

INTRODUCTION

Xeiioplioridae ar<' a u;iniip i>l cxclusivcK wann-water ma-
rine gastropods. The liioloin,. fossil liiston. and taMiiioinv
of 25 recent species has heen re\ie\\ctl In Ponder
(19S3). who recognized the single genus Xnioplidrti Fi-
scher \()n W'aldlieini, 1807, including the subgenera
Xciioplioni sciisii sfriiiii. Oiiiisfn>; Swainson, ISdO. and
Stclldrid Schiniclt. 1S:52. IIowe\cr. most workeis now
recognize these at the generic rank (e.g.. Kicijil and AH,
1999). a \iew that is followed liere. Where known, the
protoconc'h in Xcnoiilioni species consists of aliont .IS
low trochospiral whorls I Bandel. 199:3: pi. 12. Hg. 1), a
t\pe of moq^liologx that indicates planktotropliic (le\i-l-
opment, which in tiu'n suggests long-distanc'e dispeisal.
Ponder (19S.3I also described Tertian fossil species
from Australia, wliile in an earlier work. Ben (1977) re-
viewed the Onozoic Xenophoridae of New Zealand
which include an Eocene to Miocene species, a Miocene
species and a Pliocene to Recent species. Until now, Ce-
nozoic Xenoplioridae ha\e never been recorded from
Pern or Chile.

' Mailniu ackhess: Box 13(161. Burton. WA 9Sl)f3 USA.

(;k()1.()(;y of fos.sif-hfahixc i,(K,Ai,rriFs

Pi.sco B,\si\, Pkhh

The Pisco f^.isin of southern Pern i Fignri' 1 ) is a forearc
basin (Unnbar et ah. 1990) with four tli'positional se-
quences containing upper Eocene. Oligocene. lower to
miildle Miocene, and middle Miocene to lower Pliocene
shell and littoral deposits i DeWies. 1998). Specimens of
Xiiiojihiiii: are most often found in ma,ssi\'e, bioturbated,
medinm-grained sandstones of the Oligocene Otnnia
Formation behveen Paracas and i5aln'a de la Indepen-
dencia (Figure 1), together with \aKcs of Canlilti iicw-
clli Ri\era. f957. The .\'r/i(i/)//(i/v/-bearing sandstones in-
lerpicted as shallow -w .tier nearshore slielf deposits,
overlie a basal transgressi\f si'i|nence of nu>dium-bed-
ded coarse-grained sandstones with uearK monospecific
mollnsean assemblages of rtirrilcllii or Ostrcd. and nn-
ilerlie a thick se(|ueuee ol thin-bedded, tuffaceons. fine-
grained siltstones with thin-shelled \al\<>s ol Chlmiujs
and fish scales of sar(fiues and anchovies (DeWies,
I99S1.

Internal molds ol probable Xcuophoni species are also
found in a fault-bound outcrop of ]iebbly coarse-grained
sandstone o\erlookiu'j; the lower iii'o lea \alle\ i Figure
1 1. The molluscan fauna associated with the molds sug-
gest a late F.ocene age and hence assignment to the Par-
acas ( )ioiip,

NwiDM) {■"i IKMVTION. ( KNTRM, (jllI.K

Tile Navidad Formation (Darwin. 1846) is known for a
rich molluscan fauna that was last re\iev\'ed extensively
by Philippi (18S7'. Sediment deposition as indicated b\-
microlossils ranges from intertidal to outer shell. Most
of the spi'cinu'iis of Xcnopliorn descrilied iiere have
been recovered from gra\ deepwater siltstone tliat today
forms the intertidal platform at Pnnta Perro (Fignre 2).
These deposits have been datetl with foraininii'ers hv
Dremel (in Herni. 19(i9. p. 71 ) as Power Miocene (Bnr-
ch<falian>. However. Tsnchi et al. (1990) and ll)araki

Page 72

THE NAUTILUS, \'ol. 116. No. 3

75°35"W

~^

.1 ^.d^i i

75 "W

PANAMERICAN
HIGHWAY

Figure 1. Fossil loculitics lor Xcnopluini ctiiditiiH-rii in the Pisco Basin ot Pc

(1992) lia\v poiiitcil out thai an Uppci' Midfciic acjc
(Tortoiiiaii) is nKirc likcK.

.■\cklitioiial spciiinciis lia\c hccn colhilcil ihhIIi iiI llic
Ri'o Hapel and Iroiii Malanzas. l)()tli also in the aioa ol
Na\idacl (Fissure 2). Tlic scdimciils ol liicsc localitirs arc
believed to he (•(intrrniiorancoiis with tliosr ol I'nnia
Perm.

.\Iii,i.()\f;ri'; Foumaiion. Pkmn.sula Ahai'co. sen iii-
cjENiHAi. (.'im.i-:

Tlie .Vlillonirne P'onnation has heen defined from coi'es
drilled on Peninsula .\rauco. It consists ort^nix- sillstones
dated a.s Kocene (e.g.. Garcia. 1968). .\\\ similar sill-
stones of this area usnallv ha\e heen eonsidcicd to he
long in this f(jrniation. Ilovvex'er. analvsis ol tlu niollns
can (anna suggests a .Miocene age at least loi pail ol
tlie.se .sediments (.S. Nielsen, unpuhlished dataL

The Strlldria specimen descrilietl herein was loniid in
a concretionar\- nodule from a coastal cliff with e\po
sures of gray siltstones to the east of Pnnta Millongne
(Fignn^ 2). These concretionan- nodules are washed Iree

h\ the tide and iisnalK \ield specimens of the crah ('iiit-
ccr finiiii(mii\ I'hilippi, ISS7. Because this localitx has
not heen d.iled until now. the age of the Stelldria rc-
ni.iins nneert.iiu.

MATFHi Ai.s wi) Miriiions

.Specimens deselihed in mentioned in this stud\ .ne de-
liosiled in the (dileetlons ol die lollowinij; miiseiims; l)e-
partameiilo de I'.ileontolo'^ia de \ eitehiailos. Miiseo dc
liisloria Natural de la Uni\crsidad de San Marcos. Lima.
Peru (MIISM IN\'): Departamento de Paleoutologi'a de
jnxcrtehrados. Miiseo Nacional de llistoiia Natural.
Santiago de ( liile (SCO. PI): and Senckeuhert; .Museum.
Fr.uiklnrt, (a'rniau\- {SMP"). Photographs were taken iis-
iiil:, a Leicalle\ SLil camera. Images were scanned Irom
lllord LP I \2~> hlack and while .'li mm negatives using
.III \eei SeanW'if 272()S film seaimer and processed with
Photoshop (i.O.

S. N. Nielsen and T. T. DeN'ries. 2002

Fatie 73

Figure 2. Fossil localities tor Xriiniiliiii'ii
Chile.

piiuliiKir (\a\i(la(l arcal and StvUiiria kririurhdrtholili (Peninsula Aranco' in Central

SYSTEMATIC PALE0NT0L0C;Y

FaniiK Xenoplioridae Philippi, LS53

Cienus .\V/(()/)/)(i;v/ Fischer xon Waklheiiii, ISO?

Type Species: Xcnopltoni l(ic\:i'^(itii Fiselier \iin W'ald-
lieiin, 1S07 ( = Twchns coHcliijluiphonis Hoi-n, ITSO).

Xeru)phom cardHi'UTii new species
(Figures S-U)

Diajjnosis: Moderate size, moderateK tall spiic; base
witliiiut spiral sculpture: dorsal surtace with weakly op-
isthocliuc tci spiral irri"j;ul,u' lirai'.

Description: Diameter up to ofi luiu. Spire au^le 70-
yo""; whoils and spire Hat-sided to \er\ slightlv coii\e\.
Protocouch poorK preser\'ed. Base Hat lo sli'^hllx con-
cave; neither spiral nor suhspiral sculplure i lines,
threads, rugae) evident; colahral giowtli lines ;iu(l ridges
present. No unihilii'us in :idiills: none ixidcnl in ju\('-
niles; thick columella. Dorsal suilace wiHi wvaV prosoe-
line growth lines and coarse, w;i\\ lir;ic ihal arc weakly
opisthocline to irregularK' spiral. No prosocliue nor op-
istliocline axial cords. Cemented objects e\enly spaced.

about se\en pei' w lioil. increasing in size on later whorls;
less than oO*^-? ol shell co\'ercd. .Ajiertnre unknown.

Type Material: iiololxpe S\ll' :523()39 (figures 3-5),
height 17 nun, dianielcr .»(i niui; o parat\pes; SMF
.323040. heighl Id uini. diameter :5(l mm. S.\1F 323041
(figures fi-S). height l.3.."i mm. diametei- 22. .^ nun,
MUS.M 1N\' 1 ifignres Oil), height 21 mm. diameter
.32. .5 umi. MUSM INN' 2. height I.t nun. diameter 2fi.5
nnn, .ML'SM IN\ .). heiii;ht |.> nun. diameter 21.5 nun.

Tyjie L<»ealit\: D\ (S.5I-S. northwest of l^)ma (.'uesta

Chilcatax. about 1 km north of Comotrana-C:arluias road,
about 5 km east Pla\a (,'arhuas. in ridge-lorming sand-
stone bed, 120.5 m in measured section. 14°l I'Ofi" S.
7fi°0S'I7" W iPunta Ciranik^ 1:100.000 (luadrausilei.

El-Miioloj^y : Named alter ('anliln. the bi\al\c most
often cemented to tliis sjiecies. and gc;y;. the Latin loot
signif\in<4 'to beai" or carrv.

Occurrence: Otuuia l''ormatiou. Oligoccne. between
Paracas and Bahi'a de la Independencia. I'cru. I'ossibiy
(rom the uppermost Foc'cue.

Discussi()n: Specimens ol \tiHi}>hora canliti^erfi new
species dilter in several respects from those of X. roti-

Paw 74

THE NAUTILUS. \ ol. 1 16. No. 3

Figures 3-12. Tfrtian XiiKipluini. 3-11. Xciiophora canlitifit'iu ni-w .spciir.s. ;}-.5. I l(il()l\])r. .S.\ll' .32:)(1.)U. li(ii;ht IT iiiui.
ciiameter 36 niin. 6-8. Parahpe SMF 323041. lK•i^ht 13.5 mm, diameter 22.5 mm. 9-11. Parahpe .Ml SM IW 1. htiglit 21 mm,
diameter 32.5 mm. 12. Xenophora pmtlmac ne«- speeies. Parat\pe S(;().l'l 5991. (lianictcr 154 mm.

S. \ Nielsen and T. J. DeNries. 200:;

Pa<fe 75

clujliophora (Bom. 1780). a species with a fossil record
that may e.xtend back to die Eocene and wiiich is pres-
ently found off the coast of bodi eastern and western
North and Central .\merica (Ponder. 19S3!. The latter
species is characterized by rugose cords on tiie dorsal
surface; wa\y lirae that are distincth' opisthocline; base
with weak spiral sculpture: an umbilicus in ju\ enile spec-
imens: and a thin columella and umbilical callous in
adults. Specimens of .V. carditi^cro, in contrast. ha\e nei-
ther rugose spiral cords, stronglv opisthocline lirae. nor
spiral sculpture on the base, and ha\e a thick rather than
thin columella.

Xenophora carditigera most closeK' resembles .\'. ficinin-
<l) Beu. 1977. an earK- to middle .Miocene species from
New- Zealand diat may be part of a lineage diat includes
X prognata (Finlay, 1926) (late Eocene to middle Mio-
cene) and .\'. ncozehinico Suter. 1908 ie;ul\ Pliocene to
Recent). .Specimens oi' X. fleniiiigi are iilso lUoderateK' tall
and straight-sided, lack an umbilicus. ha\"e weakl\- devel-
oped to obsolete spiral sculpture on the base, and are onl\-
partialK' covered b\- cemented debris diat increases in size
towards the apeitm-e. The princip;il difference between X.
carditigera luid .V. flcmingi is diat dorsal lirae on the latter
are distincd\' opistliocline.

Xenophora pnulinac new species
(Figures 12-16)

Diagnosis: Shell large. umbiBcus closed, whorl outline
concave. Subsutural ramp reaching halR\a\ onto previ-
ous whorl. Dorsal surface between cemented objects
with coarse wa\T hrae and prosocHne axial growth lines.

Description: Shell large (liolot\pe diameter 132 mm.
height 62 mm '. with depressed spire (angle about 105°),
peripheral flange unknown. Protoconch unknown. No
umbilicus in adults (no juveniles known), whorl outline
concave due to subsutural ramp reaching about halfwav
up onto previous whorl. Dorsal surface between ce-
mented objects with coarse wav-\- lirae as well as prosoc-
line axial growth lines. Base weaklv concave, sculptured
with low, narrow, irregular, collabrid growth lirae. .At-
tached camouflaging objects seem to include either
high-spired gastropods or concave-side-up bivalves lup
to 50 mm wide). Basal apertural lip regularlv and shal-
lowiv cursed, .\perture unknown.

T\pe .Material: Holotvpe SMF 323042 (figures 13-
14), height 62 mm, diameter 132 mm. Punta Perro;
paratopes SCO. PI 5991 (figure 12), diameter 154 mm.
Punta Perro. SGO.Pl 5992 (figures 1.5-16), spire frag-
ment, height 16 nun. Matanzas.

Type Locality: Intertidal platform at Punta Perro,
central Chile.

EtyTnolog\: Named after Paulina S. N'asquez IlliUU-s.
friend and colleague, who found part of the tvpe materi;il.

Occurrence: Navidad Formation. Tortonian. Navidad
area, centred Chile.

Discussion: Xenophora patdinae new .species differs
from most other species by its large size. The onlv sim-
ilarly large species is the Eocene to Miocene New Zea-
land Xenophora prognata ( Finlav. 1926) isee Beu. 1977)
from which it differs in having concave rather than con-
vex whorls, formed by a sidjsutural ramp reaching about
halfxvav up onto previous whorl.

Previouslv. Tavera il979) stated that his Trochita gi-
gantea. also from the Navidad Area, might prove to be
a Xenophora. However, Tavera (1979) provided no di-
agnosis to separate his species from other taxa, and he
did not figure the specimen. Conse<juentlv the name is
considered imavailable under ICZN Article 13.1.1.

Genus Stelkiria Schmidt. 1832

Tvpe species: Trochus so/c/n.s i>iime. 1764.

Stellaria hriegcrl)artholdi new species
(Figures 17-20)

Diagnosis: Spire short, with narrow peripheral flange
divided into prominent, blunt digitations. No umbilicus.
Base lightlv convex, with distinct collabral growth lines.

Description: Short spire, periphen divided into about
10 prominent blunt digitations. No umbilicus. Whorl
outhne and sculpture unknown, because original shell is
dissolved except for peripherv and digitations, but out-
line appears to be weaklv convex. Base shshtlv convex,
with distinct collabral growth lines.

Holotype: SGO.Pl 5993 (figures 17-20). height 24
mm, diameter with attachments 60 mm. spire angle 9.5°.

T\pe Locality': Northeast of Plava Millongue. Penin-
sula .franco, south-central Chile.

Etvmology: Named after Rolf Kriegerbarthold. who
did the difficult preparation of this specimen.

Occurrence: Northeast of Punta Millongue. Penin-
sula .\rauco. south-central Chile.

Discussion: Stellaria kriegerbartholdi differs from all
other species of this genus in having a clo.sed umbilicus.
The tvpe species, S. solans, has tubular spines, a feature
that cannot be observed in S. kriegerbartholdi. S. krie-
gerhariholdi most resembles the Oligocene to Pliocene
S. testigera (Bronn. 1831). with its two Recent subspe-
cies, in having a stronglv digitate peripheral rim. -S. tes-
tigera lived in the Mediterranean and spread to Atlantic
.Africa and the Gulf of .Aden. .As suggested by Ponder
(1983). -X testigera could have evolved from the Eocene
S. conica (Dall. 1892' from Mississippi, which also might
be regarded as ancestral to .S. kriegcHjariholdi. S. conica
has the umbilicus edinost obscured bv the parietal callus
.Mc.Neil and Dockerv. 1984). The completely closed
umbilicus would also justilv inclusion of this species in
Xenophora. suggc>sting that this species or species line
lost its camouflaging habit and evolved from a difTerent
ancestor than Stellaria s.s. However, this view is not fol-
lowed here.

Pasie 76

THE NAUTILUS. \nl. llfi. No. 3

Figures I3-l(i. Xnwphora pauluwr ur.v spvirs. i:?-14. llolulxp... SMI' o23()l2. lu-,^1,1 h2 nnn. clianu.l.-i 1:52 un
Paratvpc- S'- .> M'l 3i)92. spire IVauiiu'iit, lici^^lil Hi iiiin.

1 5- Hi.

Remarks: '\\,< di.isiii' .I Ibi- Stclltiriti 'j;i\fii In I'diuKt
(19S3) lias to III- cnicndcd to indnclc spct-ics witli a
closed iiinhiliciis like S. kiiciicrhdrllioldi. Ponder (19S3)
noted that die placeiiieiil ot S'. icsti<iii'ra "is iml eiini-
pieteK satislacton". a eoiniiiciit wliicli is excii nunc ap-

prnpnalr Inr S, kiii-^nhinlliohli ll does nut sc.-m jus-
tilird to creel a new '^einis lor tins spet'ies heeanse a
closed nnihiliens is also preseni ni \, iinpIiKril and. tlieie-
lore. seems lo he an old in\entinn ol tlie lamiK. Cliar-
aelers separatni'j, ,S7, //r;/;f/ Irom Xiiioplii'm are reduced

S. \. Xiclsc

T. 1 i:)f\ries. 2002

Wm-

Finiirt's 17-20.

(iO iiini.

Slillariti kiif^crhiiiilioltli new species, llolofxpe, SCiO.I'l .5SJ!).). lieij^hl 24 iinji. diaiiieter witli .ittaeliiiieiits

to tile preseiiee (il spiiu-s or (limitations, an expandi'tl
peripheral flaiiij;e and a iieaiK sniootli dorsal surlaee.

CONCLUSIONS

The Cenozoic Nenophoridae ul Cilnle and I'eiii elose a
considerable biosreographic gap in the histon ol the
taniiK'. Oligoceiie records of XcitDjtliora are lew but the
genus was alread\- established in Australia (Ponder 19S.'3)
and New Zealand (Ben 1977) in the Eocene. In the Mio-
cene the genus was widespread in the Indo-Paeific {Re-
gion with tliiee species also picsenl ni the CaiiMiean
Sea. Toda\^ onK the t\pe species, X. coiulnilii'phnni. re-
mains in the Aniericas (Ponder 198.3).

Our t^\(l species ol XmopJiDni seem closeK allied w ilh
New Zealand species. Similarities between launas Ironi
New Zealand and Argentina ha\e recentK been indicat-
ed b\- Ben et al. (1997). Our records ol Xciiojiliiira show
that there are even more connections witli Pacific South

.Vnieiiea. a laet whieh is. ol course, not surprisin'4. Moil-
tl'.ius-l-'acilic' alliuities can be expeclcd Irom lurlher
studies ol Pacific South .Amerii'an launas.

Stclliiriti ctiiiicd Irom isocene beds ol .Vlississijipi ma\'
be an ancestor of .S, kric^crlxirlliold't but has an open
umbilicus like modi'in sjiecies. i'onder (198.3) discussed
,S. Icsllis.<'i''i as a possible carK ollshoot while Stcllaria
was still close to \ciioj)liorii. which ma\ also be true lor
.S. krii'<H'rh(irthol(li. llowc\cr. characters ol (lie known
Stcllaria species suggest that the lossil record ol (his ge-
nus is \(:'r\ IragmenlaiA.

ACKNow i.i:i)(:mi;x IS

Klaus Handel is diaiiked for companx in the fii'ld and
discussion on (Jiilean geoloin and bioloi^ol Xciiophora.
|ose II. Peal. .\lau Ben. and an anonxnions reviewer
made \aluablc counnents which heliied to improve the
manuscript. Kieklwork ol SNN was linaucialK supported

Pa«' 7S

THE NAUTILUS. No

Ifi. No. 3

bv the Deutsche Forsdiuiigsgeiiieinsehaft (DFG), grant
Ba 675/25. Fieklv\ork ofTJDV wa.s financed In- liimseli
and in part bv a Fulbright Senior Scholarsliip.

LITERATURE CITED

Biindel. K. 199.3. Caenoga.stropoda during .Mesozuif tiniL'S.
Scripta Geologica, Special Issue 2: "-.56.

Beu. A. G. 1977. New Zealand Cenozoic gastropods of the
genus Xenophora Fischer, 1807. Journal of the Ro\al So-
cietv of New Zealand 7: 229-24 1 .

Beu, A. G., M. Griffin and R A. Maxwell. 1997. Opening of
Drake Piissage gatewa\ and L^ate Miocene to Pleistocene
cooling reflected in Southern (Jccan nioilnsean dispersal:
evidence from New Zealand and .Xri^entina. Tectonoph\s-
ics 281: 83-97.

Darwin, C. 1846. Geological (iliscnations on South ,*\merica.
Smith, Elder & Co., London, 279 pp

DeX'ries, T. J. 1998. Oligocene deposition :uiil ('enozoic se-
quence boundaries in the Pisco Basin (Pern). Journ:il ol
South .American Earth Sciences 11: 217-231.

Dunbar R. B., R. C. Mart^-, R A. Baker 1990. Cenozoic marine
sedimentation in the Sechura and Pisco basins, Peru. Pa-
laeogeographv, Palaeoclimatolog)', Palaeoecologv 77: 235-
26L

Garcia A., F. 1968. Estratigrafia del Terciario de Chile C:entral.
In: Cecioni, G. (ed.): Svmposio sobre el Terciario de
Chile, Zona Central. Andres Bello, Santiago, pp. 25-57.

Herni, D. 1969. Marines Pliozan und Pleistoziin in Nord- und
Mittel-Chile unter besonderer Beriicksichtigung der En-
h\icklung der Mollusken-Faunen. ZitteUana 2, 159 pp.

Ibaraki, M. 1992. Planktonic Foraminifera from the Naxidad
Formation. Chile: their geologic age and paleoceano-
graphic implications: pp. 91-95. In: Ishizaki. K. and T.
Saito (eds.): Centenan of Japanese Micropaleontolog\-.
Terra Scientific Publishini^ Company, Tok\o.

Kreipl, K. and A. Alf 1999. Recent Xenophoridae. Conch-
Books. Hackenheini. 148 pp.

McNeil, F. S, and D. T Dockerv. III. 1984. Lower Oligocene
Castnipoda. Scaphopoda, and Cephalopoda of the Vicks-
burg (iroup in Mississippi. .Mississippi Department of
Natural Resources. Bureau of Geolog\- Bulletin 124. 415
pp.

Philippi, R. A. 1S87. Los losiles terciarios i cuartarios di- (^liile.
FA. Brockhaus, Leipzig. 256 pp.

Ponder W. F. 1983. Xenophoridae of the world. The .Australian
.Museum Memoir 17: 126 pp.

Ta\era, J. 1979. Estratigrafia \' paleontologi'a de la F'ormacion
Navidad. Provincia de Colchagua. Clhilc iLat. .30°50-34°
S). Roletin del Museo de llistoria Natural (diile 36:
1-176.

Tsuchi, R., T Shuto, T Takavama, I. Koizmni, A. Fujiyoshi, .M.
Ibaraki and Martinez-R. Pardo. 1990. Fundamental data
on Cenozoic biostratigraphv of Chile. Supplement. 3.
Punta Perro section north of Navidad. central Chile. Re-
ports of Andean Studies, Special N'olume 3: 70-71.

THE NAUTILUS 116(3):79-S.S, 2002

Page 79

Redescription of the genus and species Hctcrova^iiiiiui
Jimaijana (Lesson, 1830) (Gastropoda: Soleolifera:
Veronicellidae!

Jose \>'illilialclo Thome
Siizelc Rodrigues Gomes
Kosane Soiiza da Silva

Lalidratinii) de Malacologia,
Faculiliuk- de Biociencias. PUCRS
9()69()-9()() Porto \h'2\v
BRAZIL
tlioniej\\(a'pii(.'is.hr

ABSTRACT

Based on the examination of 58 speeiinens deposited in xarioiis
institutions, we redeseribe the genus ll<tcroi:a'^inina Kraus,
1953, considering it inonot\pic and with its range restricted to
Penj. The t\pe species, H linuiiicniii. ( l>esson. 1S30) is also
redescrilied, and a neoUpe de.signated for the species. The
anatonn- ol tlie herniaphioditic region near tlie female genital
pore is emphasized as characteristic ot the genus: the copula-
tion bursa is elongatetl'spheroid. tapering into a distal elonga-
tion tliat tolds and joins tlie bursa duct toward its base, receiv-
ing the junctor duct at the tip of the bursa. The rectum pen-
etrates the integument near the female genital pore i'liiic is
no penial glaud and no accessor\' gland. The aualum\ ol the
anterior male gemtal s\stem cliaracterizes the species: smooth
penis, with short cvlindrical \erge that intumesces slighth in
its medial region, forming expansions on the sidis tli:il mark
the beginning ol the glans; tlie penis is lougei llicn Hie xerge,
and tapers in its distal portion. There is no spatlia. All pertinent
hibliugiapliN is discussed :ind the SMion\iui/.ation of \'«g/;/i/i(/
(hIIuiiii I liilliiKinn. UJ2T. is justified.

IXTHODUCTION

The WronicelHdae are terrestrial sings with worldwide
ilistrihiition, stiii poorK' knowii, and ot uiKcrtain position
witliiii tlie Ca.stropoda (Hoffmann. 1925: ii\nian. UKiT:
Thome, 19SSa, 1993; Tillier et al„ 199fii. Among the
genera iiielndcd in the famiK' (Hotfmami, 1925: i'orcart,
1953; 'flioiii('\ 1975), t^\o sliare the lack ol :i penial
gland, a stiiietme foinied h\ a papilhi ;ui(l a varied iinni-
her ol tubules, wliu-ji lies alongside the penis and lias
swstt'inatii' importance, altliongli its tmietion is vet not
understooil.

Ol the hvo genera ol the \ eionieellidae lacking the
penial gland, one occurs in ,\friea. \'(i<^iniiHi Simrolh.
1S97, and the otliei', Hctcwva^imtid. is I'estrieled to
Peru. Sinn-otli characterized Vfi'jjiiind simpK 1)\ the ab-
sence of the penial gland, a,s opposed to the genus Va-

"iiuild l^ertliold, bS2'

:i sv noiiv 111 ()

\ aiiiiiithts Fernssac,

1S21). vvliicli exliibils (bat siriietiire. In |iroposing the
genus, Simidtli !lS!)7' based it on the e\ainiiialion of
two specimens without speeilie names. Oiilv later, thai
author iSimrofh. I9l.it, examining the same two speci-
mens, di'scribed iheiii as V counulli and \! loffu'nsis,
both originating Iroiii the same West .Mrican loealitv. I?iit
tliev differ In the presence of glandular jirotiibeiances
on one side ol the |)eiiis sheath in \ coiiniilti. ami bv a
smooth sheath in \' /()g<ie)is7s

In his disciissioii of the iiomenciatiire of the \ eroiii-
cellidae. Baker (1925) designatecl \'aiiiiiiiia rcnnuUi as
the t\pe species of Vd'^inhui. liollmami ( 1925) accepted
\' roiinnlti and sv nonv mi/ed W lofiocusis. maintaining
that the character used bv Siinroth (1913) would be in-
sufficient to diflerentiate them, in that he bad onlv fvvo
spi'cimens, both collected in the same localitx. Ilotf-
maiin (1925) included the genus in his then new siih-
tamlK .Meisenheimeiiinae, with the following diagnosis:
"dark notum, vvcaklv pointed; genital jiore almost at inid-
lengtli ol the bvponolum: with (he anterior lobe of the
digestive gland behind the anterior loop of the intestine;
with the pedal iiei-vcs together for a short stretch, par-
allel and then diverging: with a needle-shaped penis and
;i basal ringed tbiekeiiing: without penial gland".

I lolliiKiiiu d927) desi-iibed Vaiiiiiiiiti odiuwri based
on a single specimen of unknown origin, but stating that
it was certainlv from .Mriea, despite the lact that W/g/-
nina. known oiiK from .\liica, vv:is onlv recorded once.
Without discussing the smioiimiis he had proposed in
1925, he differentiated \'. (Hlliiicri from V. ioffjcnsis by
the shape of tlie penis (also different from the gemis-
levcl inoiphologx- lie had proposed) and respective
sheath ami asi^'cts of the copulation bursa and con-
necting ducts, lie adds tli;it in his species the lobe of
the digestixe gland is in front ol the iinterior loop of the
intestine (also dilleri'iit Iroiii the jiroposed gemis-levcl
inoiphologx 1.

Page 80

THE NAUTILUS. \'ol. llfi. \(

[•"ore-art (1953). in his monograpli on African Wroni-
ccllidae, consiclcred Vii<iitiiua as a suhs^cnns of Pscudotc-
roniccUa. stating tiiat it dilfcrs from tfic otficr snfigcncra
inclnck'd in tlu- gcnns 1)\' tfu' characteristics of tfic cop-
nfaton' organs and tfic prcscnci- of canafs and ducts iic-
tween tlu- rectniii and the int'tcr. witiiont discussing, at
neither tlie gen\is nor llie sntigenus fevel, the absence
or presence of tlic pciiial glantf. Forcart (1953) agreed
with the s\'non\ni\ of Hoffmann (1925) and considered
Rscii(li)rcr()iiicclla lA'd^iiiiiui) roiiriidti as a \alid name.
W itfiont (fiscussion an(f witfiont examining specimens,
he afso atfdecf PsciKhiirroiiicflla [\'ap,iiuiia} odhncii
(Hoffmann) to fiis list as from "prol)alil\' Africa". He also
proposed a new species, PseudovcmuiccHa (Vaf^inina}
diipoiii. obsening that this would differ from the others
in the subgenus l)\ the presence ot the penial gland
(which in fact sets the species apart).

Kraus (1953a) proposed the ta.\on IhicroiniiiniiKi as
a subgenus of Vaginina, baseil on the description of \'(i-
ginina (Heterovaginina) penivicnia, from Lomas tie .\to-
congo, near Lima, Peru, and included \' odhiich due to
the similarities of the two species. He excluded V. foii-
radti !)ecause the anterior intestinal loop is anterit)r to
tlie digestive gland, the copulation bursa lias a long duct,
and the penis has a much different shape. Kraus
(1953b), without anv reference to Baker (1925) or to
Hoffmann (1925), but mentioning verbal contact \\itli
Forcart (Basel, Switzerland) and based on the work of
Degner (1934) on tlic analoni\ ol W lugociisis. raises
Heterovaginina to the genus le\el. Kraus stated tliat, af-
ter examining histological sections of a paratvpe of H.
pcnnidiuL he found it had a cloaca and lacked the links
between the rectum and the tbnrtli nreti'r, thereb\' dif-
fering from the African genus that, according to Degner
(1934), had separate openings for the rectum and the
fourth ureter that were connected to each other b\ a
urethral canal and 5 urethral ducts (which were used in
the description \)\ Forcart (195.3)). Later on, Kraus
(1954) s\iion\nii/,ed Heterovaginina jieruviaiui Kraus.
1953, with Vagiindu\ liniai/uinis Lesson. IS.'IO. consid-
ering that, according to peisonal inloinialion from \V.
Wevrauch, "only one species occuis " ni the t\pe loealitx.

Forcart (1957), after examining I'eruxian specimens in
the I'ield Museum of Natural lliston (now re-examined
by us), found them to represeni \a'j,iiiiii)i odliiicri. Iml
concluded that this species should lie included m the
genus Heter()va<iinina because il oecurs m southern
Peru. He thus agreed with Kraus (1953a). allliou^li lie
did not slati- so directK.

Thome (1969) redescribed the hololxpc <)\ \agininii
(Heter<n:aginiuti ' peniviana Ki'ans, 1953, depositc'd in
the Seiickenberg .Museum. l'"rankfurt-am-Main, williout
discussing the .s\iion\inv. The copulation buisa is. ae
cording to Thome : 1969), sac-like and sessile in \: i // '
peniviana (which now we know to be present oiiK in
juvenile specimens L Thome (1975), in a revision of tlie
American g(>nera of \erouieellidae, recogni/ed and re-
describetl (still only on the basis of the juvenile t\|ie
specimen) Heterovaginina. keeping il iiionol\]iic i // j)e-

nn ianai and with otcurrence restricted to Pern, lie did
not mention \ odiineri.

Ill addition. Thome ( 19S4) ri'described the liolot\pe
of //, odiineri. di'positetl in the Natiniiistoriska Kikmu-
seet ol Stockholm. Swetlen. .According to Thome ( 1984),
the cdpulatiou bursa is s|iheroid. seated ox'er a thick cv-
lindrieal uniform duct and has a short, slender junctor
diK't that penetrates the copulation bursa duct at niid-
leiigtli. An inspection of its illustration (Figure 18, p. 44)
and kiiowiedm' that the material was damaged, lead us
to infer that the (.(ipnlation bursa had actually been
pushed (.lown along its duct. Thus, the junctor duct is in
part adlu'iing to the bursa duct and does not penetrate
it. 'I"lu' junctor duct must, certainly, penetrate the tip of
the damaged copulation bursa. The penis is short and
broad, with a short, level/convex verge that continues
toward tlie spheroid glans. This latter is trans\ersall\'
widened, witli a lateral conical protuberance projecting
from the le\el face of the verge. Thome et al. (1999)
reported a preliminaiy illustrated redescription of the
genus, which is now presented herein in an expanded
format.

During the examination of the specmu'us aiut ol tlie
literature to define the species of the Wronicellidae oc-
curring in Peru, several specimens. Iiotli juvenile and
atliilt, were found in the examined collections. These
were positixelv identified as Heterovaginina limaijana
(Lesson. 1830). Based on this material we propose the
validitv ol the genus and the species, with their respec-
tive redescri|itioiis and the designation of a neotvpe.

MATERIALS AND METHODS

Fiftx'-eight specimens from 26 lots were examined, from
the collections of the Museo de Historia Natural. Univ-
ersidad Nacional Mayor de San Marcos (MUS.M), Lima,
Peru; Senckenberg Museum (SMF), Frankfurt-am-
.\laiii, Germany; Field Museum ol Natural Histon-
(F.MNH). Chicago. USA; British Musenni of Natural
lliston iBMNH), London, England; .\luseu de (aeucias
e Tecnologia, Pontiticia Unixersidade C^atolica do Rio
(;;rande do Sul (MCP-PUCRS), Porto Alegre. RS, Brazil;
-National Museum of Natural Histoiy (USNM), Wash-
ington, DC, USA (Thome et al., 1997) and Natnriiisto-
liska liikmnseet (NRS), Stockholm. Sweden (Thome.
10841.

For studv of the internal anatomv. the specimens were
dissected, starting with a posterior to anterior longitu-
diiKil ventral incision, follovvin-j, llie iiictliodologv ol 'l1io-
nie and I ,i ipcs (19731 ;ind siil)si'(|iient iinprov emeiits.
Tlic iiiaiidililc ( = j;ivv. ol sonic .mlliorsl and radula of
two specimens lidm lots \1CP 6594 and MUS.M 251.'Vb
were e\li;icled inidci the slereomicroscope and later ;in-
;ilv/ed ;ind iiie;isiired under the sc;iiiiimi,i elcctiiiii mi-
croscope. The ;iv('rages ol length, width, and disl.inci'
betxveen the region of greatest width :ind the anlerior
lip of Ivvc'jitv rac'hidiaii teeth ;ind twentv lateral teeth
vveic c;ilciil;ited. I'lic anatomic, generic :iiid spi-cifit'.

]. w. Tl 11)11 K- ,i III. -loo-:

Vim- SI

cliaiai-tt'ristifs are einpliasizrd, Inllow lipj, I loll niaiiii
(1925) and TlioiiK' (1975. 19SSa. I9SS1), I99.)i.

SVSTEMATICS

Ck'iius Hctcntiaoinina Kraus. 1953
Hctvrovd'^itibui Kraus. 1953a: fi.3-(S5.

Type Species: VdiibiiiKi 1 1 hliini ii^j^iniiui ' jirnn'uiud
Krau.s. 1953a: 63-65 (= Urlciiu ii<j^iitiiiii liiuiiiiuita (Les-
son. 1S3())).

Diagncsis: The reetiiiii peiietiates (lie iiite'j;iNiieiil
near the female t^enital poi'e. The tliiii-tissiied edpiilatioii
bursa is spheroid, taperiinj; in a distal leiiiitlieniuL; that
lieuds and joins the bursa tluct toward its base, reeeivint;
the jiiuetor duet at the bursa tip. The biirs:i dnet. nl
iiioi'e rit^id tissue, is exlindrieal and e\treiiiel\ short in
the juxeuile aninial. but lengthens in the adult, bet-oin-
in<r an eiontiated/spheruid bursa. \o aeeessoi-x inland and
no penial ^laml are to be found.

Distrihiilion: Kestrieted to I'eiu. .South .\iiieriea.

Hiii-niniiiiitiiiii liiiKii/diiii (Lesson, 1830)

\'irj^iiiiiliis liiiiiiiiiiiiii\ LcsMiii. 1S30: 302-303. 471 pi 1 I, fi<^s.

I I \
Vifnimlii liincdciiui Deslia\es in Ferussae and Deshases. 1S19/

1S."j1: V)(S'-9fr. pi. .SE. fig. 11 (exact copv of Lesson. 1S30).
\'ii'j,iiiiiiii ndhiirh Hofhnann. 1927: 2(i. fit^s. II II 'I'liciine.

I9S4: 34-3fi. figs. 14-19.
Psi-itdovi niidiilhi (Va0inn(i' (Hllnuir l-'oitart. 1953: (iO.
Vii'^iiiiui I Ihtirdia^niiKii jicnuKdui Kraus. 1953a: ^-(iS:

Tlioine. 1969: 357. figs. 29-30. 4S
\'ai2iniiia \HcterdV(i0iiiiui < oilhitcii: Kraiis, 1953a: fi3-fi5.
Hetcrovii0iiiii/i liiiKii/aiia: Kraiis. 1954: S2-S3.
Hctcnivdi^iidiiii dilhiirri: Forcart. 1957: 9(S.

Diagnosis: Penis smooth, short exliiidrieal \erge that
intinnesees a little bexoiul the middle ol its length, lonie
ing expansions on the sidt's that mark the beginning ol
the glaiis. tdaiis longer than th(^ \-erge. tapering in its
distal portion; generalK' euncd. There is no spalha.

Redeseription: Extcnuil Aiuiliddi/ A large pmiioii
ol the I'xainineil material is externalK xen diseoloicd.
However, the color pattern on tiie iioliini sliowed ihe
tollowing general characteristics: specimens with light
chestnut colored notuin with irregnlaiK distributed
black dots and splotches. ue\crtlieless ciearK showing
two longitudinal black lines that deliinit a lo/enge-
shapeil mid-region. Between these two lines a fine light
streak occurs centialK in the uotiini. starting :i|iail Ikhm
the anterior pt'riiiotiiin at about '4 ol llie wa\ down the
lull length of the animal and cont inning to I hi' n'ar witii-
out reaching the posterior perinotiim. (.Veragi's ol the
external dimensions, in 20 specimens (measurements in
mm): notum length = 29.20, width = 13.90. height =
7.14; width of the sole = 4.41; width of the right h\-
pouotimi = 4.14, distance from the leiiKile geiiit;il pore
fon\artl = 14.50, and behind = 12.90: distance Iroiii the
female genital pore to the pedal lurrow = 0.73.)

Inlciddl Addloidij: Sali\ar\ gland well de\i'loped.
loniii'd b\ large and prominent acini, some (jiiile loose,
others ((Jiiipaet. .Anterior iiitestiii;il loop. c-o\cred b\ the
anterior lobe ol the digestise gland.

The two nerxe pairs, pedal ;iiid palial. originate \vy\
close to the respectixc ganglia ol the periesopiiageal ring
and stretch through the ceii(i;il ca\it\ in two sets, right
and left, which se|i;iiate from e;icli other about a foiirtii
ol the w,i\ liiiiii liieii sl;ir(iiig phice. continuing thus al-
most to the end ol the cenliai bod\ ca\il\. where thi'V
c-oiiie siighlK closer and peneti.ile llie inlegmnent (Fig-
ure 3). The antcaior aortic arler\ meets the nent's near
their beginning I in ;interior direction) (Figure 3). The
posterior pi'dal biiinih ol tin- aortic ;irtei-\ was not
found. .\\er;ige measurements ol the uenes in 20 spc-
cimeiits ill mm: tola! leuglli - 29.9: together = 4.48;
apart = 16.5; maximum sep;iration = 2.72 nun; meeting
ol the aortic artel^ = 1.76 mm from the origin.'

Pedal gland (Figure 4) shoil . Ilalleiied bro.id m ils
proximal portion, lun'rowing in the middle with the pos-
terior extreiiiit\ rounded and somewhat broadened, dor-
sal opening s(.'allo[)ed: longitndinalK three areas are \is-
il)le. ,\ tninsliicent outer one on each side, and a slightlv
roiigli eeiiti'iil one that is higher, narrower ;iiid \ellowish.
.\t the distal extreiiiit\ ol this central area is the pene-
tration scar ol till- inferior bi;nicli of the aortic arteiA',
right under the lu'nc giingiia. e\\erage dimension of the
pedal gland in 20 specimens (in mini: length in the ii;it-
ural position = 4.20; distended i dorsal smiace> = 4.44;
greatest width = 1.90. (

The thin-tissued copnl.ilion bursa .— spernuithi'ca or
spermatoKtic gland, ol some aiilhorsl is spheroid, ta-
pering in ;i distill lengtliening that folds and joins the
bursa duct tow;ird its base, receiving the junctor duct ( =
canalis junctor ol some authors i at (he bnrs;i ti]). The
bursa duct, ol more rigid tissue, is e\lindrical and ex-
treineK short in the jnxcnile .ininial. but lengthens in
the iiilnlt. becoming an elongated s|)lieroi(l bursa. The
imietoi duel is lliiek and siimons. luu'rowing siightiv in
its Hnal portion, when, alter entwining part ol the base
of the burs;i duct, it j)euetrates into the nairowest part
of the copukition bursa itself The rectum penetrates tile
integmnent near the leinale genital pore (Figures 1. 2).
No accessor gland : = accesson bursa, ol some authors)
wiis found.

Penis smooth, with a short, evlindrieal \crge. swelling
;i bit bcMiiid the middle ol its lull length, forming on
the sides two ex|iansions tluit iiuirk tlie beginning ol tlie
glaus. (dans longi'r than the \erge, becoming slender in
its distal portion; generalK cuned. i^etractor muscli' of
the penis \er\ \ariable in lengtii (Figures 5-16t. '{"here
is no sjKitha. (.\\erage dimensions ol the penis in 20
specimens tin mm); total length = 2.26; greatest diiun-
eter of the glaus = 1.45; \erge length = 1.02; \erge
diameter = 1 .07. )

The sheath surrounding the penis i I'"igure 17' has two
distinct regions. clearK' differentiated b\- the well-de-
fined presenci' of two different t\pes of tissue. The prox-
imal portion of tlie sheatli. which surrounds tlie penis.

Paw 82

THE NAUTILUS, \"ol. 116, No. 3

prt

dpp del teg

vao

Figuivs 1-4. ilctrrornoiuinii liiiuii/aim. 1, 2. I'.ul (il llic postrnor <j;cnilal <.p,;uns anil nl llic
..Itlu' pedal ncr\X's .M(:ra594). 4. IVdal Klaiul (MC:i>6594). al.c: opcninsl; aiir: iirnv m.U: art
bursa; dhc; copulation hiu'sa dnct; del: junctor duct; ddni: middle ileferens; dpd: distal poste
deferens; glp: part of llie pedal iiland;' nnp: retractor niiisele ol tlie penis; npp: pair ol pe(
oviduct; pi1: prostate; rec: central rei;i()n; rep: perieardi;il re',ii()n: ret: part ol the reetuin: te>i:
branch oi tiie aortic arteiT.

leehini i \1( •
: anterior aort
rior delerens;
hil and palial
intei'unieiil;

(i594i. 3. .\
ic arten; hoc
dpp: proxiin
iienes; (i\ i:
\;io: part ol

r;nit;enieiit
copulation
al posterior
part of the
the inferior

W. Tlioiiie ct al, 200

Page 83

2 mm

2mm

Figures 5-16. llrtcnn H'^iiiiiui hiiuiiiniiii 5-13. 'Hiicc (lilTcriMit \if\vs ol' the penis, in -3 adult specimens ( respecti\el\-: MUSM
2513/h iFii^ines 5-7). MU.SM 145S iFii;ures S-l()!. KMNH 594SS/a (Figures 11-1.3)1. 14-16. Three difCerent \ie\vs of' the penis
in one juvenile specimen (FMNH 594S6/1)). dda: antcnor poiliou of the deferens duet: glp: ghms: nnp: retractor nniscle of the
penis; soq; \erge.

Pa^e 84

THE NAUTILUS. \"c)l. 116, No. 3

2mm

Hrlrn,va"i,m„l li„m,,an„ 17. Shcall, that r,n,-lops llir p<-,ns. „pr,u.l Inn.^.t.ulin.dK '^U^^if^^J^/bl/^- ^'"7
sheath sliowinil the shaip l<,nuitucl,H.Ll folds ul thr iMt.rnal tis.su.^ (MUSM 2ol;Vh). 19. Shea h.

Figures 17-20

of the ilistal i)oi'tli>n (il the penis siieaiii, sinnviiin ujc mi,u|j iw.i-i,.ni....i. ,..,,. i i i l l

opened lonf^itndinallv. sliowing the presenee of a sali.Mit i;l..hnlar lonnatuH, nt du- intrrn.il tissue, similar t<, that desenhed by
Hoffmann (1927- lor //. odhm-ri as a crland (FMNH 594S(rl.'. 20. He-un. ..I (hr sah. „t .^Inl.nla, lonnat,,.,,. ^n-atlN rnlaru.'.t. also
,shoNvine details ol the tissue lolds (FMNII .59486/1)). l.ai: shr.itli: d.)l): lolds; i;ll); saheiit i^lohnlar tonuatu.ii.

|. W. ThoiiK- (■/ ai. 2002

Page 85

is e\trt'iiiel\' tliin. often allowiii'j; \isiializati(iii of tin- pv-
iiis b\ transparent'. The distal portion. startin<j; Ironi a
si<4nificant constriction ri<j;lit alter tlie tip of tlie penis,
which is swollen, has numerons lon<jitndinal lokls inter-
nally and a porous and velkjwish (glandular?) aspect.
These folds nia\' lie more or less tight, neatK' arranged,
and rectilinear lespecialK in ju\enile specimens) or in
disarras' (Figure IS), Exen in juvenile specimens where
the male region is little developed, the minisculc folds
(Figure 20) can be seen inside the slieath. The penial
gland is absent.

The mandible is formed bv an average of 2.'3 laths ( =
ribs, of some authors), verv visible and somewhat sepa-
rated one from the other. The most central laths are a
bit taller than the others. \o transverse grooves were
noted, perhaps due to the obvious wear. The average
length and greatest width of the mandible, in the same
lots utilized for extraction of tlie radulae. were 2.1 mm
and 0.5 imn respectivelv (Figure 21).

The radula has a great number of lateral teeth, dis-
tributed rectilinearlv in columns parallel to the single,
central line of rachidian teeth, which are alwavs smaller.
An average of 10.3 coUnnns and 120 transverse rows
were counted, resulting in a calculation of 12.360 teeth
per radula. The average dimensions of the radula were
5.85 mm length bv 3.41 mm at the greatest width. The
average dimensions f)f 20 lateral teeth were: length 56.2
[xm, width 24,5 ixm, distances from the greatest width
to tlie tip 29.43 |xm. The average dimensions of 20 rach-
idian teeth were: length 32. S |i.m, width 12.8 |i.m, dis-
tance from the greatest width to the tip 17.0 p.m. (Fig-

ures

oo_o

■23).

NeotApe (herein designated): Ml S.\l 4001, from
tvpe localitv, 24 |an. 2002, Icii. Mina Ivanii'rc/ and L'lises
Zamora.

^^Jie Localitv: Peru, Dpto. fJma. Oerro San Cristo-
bal, ( 12-01 '4.5', f \, 77'00'52.6" W), 215 m aliove sea
level.

Other Material Examined: .\1LS.\1 4(iOl neotvpe),
Peru, Dpto. Lima, Cerro San Cristobal i 12 01 '45.1" N,
77°00'52,6" W), 215 m above sea level, 24 Jan. 2002,
leg. Rina Ranu'rez and Ulises Zamora; MUSM 4000, 2
specimens, same data as the neot\pe: MUSM 14.58, 1
specimen, Peni, Dpto. Lima, Bcjsque Zarate i ir54' X,
76°29' W), 3000 m above sea level, 24 Mar. 1978. leg.
X'alencia and Franke: MUS.V! 1460a, 1 specimen, Peru,
Dpto. Lima, Bosque Zarate (ir54' N, 76°29' W)
(Chourritos II). 2880 m above ,sea level. 01 Mar,1980,
leg, X'alencia and Franke; MUS.M 1660, 4 specimens,
Peru, Dpto, Lima, Lomas ck- .\tf)Congo (Steinloma)
(12°13' N. 76°54' \V), 28 Sep. 1951,' leg. Koepcke
(Kp340m); MUSM 18()7a. 3 specimens, Peru, Dpto.
Lima, Surco (11°52' \, 76=28' W), leg. Koepcke
(KplOlOa); .MUSM 2388, 1 specimen. Peru, Dpto,
Lima, Surco (]r52' X, 76''28' W"), 2100 m above sea
level. 17 Aug, 1965; MUSM 2389, 1 specimen. Peni,
Dpto. Lima. Distrito de San Borja, (Rio Surco), (I2°06'

X, 77-01' W). 08 Juu, 1986, leg. A. Rossi: MUSM 2513.
3 specimens. Pern. Lima, Ihiachipa (12()0' .X, 76°56'
W); .MUSM 3251, 2 specimens. Pern. Dpto, Lima, Can-
ta (Culluhuav) (iri9' X, 76°33' W"), 3550 m above sea
level, 24 Mar. 1989, leg, H. Sisniegas: SMF 108548 (ho-
lotvpe of Wifiiniiui {Hi'lcrova^nina) peruviana). Peni.
Dpto. Lima, Lomas (k) .^tocongo (i2°13' X. 7(-)°54' \V),
2S Oct, 1951. \eo. Koepcke: S.MF 32.3293/1, 1 specimen,
cv-NUSM 40061, same data as holot\pe; SMF 140062/
7, 7 specimens. Peru, Dpto. Lima, Cerro San (Cristobal.
leg. W. Wevranch; F.M.XII 59486, 8 specimens. Pern,
Dpto. Lima, April 1948, leg. W. Wevranch; F,MXH

59487, 6 specimens, Peru, Dpto, Lima, Lomas do .\to-
congo (12"13' X, 76°54' W). leg. \V. Wevrancli; FMXH,

59488, 3 specimens, Peni, Oconeque, Puno, 4 Oct. 1941.
leg. C. C. Sanborn; FMXH 300744, 1 specimen, (ex-
MUS.M 4000), same data of the neotxpe: BMXH
20020126, 1 specimens. (e\-MUSM 4000),' same data of
the neotvpe; MCP 6501, 1 specimen, Peni, Dpto. Lima,
Bosque Zarate. ill°54" X, 76°29" W"). .3000 m above sea
level. 24 Mar. 1978. leg. \alencia and Franke: MCP
6502, 3 specimens, Peru, Dpto. Lima, Surco ( 11°52' X,
76°28' \\'), 2100 m above sea level, 17 Aug, 1965; .MCP
6504, 1 specimen, Peru. Dpto, Lima, ("anta iCullnhnav)
(iri9' X, 76°33' W"), 3550 m above sea level, 24 .Mar.
1989, leg, H. Sisniegas; MCP 6593. 1 specimen, Peni.
Lima. Huachipa (12^00' X. 76°56' W); MCP 6594. 1
specimen. Pern, Dpto, Lima, Apr. 1948. leg. W. Wev-
ranch; MCP 7988. 2 specimens. (ex-MUSM 4000), sanie
data of the neotvpe: USXM 574941, 1 specimen, Peni:
XRS 949(1052! (holotvpe of Vaginiiia ixlhncril

DISCUSSlOX

Vdilinitliis liiiuii/aitiis was brieflv described bv Lesson
I 1830), who used a few moiphological featiiri's in his
description. Lesson also offered a color plate, probably
prepared in Peni, including dorsal and ventral views of
the animal. This specimen was not deposited at the Paris
Mnseiim, and could not be found, (k'spite our efforts,
in anv other major ICuropeaii .Museum. It is vim-% likeK-
that Berard lost the spec imen after the artwork was com-
pleted. We conclude that there vv;ls never a holotvpe.
The description and figures were used bv Deshayes
( 1830-18.32) in Fenissac :ind Desliaves, with no further
comments.

.\lth()ngh the holotvpe Vaginirw (Hetcrocagiiiina) pe-
nniaitfi Krans, 1953, is an immature specimen, the svn-
onvmvzation of this latter species by Kraus 1 1954) based
on information from W. Wevranch is confirmed alter its
examination.

Recent collections ;ind observations of live specimens
made bv our collaborator Rina Rami're/ confinn that
there is onlv (me species at the tvpe loc-alitv and its vi-
cinities.

Therefore, we herein propose the designation of
a neotype for the species (see above), deposited at the
Miiseo (le Historia Xatnral. Univcrsidad Xacional Mavor
de San .Marcos (MUSM). Lima. Peni, MUSM 4001, and

Page S6

THE NAUTILUS. \ ol. 1 Ifi. No. 3

■'■^^'*^^->V

I iguii- 21.

(.> i„ i,.i .., .V ,,1 ,, iM.Muiii.ir Mcr :')i.i 1

Figures 22-23. Uctciiniiffnina liiiifii/aiKi: Mnlillr region ol tljc ladiihi, 22. sh^lilK .ind 2:5. '.;ri'a(l\ ciil.ii'.^id .\l(d' 251
lateral lootli; (In racliidian toolli k-ciihal).

:Vl,i. d'

W. TIioiik' ,1 a!.. 2002

VdMv S7

collecti'd at tlu' nriifiiial tvpc Imvilih dllln' species. ( ^ei-
ro San C;rist()l>al.

Holfiiiann (1927) pnipcised tlu- species \'(iiiiitulii
odhncri, especialK due to tlie presence in tlie iienis
slieath of a lilandular protuberance. Tlioine ( 19S4I in liis
rede.scription ol the holotxpi' oi \' itdhmri. made no
referenci' to that di.stinctix'e protuherance liecanse the
holotA]ie was extensively daniai^ed.

In tlui-e jn\-enile specimens (SWV 10S.54S. MCF
6593. and FMHN 594Sfi/dl. amomj; theni the holot\pe
ol //. jicnnidiKi ( = H. liiiuii/tniiiy an ahiiost cncnlai
protnlieiance on the inner w.ill ol the penis sheath was
tomnl. \ei"\ smiilai' to that described 1)\ iloirmann
(1927) for //. udhiicri as a "'^lantl". Tlie ■^land" de-
serilied tor //. odhiicii. used lo <hstin<i;nish it from //.
liiinii/diiii ("sensn" Kraus). appears to be oiil\ a more
salient, globular lorniation ol the internal tissue ol the
sheath, because its texture is the same as this tissue (not
a "gland"') (Figures 19-20).

Despite the additions antl changes uiaile lien-in. lh<'
dichotonions key tor determining the .\mericau geneia.
proposed bv Thome (1975). contimies to be valid lor
Hcfcroc'dgiitiiia.

a(;k\o\\ledc:ments

W'e are gratelul to William Beltou, (uvat Cacapon, West
\'irginia, U.S.\, lor translating this article into English; to
Jose H. Leal, .Sanibel, Florida. L'S.X. loi' atlditional help
widi the language; to technician .\ugnsto Mattos. of
PUCRS, for the SEM photomicrographs: to Rina Ka-
mirez-Messias, Coordinator of the Department of Mal-
acolog\- and Carcinologv, Museo de Historia Xatinal.
Universidad Nacional Maxor de ,Sau Marcos, biiua.
Peni. for collecting, donating, and lending specimens; to
-Margaret Baki-r anil Martin Pnzdia. Division ol Inver-
tebrates. The Field Museimi of Natural Histon, Chica-
go, Illinois, USA; and to Ronald [anssen. Curator ol the
Mollusea Section, Farsehimgsinstitut und Naturnmseinn
Senckenberg, Frankfnrt-ani-Main, (lernianv tor the loan
and donation of specimens. Work prepared with finan-
cial support from CNP<i, CAPES, FAPEHCS and
PUCRS (Brazil).

LITER.4TURE CITED

Bakci'. II, B. 192.5. Xoineiiclatnn' of Xrionicclliilac f \ a^iiiiili

dae). The Nautilus 39: 13- IS.
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Tliicrreichs, ans dem Fran/.oesisclicii iiiit .\iinicrkiiiigen

und Zusaessen. Weimar. Wmi; Cr II. S priv. L:in(lcs-lH-

(Instrie-Goniptoirs.
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tiLxiden. Helicarioniden, X'agiuuliden. Zt)ologisclie [alir-

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Particuliere des Mollusques Terrestres et Fluviatiles. Gliez

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llviiuui 1. II 1967. The liivcrtrlir.iles: Mollusc;, I. McCravv-
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Il'illiii.inii II 1925, Die \ aiiiuulideii. l'",in Beitrag zur Keniituis
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61( 1-21:1 :17I '

llolliiiaun. 11 1927. Ueher Xai^iMuliden iins deni HeicliSMiu-
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kunde 82: 6.3-65.

Knius. (). 19.531i. Zur ,\iuiU)iiiie uud svsteiicitisclicn Stcllung
\c)ii }liiii(ii ii'j^iuiiKi Xirliiv lili Molluskenkuiule S2:15.3-
154,

Kr;ius, (). 19.54. Wcitiav X'eroniccllickic ;ius Peru. .Arcliiv liir
Molluskenkunde 83: 81-8.3,

Lesson 11 V Zooloirie. 18.30. In: Duperrev. L. I. — Wnage ;m-
tour du uioudc, execute par onlre du Roi, sur la Corvette
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Siuuotli, II. 1897. Ueher vcrscliiedene \;icktscluieckeii. Sit-
zungshericlite der \;ilMrlorschenden Cesidlsclialt zn
Leipzis; 22-2.3: 140-1.5-1.

Sinirotli. II. 1913. Uelier die von llenii I'rol. X'oeltzkovv anf
Madagaskar uud in ()slalrika erheuteten \'aginulideii.
nehst V('nv;uidtciii M;aeri;il von gauz Afrika. In: \beltz-
kovv. A,— Heise in (>sl;ilrika in den Jahren 190.3-1905,
luit. . .- WisseuelKiltliclie lM<j;el)Tiisse. Stutlnart. F.
Schvveizerhiirt'sclie Wrkigshuchluindluui;. Nae<;ele 6c Dr
Sproesser. v. 3, Svst. Aht^. Heft 3. p. 127 216. pi. 1.3-17.

4"lionie. I W 1969, haiieulc Beselueihung neotropisclier \e-
ninici'llid;ie- Tvpen iMolluse;i. (^astropodai. II .Arten aus
der Sauuiiluui; des Seuckenherg- .Museums in Frankfurt a.
M.. Arcliiv hir .Molluskenkimde 99: .33I-.36.3.

TlioiiK'. I \\ 1975. ()s generos da Familia \Vronicellid;ie nas
.Xmcricas ( Molluscii; (iastrojiodaK Ilieringia. serie Zoolo-
gia. 48: .3-.56.

Tliome, J. W. 1984. Wiomcellitkie .Mollusc;!. (;;Lstrop(Khi' paii-
trojiicais: III. Hedeseiivao de 5 especies, com IxLse no px-
ame dos tijxis depositados no "N';iturliistoriska Hiknni-
seet ", de Isstocolmo. Siieei;i. Ilieringia, serie Zoologia. 64:
29-46,

Thome. J, W. 1988;i. luc;ir;icleriz;n,;'io de Vcmiiirrlld sloiiiwi
((.aivier 18171 (Mollusc;i. (iiLStropoda. \'erouic<-llid;ie).
Iheriugia. serie Zoologi;i. 67: 1.3-35.

Thome. I W 19881) Hedescrivao dos tipos de N'eroiiicellidae
iMollusc;i: (;;i.stropo(lal Neotropieais: XI. Especies na
",\cadeinv of .X;itund Sciences ", Philadelphia. Ilieringia,
serie Zoologia, 69: 2-5^39.

I'honie. J. W. 1993. Fsl;ido atii.il d.i sislematic;i dos Wronicel-
lidae (Mollusea. (iastropodal americanos. com comentar-
ios sohre su;i importruicia ec-onomic;i. amhieutal e na sai'i-
de. Bioeieucias 1: 61-75.

ThomO. |. W., S. H. Comes ami K. S. Silva. 1999. Recaracter-
iza^ao do nenero ilrtcnna^iitiiia Kraus, 19-53 (.Mollusea,
Ca,stropod;i. Wionicellidael. In: (a)ngreso Latinoameri-
cano de .Malacolosji;!, 06-10 Septiemhre 1999, Oxjniniho,

Paire SS THE XAITIIA'S. \ol. 1 IR, NO. 3

Cliilc. HfSMim'nes, llnixcrsulad ( ^.itolica Del Nortf. Cx> ington, D.C., U.S.A. (Molhisca: Gastropoda; SoleoliicraV

(|uiiiil)(). pp. lOfi-lOT. Proceedinss of the Biological Socieh' oi Wa.sliinijton 110;

TlioiiuK J. \\'. am! \; L. H. Loprs 197.3. Aida.s pnitica.s de Zoo- 520-.5.36-

losjia I. Dissccavao dc mil luohi.sc'O ga.sti"6])odi' dcsprovido Tillicr, S.. M. .Massilot and .V. TiUiLi. liJIJd l'li\loi;riu4ii- re-
de concha. Iheiiiigia. .serie Diviilgavao (3): 34—45. lationsliips of tlic Puhnonate Ga-stropods from rRNA .se-

Thome, J. W'.. P. II. dos Santos and L. Pedott. 1997. Annotated ijnences. and tempo and age of the StNlonmiatophoran

list of Wronicellidae from the collection.s of the Aeadenn radiation In; Ta\lor, J. D. (ed.) Origin and E\c)lutionan

of Xatnral Sciences ol Philadelphia and tiie National Mn- radiation ol the Molln.sca. O\lord l'ni\(isit\ Press. New

senm of Natural lliston. Smithsonian Institntion, Wash- York, pp. 267-2S4.

THE NAUTILUS 1 lfi(.",):S9-94. 200

Page 89

Aiiatoniy of the dromedaiy jiiniping-slng, Hciiij)hilli(i
dwmcdanus Branson, 1972 (Gastropoda: St)l()nnnat()plio)-a:
Arionidae), with new distributional records

Kristiina ()>aska

Hidliiix EnxiroiiiiHiit.il Hisiarcli lid,

4 ISO Clinton V\mv

X'ietoria, British ( Nilnniliia

CANADA \'SZ (SMI

and

Department of Forest Seicincs

Universit\ nl liritisli (jiliiinlna

Vaneoincr, Britisli C^oliniihia

CANADA V6T 1Z4

kcnaskaCn'jdniicni.eoni

Lylf C'liiclu'sliT

2S()5 (Jreenhriar Uonlexard
Wellington. FL33414 USA

Ileike Reisi-

State Miiseniii ol Natural ilistoiA-
PF3()0 ir,i
D-()2S0(i Coilit/
CKHMANY

\\illiain P. Leonard

223 Foote Street NW
OKnipia. \\A 9S5()2 USA

Jim Haiii^li

2()|S Dn Creek Hoad
Flleiislmp' W \ 9SV)2(i USA

ABSTRACT

rile '^eniis Hiiiijiliilliii I'epicseiited 1 1\ al least 7 spi-rii-s, is a
poiiilv kiiiiwii '4riiiip III sjii'^s I'lidiiiiie Id wcslirii \im1Ii \nier-
iia We niNcsti^atid tlic disl nl iiilii m ,uid liahitats ol // dni-
iiiiddiiiis Branson. 1972. and present a redescription ol tlie
anatoins ol its distal genitalia, nsetl tor speeies identiliiation.
We collected specimens from fi localities on X'aneoiner Island,
British Columliia, Canada, and hom 9 localities in Wasinngton
State. USA. The \'ancon\er Island localities repnsmt a north-
ward range extension loi' the species and first confirmed re-
cords Iroin ("anaiki rlii- species occurred from ncLir sea le\el
to an eliAatioii ol I :i70 in. wilh iiiosi localities alioxe 701) nl.
In all disscctt-tl specimens lat least I per local it\! the penis was
priiximallv broad and distalK taperetl; a \\(ll-ile\ eloped \erge
was present witliin the penis sac. Contran to the original spe-
cies description, the penis had an accesson sac that inserted
distalK near the gonopore, .Although the anatonu of the gen-
italia did not match the original description, the dissection ol
the t\pe material confirmed that onr speiiineiis repnsented //
ilntiiiiildriii.s.

IXTHODUCTION

The genus UcuipJiiUid lilaiul and Hniiie\. 1 S72 eiinsksts
1)1 a giDiip <il arloiiici slugs endeinic In weslein North
America. Slug.s of thi.s genus are cliaiaelen/ed li\ a vis-
cera! ca\ih' ele\ated into a pniiiciiinced. dorsal liiiinp. a
.shell-plate that i.s partialK exposed llirongh a slil in liie
mantle (Pilshn. 1948). and a remarkable anti-|)redatoi-\
hehaxior: the\ txpicalK exhibit "xioleiit writhing and

leaping in response to iiiolesladoii I'ilslirv. I94S; 73S).
Sex'eii species are eiiireniK reeogni/ed // 'J/liidlilostl
Rlaiid and Hiiniev, 1S72: // hnrrhfjoiii I'ilsbrx. 1948:
//, pinillirmiii lii.inson 197") // idiiichis l'ilsbi"\ and
N'anatta. 1897: // ilriiinitltiiiiis liianson, 1972: //. daii-
iiisi X'aiiatta. 191 I: and //, iiiulonri Pilsbn. 1917: Tiir-
geon et. al., I998i. i)iit :iddilli)ii:il. midescriiied species
ma\ exist i kelle\ el al,. I999v I'lie geims has receixed
little attention lioiii ell In a' s\ slenialists or ecoiogists. and
file taxononiie nkitionsliips. disliibnlion. and ecologx ol
all speeies ;iic poorK known.

Hr;nison 11972' described // drdmidiiriiis based on
specimens Ifom the OKnipie Nh)milains. Washington.
His di'seription was based primariK on external chaiae-
leiislics w illi lillli' rerer<'nce lo llie anatonix' of the ilistal
reprodiieli\e s\sleiii. wliieh is ollen extremelv useful or
e\fii essential loi' the idenlilicatioii ol shigs (Keniex and
Cameron, 1979: dompa. 1984: (;onie/,. 20()Ll Onr in-
leicsl ill llns speeies arose alter the diseovcn in 1999 ol
ii large ((iO iiiiii In length; jnmpiiig-shig Ironi N'aneouxcr
Isl.uid. Hnlisli ( ohmibia. (Canada: onK //. '^laiidulosa. a
iiiiieli sni.illc'r speeies. was pre\ionsl\- known Iroin tiie
west coast of liritisli (.'ohunbia i Pilsbrx I94S: Cameron
198fSi, but an earK report of a huge, unidentified //(7»-
phillia loriginalK niisidentified as //. inaUmei) existetl
from \ancou\cr Island (Ihinham. 192fi'. Our attempts
to \erif\ the identitx of the new s])e(,'imen were Irus-
Ir.iled l)\ ,1 lack ol piiblisiied information on the genitalia
of // di(iiii(d<inus. iM'omjiting us to examine the t\pe
mati'rial and to collect additional specinu'us from Wasli-

Pat:.' yo

THE \AUTI1AS, \nl. lift. \<

iiiiitoii State. United States, lor eoinpaiisoiis. Here we
present a description of tlie distal reproductixe anatonu
of//. (Iroiiu'daiiits based on tliesc findings. We also \ivc-
sent new distribntional records tor the species, inclndiiig
a northward range extension and the first confirmed doc-
niiientation of the species from ( 'anada.

MATERIALS AND METHODS

We colleetetl slugs from forestetl areas on Nancouvt'r Is-
land. British (]olmnl)ia. (Canada, and in Washington
State. United States, between 1999 and 2002 (.seeAp-
pendi\ loi' collection localities antl tlatcs). On X'ancouver
Island, the first specimen of H. (Iroiiicihiriiis was found
incidentalK during sunevs for terrestrial gastropods in
the autumn ol 1999. but sunexs in snbseijnent vears
focused specificalK' on potential habitats tor this species
(a total ol 104 forested sites were suncxed on \'ancon\er
Island and 3S sites on the mainland of British Oolnm-
bia). Specimens from Washington wfre colleetetl either
for comparisons with material trom \'ancou\er Island or
during opportunistic suneys in \arions localities. We lo-
cated slugs b\ turning over woocK debris and b\' xisualK
searching other potential microhabitats on the torest
floor. Specimens collected were killed b\ drowning anil
preser\^ed in either 10% ethanol or isopropxl alcohol,
and vouchers were ileposited in the eolk'ction at the
Ho\al British Oolumbia Musemn (RBOMl. Mctoria.
British C^olnuibia. (.'anada.

We dissected at least one specimen from each localit\'.
The dissection ol new material was pertormed b\' LC
(most specimens) and KO under 7-30X magnification
using a dissecting microscope. Drawings of the repro-
ductixe sNstem of selected sjiei'imens were made using
camera lueida attached to a (.lissecting microscope. The
dissection and examination of the holotxpe (USNM
577690; National Museum of Natural Histon, Washing-
ton, DO) and txvo paratxpes (FMNH 173022', Field Mii-
.semn of Natural llistorv. Chicago; DMNH 43029. Del-
aware .Museum of Natural llislon. Wilmington) was
performed by HH.

RESULTS

On Nanconxcr Islaml we located // (Inxncddiiiis al six
localities ranging from ni'ar sea le\cl lo ;iu elevation ol
1060 m :.\j)pen(li\). These loc-alities represent ,i north-
ward range extension for tiie species ( Eignre I 1. I iabilats
consisted ol conilerous forest, dominated !)n western
hemlock [Tsnou liclrrophijlla). western redeetlar {'ihuja
pUcata). and am.ibilis fir (Ahics aniahili.s)- the forest tloor
wa.s txpically composed of thin, compact mcdle littei
Fixe of the six loe;ilities xxere in ri'nmant patches ol Old
groxxth forest (with trees > 250 years old); one loc;ilit\
(record 3 in .\ppendix) was in second growth torest with
some old trees. All sites contained abimdant coarse
woody deliris, inclnding large-diameter logs in atlxanced
.stages of (k-cay. In Washington, habitats of H. drouw-
dariuH consisted of both old-groxx'th and older second-

growth conilerous forests dominated bx Douglas-fir
[Fsvudotsiiiid iiwiizicsii) and xxcstern hemlock; mountain
hemlock (7's7(g« mciicusidtKi^ anil subalpine fir iAhii's
hisiocarpa) xvere present at the highest elexation sites.
Within these habitats, xxe tonnd //. drotiicddriiis under
and xxithin decaxing logs and lallen bark, under rocks,
and. once, actixe on the surface of the litter. Most sites
xxere mesic, but at three high-elexation sites on the east
slope of the Cascade Mountains (records 10, 11. and 13
in Appendix 1) the slugs occurred on drier talus sub-
strates.

.Moiii'iioi.oi.x wii Intkhnai, .AxxToxiv

Extenuillx, all specimens xxere similai' and contormed
xxith the tiescriptioii b\ Branson (1972) (Figure 2). The
pretlominaut color ol the mantle and foot xxas various
shades ol grax xxith cream-colored mottling on the sides;
the sole of the loot xxas pale-xelloxx; orange-xelloxx'. or
cream. Otic inilixidual (Irom loealitx 1 in Appendi.x) xxas
bull broxxn xxith a bright xelloxx sole. The tail-portion ol
the foot xvas laterallv compressed, keeled, and tipped
\xith a horn-like protuberance leaudal horn), as de-
scribed bv Bran.son (Figure 2. insert). The nine \'aneon-
xer Island specimens ranged from 40 to 60 nun in
length; three specimens trom Washington (record 9 in
.Appendix) xx'ere 32-37 unn (all measurements trom
fresh, reliLxed specimens after droxxuing).

Inteniiillv. the oxotestis xxas grax-broxxn antl similar in
color to the digestixe gland, ("lose to tlie entrance ol the
ducts ol the digestixe gland there xxas an intestinal di-
xerticnlnm ol nnknoxxii hmction. The penis xxas proxi-
inallx bnnul and tapering distallx'. and a xxell-developed
xerge x\as present (Figure 3). The xerge filled most of
the broad end ol the penis sac; there xxere no other
intra-penial structures. The penial retractor nmsele in-
serteil at the long and slender epipluillus. The penis had
an iiccesson s;ie. xxliicli eonsist("d of ;i narroxx tube that
broadened pi'oxiuKillx and iiregularix i Figure 3!. The tu-
bular base of the accesson sac inserted toxxards the dis-
tal end ol (he penis ne;u' the gonopore. The bursa I'o-
[iilkitrix (spermatheeal consisted ol a stout tube (h;il
broadened proxiuuilK into a globular sac. more jiio-
nonuced in some iudixidnals than in others. The peins,
Iree oxiduel, anil liiusa eopillatrix joined close to the
gonopore; iheie \\,is no distinct alrinm.

The holotxpe is a lullx iidnll specimen with ;i Lirge
.illinmen ghmd. ;ind its genital aiuitomx is generalK the
s;ime as m oni siiecimens. The distal part ol the penis
is liilhei- u;iii"o\x, ,ind the proxim.il p.irt is wide. There is
a penial ;i|)peudi\ that insi'rts dist;illx near the gonopore.
■file retr;i(lo|- inserts .it the <'piph;illus. Intern;ll penis
stnietures xxere not inxestigated to minimi/.e damage to
llii' lx|)i' Sjieeimeu. Both |)aratxpes are juxenile. ;ukI
their genitalia are in somexxhat miu,itm:il positions,
which furthei' hinders the inxestigation (parts ol the dis-
l.il genitaliii ol speeiiiieii DMNH 43029 are xxithin the
right leutacle. and those ol specimen F.MNH 173022 ;ire
partlx exerted through the genital pore — both probablx

K- ()\aska i1 al. 2002

I'a.'c 91

Figure 1. Distil I iiitidii map lor lotalitx iccdnls ol U(iiii>hilliii (Inmnddniis Solid t-ircles: this si ink i iiiiiiilicis {oncspoiid to those
ill AppiiulK); open circles: piv\iiiiis Iciealltles (Branson, 1972. 1977, 19S()). Arrow points to the t\pe loealitv iHrarisoii. 1972).

artifact.s of killing). Nexertiiele.ss, a penial appendix i,s
pre.seiit at lea.st in specimen DMNC 43029, and the
stnicture of the everted genital complex in s|)('ciiiien
FMNH 17.3022 suggests its presence. \\\ lliicc t\pc
specimens ha\e the intestinal dl\(■l■|i^■llllnll inciiliinicd
above.

DISCUSSION

The geographic tlisti'ihntion ol //. (Ironicilnniis. like thai
of man\ other terrestrial gastropods in western North
.America, is poorK knowii. Pre\ious piiMished loealit^ re-
cords aic iioiii the Cascade Range and ()l\in|)ic I'eiiiii-
snla in western Washington, including the ()l\iiipic and
Mt. liainier National Parks ami the .\lt. Baker and Suo-
(juahnie National Forests (Branson, 1972. 1977, 19S(),i
(Figure 1). In addition, some ol the records Iroiii west-
ern Washington reporteil 1)\ i'ilsliiA (i94Si lor //, inn-
loiui. prior to tlie description ol //. (Iroini-iliiriiis. may
lielong to the latter species. Our records Iroin Nancouxcr

Island represent a nortliuard range extension and first
c-onfirmed records from Canada. On the mainland, both
the iiortliern anil southern extent ol the species' distri-
bution remain poorK known. The norfliermnost main-
lanil n'cord is Iriiiii the Cascade .Mountains. Wasiiington
(SiKcr I'ir Camp. .Mount i5aker National I^'orest: Bran-
son, 19S0). close to tile Canadian border ( Figure 1 ). The
sontherimiost confirmed localit\' is from the (iillord Bin-
chot National i'"orcst (record S in .\|')]iendix). lliiiijiliilliti
thai arc similar in si/e and external appearance to //
(Iroiiiciliirius lune been iomid lartiier south in the (Cas-
cade and C^oast ranges ol northwestern Oregon (Tom
Biirke. pers. conmi. b\ WBI,: Carol i^icklord, pers.
coiiim. b\ Wl'I/i. but iiecause these were not dissected
sjiecics i(k'ntificatiou remains uncertain.

We fonnd H (Iniiiu-tltirius at six localities on X'ancou-
\cr Island. \ prexioiis report (llaiiham, 1926: 143) exists
of tx\-o specimens of a large, black jmnping-slug from

. . the iioriler ol a good-sized lake |most likely Ifolyoak
Lake! on Mt. Brenton. \'ancou\cr Island, at an elevation

Page 92

THE NAUTILUS. \ol. llfi, NO. 3

Figure 2. Specimen ot Hciiiphilliii droinfdarius Irniii Loss
Oreek. N'iiiieonver Island, Britisli Colnmhia. C^anada (l()calit\'2
in Appendix; KBCM ()()1-0()2S(M)1)I ); candal liDin (.1 llic same
animal linseitl. Tlu' Irni'th ol tin- li\c' animal was 55 imii.

Iiitween 3500 and 4000 ft". The slugs were identified as
// iiifiloiu'i In Walter |. Excrdani (of .Seattle. Wasliing-
(i)ni. apparentK nn the hasis ol Ilaiiliani's later descrip-
tion oi their color; no spi-ciinens were a\ailal)le. as their
soft anatomy had disintegrati'd before the slugs coukl be
preserved (Hanham, 1926). Later authors (Pilsbn', 1948;
Kozlolt and \'ance, 1958) questioned the identification
ol this rcioicl. which has remained enigmatic. We con-
firmed the presence of H. clmnwdariiis near Holvoak
Lake I record 4 in .Appendix); most likelw the slugs re-
ported b\ lianliain Ironi tlie same localitx' 75 vears pre-
\iousK were also ot this species.

Our localit\' records trom \'ancnn\ei' Island were Iroiii
older torests at elevations lioni near sea le\cl to lOfiO
m. A\\ three localities in the drier southeastern and
soutli-central parts ol the island were abo\'e TOO ni in
elexation. and the species seemed to be absent from
lower slopes ;ind \alle\' bottoms. In contrast, the three
localities on the wetter, west coast ot the island were
near sea-level, in Washington, pre\ious Records for the
species existed tiom elevations of 23S m to 1436 m

1 mm

Figure 3. HcpnuliictlM- system oi llcini}liilliii (Inmiiilfiiim drawn iroiii 1 spciiniciis Ironi Wliilc Pass. Lewis Co.. Wasliiiiuldii
(HB(,M 0()1-2HB-()()I: locality S in .\ppendixi; genitalia of specimen 1 and lateral \iew of the verge, dissected from tile peiiial sae
of specimen 2 (insert I. .■VC: aecesson- sae of penis: .\(.',: alhnmen tjland: BC: bursa eopniatrix ispermatheea); EP: epiph:illns; III);

hcrmapliroditie (hiet; OT; ovotestis; V penis: I'W: pcnial v\all. ]i(c'led liaef: HI pcni;il retnietor; Sl'0\" — sper xulud: \'; verge;

\'S: Vits ileferens.

K. Ovaskar/ ,;/.. 2002

I'asic 9.)

(Branson. 1972. 19S01. .Ml hut one iifiiiir n'conls Ironi
W'asilington (ivconl 7 in .\ppt'iuli\) were lioiri rclatixcK
higli elexation.s (744-1370 ni). On the east slope of tlie
Cascade Hant^e (records 10-15) the species appeals to
be confinet! to woodeil nioinitaiii peaks amidst drier
grass/shrul) hahitats. Iiiterestiinj;l\. at tiiese sites the slugs
inhabited sparseK' wooded siibalpine tains.

E.xternalK- onr specimens conlormed with tin- descrip-
tion for //, (Inniuilarius (Branson. 1972'. aIthoir.^h oin
measnreineiits ol the length of Iresh specimens Iroiii
\ancon\fr Island i40-tS() mm' were considi'rabK' greater
than the length (24-31 mm I reported In Branson. .\s
the t\\(i juNcnile parat\pes suggest, most ol Bransons
specimens might not ha\t' been hilK grown. .Xlthoii'^h
tlie anatonn' ol the genitalia ol our specimens difleied
from the original description, thest- specimens were
comparabh' with Branson's il972i t\pe material. Tiie
original di'scription referred to the anatomv of the gen-
italia three times: in the idetitilication ke\ (p. 103): "pe-
nis broad, lacking an accesson gland"; in the corrobo-
rati\e di'Scription (p. 105): "the inllated penis does not
bear an accesson sac": anil in the diagnosis (p. 106): "It
[H. clniincddriii.s] differs from //, (laiiiclsi in matters ol
coloration and in possessing the inllatetl penis and in
lacking an accessoiT stinnilator". We presume that the
accesson- organ in question refers to the same stnictnre.
In the ailiilt spei'imens iinestigated b\ iis. onK the prox-
imal portion ol the penis was broati, whereas its base
was long and rather narrow. Also contran' to thesi' state-
ments, ill the specimens we examined, including t\j)v
specimens, a penial accesson sac was alwaxs present.
However, the liolotxpe as well as the- two imestigated
paratxpes hail not been disseited In Branson, so it re-
mains unknown wiiat specimen, and therefore species.
he might ha\e cxaiiiiiK-d lor his anatomical descriptions.

In H. (Iroiiu'claiius. there were no intrapenial acces-
son' stnictures, such as the "stimulator" described lor H.
Iiuiloiwi (Ko/lorr and \aiicc. 195SI. .\ more detailed,
comparative inxi'Stigation ol the xcr^e and other iiitr;i-
penial structures among species ol I Iciiijiliilliii awaits in-
\estigatioii.

Three other larij;e-liodii-d (>.30 mm in Ic-ngth i sjieiies
ol Hciiijihilliii are currently recognized iH. nuilmici. II
caiuclus- and //. diinielsi). A penial accesson sac is ab-
sent in // inalonci (Pilsbn'. 194S; LC". unpublished),
whicli is sxnipatiic with // ilroiindiirins in the OKnipic
Mountains ol Washington. I'AternalK. the tx«) species
can be distinguished In the absence ol a caudal hum
and the presence of a prominent, light-colored, mid-dor-
sal stripe on the tail in H. iiKihinci. Hcmpliillin rniiitliis
from eastern l^ritish Columbia and Washington also
lacks both a caudal horn and a penial accesson sac (Pils-
bn', 1948). Of the described species, H. droiucdarius
most cIoseK resembles //. dtni'uhi^ which occms in
Montana (Pilsbn', 194S). ami our speciniciis ke\c(l out
to this species using Branson's (1972) identification key.
Sx'steinatic relationships within Urinpliillia and among
related genera ha\e not been examined recentK, and

comparative studies ol reproductive analonix and genet-
ics ol Hiiiijiliilliii are in ordia'.

ACKNOW II'DCMIA rs

I he first //, di'oiiicddriiis Irom \aiicouver Island was
lonnd diirinii gastropod snnevs sponsored bv W'evcr-
haeiiser Canada ( N'anaimo Office*. .Subsccjiienl financial
support lor sunevs in British (.'olnmbia came from the
Endangered Sjiccics Hecoven fund. Wildlife Habitat
Canada. :uid Dipartmcnt ol forest Sciences. L'ni\ersit\
ol British (!olimiliia ithrouiih a Forest Bencwal Hrilisli
(^olnmbia grant to |ohn Hichardson!.

We thank Su/annc Beaiichesne. Christian Engelstoft.
Hobert l-'orsvth. Brent lladdawav. Lee.Ami Hancock.
Kellev joigcnseii. 'rum Kogiil. Brad Moon, Bill Null, ami
L.emiart Sopnck lor entliusiastic help with the collection
of speciiiR'HS. Bobert Forsvth provided literature refer-
ences and contacts. Clen Dunsworth and David Lindsav
]ii'o\ided access to forestn lands. KelK Sendall and Phil
Lamiiert allowinl us to use the tacilities at the Boval
British Columbia Museum and acce|)ted our specimens.
We are also grateful to Hobert llershler Timothv Pearce.
and |ii(licii (icilier lor the loan ol tvpe specimiMis. Ira
W illev hclpiil with the iiiKi^inn ol tlie figures,

LITEB.\TUHK ( ITEl)

Bransdii. B. .\ li)72 lU iiiplitllid ilioincdiiriiis. a new ariouiil
slu'4 Ik III! \\asliin>j;t<iii. 'I'lic Xaiililiis S5: 100-106.

Branson. H, .\. I!)77. I'rcsliwatcr ami terrestrial .Mollusca ol
the Olv iii|iK' I'ciiiiiMila. Wasliiiiiildii. 'I'lic N'cli^er U): 319-
330.

Biaiisoii. B, \. I9S0. (.■()llccti()iis ol ijastropiKis Iroiii the C'as-
(■;i(lc Mduiitaiiis ol W'asliiiimon. 'I'lic N'eliiler 23: 171-17(1

(ioiiuv. H I 2()(ll Slnirture aiiilliiuclioiiing 1)1 the reproiluc-
tivc svslciii In B.irkcr <.'■ \1 c'd.!. The Biolos^ol Ter-
restrial M.illiiMs ( \BI ruMi-.liiiiu, Wallinuford.pp. 307-
330

llaiiliani. .\ W 1020. Ih iiiphillui iiuilnini \'aii. The N'.iutihis
39: 143-1 It,

K.llev. H., S. D.mhiii. \. Duncan and T Biirke. 1999. Pield
Guide to Suncv and .Maiiaije Terrestrial Moliusk Spi'cies
from the N'ortlivvest Forest I'laii. Bureau ol Laud Man-
auinicnl, Orenoii Slate Office. 114 pp.

Kiincv M I" and B. .\. D. Caiiierou. 1979. A Fiekl V,\m\c to
land Snails of Britiiiu and Nortli-we.st Europe. William
(;()lliiis and Sons. Ltd.. Foudoii, 2S8 pp.

Kozloff F.. \. and 1 \ancr. I95S. Svsleniatic status oi llcin-
phiU'ui iiiiili'iiri the Nanlilus 72: 42—19.

Bilshi-i. II A, 19IS. I.,ind Mollusca ol North .America i north
of Mexico'. The .Acadeinv ol Natural Sciences ol I'liila-
delpliia. Mononrapli 2: 521 - 1 1 F5. i-xlvii.

4'iiinpa. A S I5)SI I .and snails iStvlommatopliora'. In: Toiiipa.
.\. S.. N. II. X'erdonk and j. .\. M. van den Biggelaar
(eds.). Tlie .VIollusca, 7: Beproduction, .Xcademic Press,
Fondoii ;nid New York. pp. 17-140.

riip.icoii. I) 1) I I' (,)uinn jr F. \ Co.ni. F (.'■ Hochherj;.
W. (;. Iaoiis^ I' M Mikkilsen. B. J. Neves. C. F E. Boper
(;. Boseuherii. B Both. \. Selieltema. F. G. Thompson.
M. X'ecchioue. and j. 1). Williams. 199S. Common and
Scientific Names oi .Xciuatic luvertehrates from the Fnitcs

Page 94

THE NAITIIA S. \ ol. IKS, \o. 3

States and Canada: Mollusks. Secfind edition. American
Fisheries Society Special Publication 26. 526 pp.

APPENDLX

Lxjcalities for Hemphillia (Iroinedarius examined for this

stiid\'.

Vancouver Island. British Columbia. Canada:

1. 5 km northeast of ^^'ild Deer Lake, 9 km soudi-
west of Duncan (logging road LIOOO): elexation
700 m above sea \e\e\ (asD: 4S°41' N. 123°46' W:
S October 1999 (collected b\ K. 0\aska. L. So-
piick. ;uid S. Beauchesne): RBCM 001-00279-
001.

2. Loss Creek, at Www 14 southeast of Port Ren-
frew; 48°28' X. 124H.5' \\'; elevation 49 m asl: 14
April 2000: (collected b\ R. Fors\th and K. 0\as-
ka): RBCM 000-00145^)01: lo'june 2001 (col-
lected bv K. 0\aska and C. Engelstoft); RBCM
001-(X)2'S0-001 (Figure 2).

3. Banifiekl West: 4S=.5()' X, 125°8' W: eIe\adon <
20 m asl; 23 JuK' 2000 (collected b>- K. 0\;xska:
maintained in capti\ih until 19 December 2000):
RBCM 001-00281-001.

4. \'anc-ouver Island, Mt. Brenton (south end of Holv-
oak Lake), ca. 8 km southwest of Chemainus:
48°.54' X, 123°50' \\^; elevation 1060 m asl; 25
.August 2001 (2 .specimens collected b\' K. 0\aska
L. Sopuck. and S. Beauchesne); RBCM 001-
002S2-001.

5. Mt. Hooper (shores of an unnamed lake), ca. 20
km northwest of Youbou; 48=^60' X, 124°29' ^\";
eIe\ation 850 m ;isl; 10 September 2001 (2 spec-
imens collected bv L. Sopuck, C. Engelstoft, K.
Ovaska, and S. Beauchesne); RBC.VI 001-0028.3-
001.

6. Indian Creek, ca. 9 km X of Ucluelet on KemiecK
Flats: 49°02' X. 125=32' W": ele\ation < 20 nv. 18
Xo\ ember 2001 (collected by L. Sopuck, B. Beas-
ley, K. 0\aska, W. Leonard, G. Shreiber, and J.
Ziegltnim) (specimen used for genetic studies:
photograph in persoiuil files of KO).

\Va.'ihin0on State. United States:

7. Clallam Co.. 4.8 km south of BKii (off Woods
Road), Olvmpic Xational Forest; 47°59' X,
123°00' W": elexation 150 m asl; 20 Ma\- 2001 (col-
lected b\ W. Leonard and B. Moon >: RBCM 001-
00285-001.

8. Lewis Co., Gifford Pinchot XF. 14.5 km west of
\\'hite Pass (north side of SB 12); 46°4()' X,
121=31' W; elevation 853 m asl; April 2000 (8
specimens collected bv K. Jcirgensen and W. Leo-
nard); RBCM 001-00286-001 ; 12 April 2002 (2
specimens collected In W. Leonard "i; RBCM 001-
286-001.

9. King Co.. Mt. Baker-Snocjualmie Xational Forest,
se\eral locations from 0.8 to 1.6 km west of Ste-
vens Pass; 47°54' X, 121°06' ^\"; elevation 1100 m
asl; April and Mav 2001 (3 specimens collected
bv W. Leonard and W". XulD: RBCM 001-00287-
001.

10. Kittitas Co., Wenatchee Xational Forest, Ski
View; 47°23' X, 121°22' \V: ele\ation 1370 m asl:
17 October 2000 (collected b\ J. Bauijhi: RBC:M
001-00359-001.

11. Kittitas Co., Wenatchee Xational Forest, Granite
Creek: 47°09' X. 121°05' W; elevation 927 m ash
19 October 2000 (collected by J. Baugh) (dissect-
ed specimen lost)

12. Kittitas Co., Wenatchee .National Forest. Cold
Creek; 47°20' X, 121=22' W; elevation 80S m asl;
1 Xo\ ember 2000 (collected b\ J. Baugh V RBCM
001-00361-001.

13. Kittitas Co.. Wenatchee Xational Forest. Mos-
quito Creek; 47°18' X, 121°20' W; elevation 775
m asl: 26 October 2000 (collected b\- J. Baugh);
RBCM 001-00360-001.

14. Kittitas Co.. Wenatchee Xational Forest, Swamp
Lake, 47°19' X. 121=18' W. elevation 744 m as!:
9 Jtiiv 2001 (collected h\ J. Baugh): RBCM 001-
00289-001.

15. Kittitas Co., Wenatchee Xati(mal Forest. Roaring
Creek: 47=20' X. 121=22' W; ele\ation 810 m asj;
I 1 JuK 2001 (2 specimens; collected ii\ J. Baugh);
RBC-M 001-00288-001.

THE NAUTILUS 116 3 ;95-104. 2(>ii£

Pa^f 95

Xe\\- species of deep-water Cancellariidae (Gastropoda) from
the southwestern Pacific

Philippe Bouchet

Museum National d'Histoire Naturelle

55 rue Buftoii.

75(h:»5 Piuis

FRANCE

pbouchet@ cimrs 1 . mnhn . fr

Richard Petit

P.O. Box 3(1

North MxTtle Beach. SC 29597 US.\

r.e.petit@att.net

.\B.sTR.ACT

One new genus and nine new species of Cancellariidae are
described from New Caledonia from deptlis behxeen 2l)() and
6(X) meters. Thev are: Africotriton mhJphum new species. Mir-
auilaplwra new genus. MiraudiipJuni cai/rci new species. Mir-
audapJwra maestratii new species, Mcrica mtiriscti new spe-
cies. Sii-ltia rocroii new species. Sifhia spleiulidiila new spe-
cies. SippoiuipJicra pardalis new species. Siiipoiiaphera ctj-
phoma new species, and Sipponaplwra goniata new species.
Africotriton adclphum new species is the first species in that
genus known from outside South .\frica and .\ustralia. The new
genus yiirandaphera is characterized b\ its broad, non-unibil-
icate shell witli \er\ lar^e crenulated axiiil ribs, and axial col-
umella. The genus is composed of die new species described
herein, Mirandaplwra macstriitii new species ;uid \f. caijrei
new species. ;uid two odier species: A/, iosaensis iHabe. 1961
new combination and M. arafurcusi.s Aerhecken. 1997' new
combination, from deep water off Japan and the .\rafura Sea
respecti\ely. Triaoiiaphcra tcramachii Habe. 1961 ;uid Aiiatrix
nodosiharicosa Petuch, 1979 are tnuisferred to Xippoiiiiphera.
New species of Merica. Svrltia. and Sippoiuiplicni are the
deepest dweUing knowii representatix es in their respecti\e
ijenera.

INTRODUCTIOX

Tiie tropica] deep-water taima oi the soutliwestem Pa-
cific w;is xiiinuilK unknown just two decades ago. but
considerable adxiuices ha\e since been made both in
tenns of sampling in die field and ta\ononiic;il descrip-
tions ot the materiiil collected. Empiuisis h;is been
placed on die exploration ot the iu"ea around New Ca-
ledonia, resulting in the descripdon of se\ end hundred
new niolluscan species isee. among others. Crosnier .uid
Bouchet. 1991: Bouchet. 1995: Bouchet and .Marshall.
200U, .\lthough the inventon is still tW from c-ompiete.
it h;is ;ilread\ been revealed to be one oi the most — it
not the most — dixerse an\-wliere in the world, with an
exceptional dixei-sification in such families its .Seijiienzii-
dae (55 species: .\I;irsIuill. 199U. Muricitlae ^90-201)
species: Honart. 2001 K or Scaphopoila [73 .species: SciU"-
abino. 1995).

ba.scti on tile literature, the tamiU t^aiici-iianKlac ap-
pears poorly represented in the tropical South Pacific-,
xvitli h;df a dozen nominal species rec-orded from shalloxv
xxater in Nexv Caledoni;i. to just h\o reported from Fiji,
and a single species from the Socieh Islands. Bx con-
trast, die material originating from tlie recent expedi-
tions in Xexx C-aledonia. X'amiatn. Fiji. Tonga, and \\'allis
iind Futiina contains about 50 species of Cancellariiilae.
about -W ot xxhich are undescribed. mainlx- from depths
betxxeen 2CX) and 6tX) meters. The purpose of the pre-
sent paper is to proxide descriptions of die more spec--
taciJiir ot the nex\ species -X Tiiore c-omprehensix e
monosrapli. describing an ing the entire can-

ceUariid fauna ot the regio;.. ox\. We restrict our-

selxes in die present paper to the description of shell
moqihologx. ;md deter to the sect)nd article descriptions
of radulae ixxlien axailable' and discussion ot patterns of
batlixnnetrical ;md geographical distributions.

M.\TERLALS AND TEXT CONN EXTIOXS

The nexx- species orisjinate troni several dredginsj pro-
grams conducted under Dn Bertrand Richer cie F"orges.
of Institut de Recherche pour le Dex eloppement IRD.
foniierix- ORSTOMl Xoumea. The cxnilext oi the pro-
grams. Uiirratixes of the expeditioiis and lull station lists
are presented bx Richer de Forces 1990' MISOR-
STOM 4'. Richer de Forges 1991 iProgranune L\-
GOX'. Richer de Forges 1993 S.MIB 5'. and Richer
de Forges imd Chexilion 1996 S.MIB S. B.\THUS 2.
3. and4\

In the lists of t\pe and other materiid examined, in-
dixidnal lots in NIXTIX are unambiguouslx designated
bx- die combination of cruise acronxni and station num-
ber. D\\" refers to dredge hauls. CP to lieam traxvls: l\
refers to live-taken specimens, dd to emph shells; spnis
to commerciallx ontained specimens, state xx-hen
dredged not kniowii.

specir

SYSTEM ATICS

Fiuiiilx Canc-ellariidae Forbes and Hanlex. 1S51
Snbfaniilx- Plesiotritouinae Ben iuid Max-xvell. 19S7
C.enus AfiicotritoJi Ben and .Maxxvell. I9S7

Pa-'e 96

THE NAUTILUS. \ol. lift. NO. 3

Figures 1-S. Xcw deep-water C'anccllariidar l-.'5. .XfiicdliUdii inliliiliiiiii mw s|H'eics \. Il(ilcil\|)c lici^lil 22,4 iiiiii scmtli of
New Cialedonia. SW of He des Pins. 22°.1T' S. I(i7 22' K. .590 m. 2. I'kiIikiiikIi of siiccimen Iroiii SW ol lie des I'ms. 22 4.S' S.
Ifi7°lf>' E. 444 -t4.5 in |HATIIl)S 2: sla. D\\7I9|. 3. Teleoeoiuli iiiuiiisciilpluic, s.inie specimen a.s Figure 2, 4-tt. Miraiidaplwra

timcstnilii new specie.s. 4. Holotvpe. Iieiijlit 14.0 ii nlT nordieastern eoa.st of New ( .'aletloriia. Passe de llientjliene. 2()".'3;3' S.

I64'.5T' v.. .5:«-61() ni. 5. ProtDconcli of speenneii Inmi Wallis and Kntnna. I.ri9' S. 17(i°l7' W. .3.50 ni [Ml!S( )R.ST()M 7: sta
D\\fi01|. 6. Teleoeoneli niierosenlptnix-. same specimen as Fii;ni'e .">. 7-8. Minniildphcni ciii/iri new species. 7. Specimen. Iniiilil
27.3 nnn: norlli of .\ew Caledonia. UfOT S. I(i3°13' K. .34I-.3.5I ni iB.VriirS 4: sla. in\»021, S. Ilolotvpe. Iieiijht .3S.7 mm;
South of New Cal.-donia. off Passe de Kouare. 22" W S. Ui6 l.'j' K. .3(lll-.)70 m. Se.ile Imes - 100 |i.m (3. (Si and .'lOO |jlui (2, .5).

Type Specie.s: Epiilnmiiis (■rchrilinitiis i,. 15.
Sowerbv III, 190.3. In orit^inal designation.

Africotrilon adclpiiiitn new species
(Figures 1-3'

T\pe Material: Ilolotvpe (22.4 mhii lieiglit X 7.S iiiiii
width) (1\) (Figure I) and 3 paratopes (ddi. all from the
t)pe localitA. in MNHN.

'INpe Loealilv: .South ol Nt'w (^aK'douia. SW ol lie
di's Pins. 22°47' S, lfi7°22' P]. .390 ni | MUS( )HSr( )\l
4:sla. nW22fil,

Oilier Material Fxainined: South ol New Caledonia.
HlOCAl.: sta 1)\\44. 22 47' S. Ifi7"l4' E. 440-4.50 ni,
2 ju\. K. I in\. (I(L- S.MIH 2: sla. DW'l, 22°53' S.
167' I. r E. 43S-444 in. I jii\. K.— S.MIB 3: .sta. D\V2fr

p. Bnuclirt and R. Petit. 2002

Pasie 97

22"55' S. 167"16' E, 450 m. 1 K.^BATlllS 2: sta.
DWTUJ. 22°4S' S. lfi7°lH' E. 444-443 m. 4 K iFi^uivs
2, 3).

Description: Protde-oiu'li iiiaimiiilatf. paiii'ispiial. of
1,1 whorls, diameter 1.25-1.30 mm (Figure 2). Traii.si-
ticiii to teleoconeh marked h\' oii.set of closeK .spaced
spiral cords. Teleoconeh ol' sexen moderatcK ronnded
whorls; spire angle 2fi'-27^ Prominent ortliocline
rounded \arices tliat extend al)o\e the appressed suturi'
are formed irregularK. hut nsualK at ahout 240" inter-
vals. Spiral sculpture ol over (SO low closeK spaced spiral
cords. ."Vxial sculpture of widel\ spaced, rounded ncm-
collaiiral rihs, 7 between final two varices, witli closeK'
packeil microscopic growth lines (Figure 3). .\perture
narrowK' o\"ate. Outer lip with weak crenulations on
edge hut smooth within. .\ shielddike callus produced
on the inner lip. weak ovi-r the parietal area h\it well
produced oxer the imluctural area and extending o\'er
the siphonal constriction. Columella ahiiost axial with a
narrow fold at the center. A second. \en weak fold pos-
terior to the center fold is xisihie on some specimens.
Siphon. il canal narrow, opt'ii, strongK recnnetl. Shell
color cream to vellow-hrown with fi or 7 red-hrowii spi-
ral bands, narrow on small shells but wiile on some
adults, appearing as weak hands except on the tops of
the varices where the\- appear as narrow stripes.

Etvmolog)-: The Greek ddcliihos. brother, utilized to
highlight the resemblance of the species to others in the
genus; treated here as a Fatin adjective.

Remarks: Of the si.\ species inchuled in this genus when
introtluced, five are from South Africa and the sixth, .A.
carifiiipcx Beu and Max-xvell. 19S7, is from off New
South Wales, Australia. The new species differs from A.
carinapex in being much more slender with less convex
whorls, and in having stronger spiral cords and weaker
axial ribs. Africotriton mlelphum resembles more the
South African A. Mlhurni Beu and Maxxvell, 1987. but
that species has more prominent spiral cords and weaker
axial ribs.

Subfamilv Cancellariinae Forbes and Hanlev. 1S51
Genus Mirandaphcra new genus

Type Species: MiraiuUiphcrd c<np-ii new species

Description: Shell rehitivelv large, reaching over 35
mm. Teleoconeh of 7-9 whorls. Shell somewhat atten-
uated vvitli large axial ribs that dominate the shell sculp-
ture. Ribs angk'd or loimdcd at the shoulder over wliicli
tlun project ;is sm;ill coi'onations but rarelv extend hack
to the sutmc. Sutinal ramp depressed. The sutiuc is
onlv slightlv impressed. Small nodes are formed where
weak spiral cords cross ribs. Spiral cords are nnich stron-
ger anil more evitlent on the ribs than in the interspaces.
Outer lip tliick. slightlv flared on outer edge and ex-
tending back over tlie paiietal wall as a rounded shield
that extends out at tlie imluctural area. No stromboid
notch visible on outer lip. Columella axial with two

strong folds and a thiid fold which slls on the ediie of
the siphonal fold before turning avvav to parallel other
folds. Folds do not extend out to the edge of the iiidue-

Inral shelf I nibilicns absenl.

Discussion: ik'sides the twd new species described
here, we include Tiitidiuiphcrii lo.sdciisis Ilabe, 1961 {re-
centlv |il;iced in Sohilia In llasegavva, 2001: 5S3) and
Solatia arafiirciisis N'erheckeu. 1997 in Mirandaphcra
The resemblance of these t;ixa to Si>latia is onlv ven
superficial. The tvpe species of Solatia. S. piscatoria
(Gmelin. 1791 ). has weak colimiellar folds, an apjiressed
suture, and a flaring apertmc among otiier differences.
Species ol Miunidaplwra iiave a superficial resemblance
only to high-spired species ni' Scalptia Jousseaume. 1SS7
from which diev ditler in having an elongated tabulate
shell with vvi-v large crenulated axial ribs and a axial col-
umella Minindaphcra. as understood here, is a deep-
water genus. No fossil spi-cies luivc been reco^jni/ed in
the literature.

Et\'moloii;\ : The Fatin iiiiraiidus. nn'aning wonderful
Ol' sli'iinge. eoinbined with aplwra. Apliera was intro-
duced as a g(.'uus in Cancellaiiidae and later used as a
stem name for various cancellariid genera, all treated as
feminine. A search for its origin reveals onlv a few us-
ages, the earliest of which are as the given name of wom-
en in Eni'Iand in the mid-17''' C'entnn.

Mirandaphcra cai/rci new species
(Figures 7-S)

T%pe Materia!: Ilolotvpe i3S.7 mm height X19.3 nun
width) ilv 1 in MNllN i Figure Si.

Type Locality': South of New Cafedonia. oft Passe de
Kouare. 22"49' S, 166°45' E, 300-370 m [B;VTHUS 2;
,sta. D\\'7311.

Other Material Examined: North of New Caledonia.

B.-VTllUS 4; sta. 1)W9()2, 19°()1' S. fa3=15' E. 341-351
m. 2 dd (Figure 7i. — l-'rom conunercial sources, boat
Tui IF off I^elep Islands. 3 spms.

Description: Protoconch smooth, of 1.3 whorls, diam-
eter S30 [jLui. Transition to teleoconeh marked bv strong
axial ril). Teleoconeh of about 9 elongate whorls. .■\.xi;il
sculpture of strong rounded ribs, about 12 on bodv
whorl and 14 ou penultimate whorl. The ribs are bian-
gular on the shoulder over which thev recurve and ex-
tend partialK onto the othenvise smooth, fiat sutural
i'am|). Sulni'e barelv impressed. S]')iral sculpture of about
S priman eoi'ds that form prominent noiles wiiere they
cross the iixiai ribs. Nodes strongest on (he shoulder.
wide, biangular .\bout 4 priman spiral cords visible on
penultinuite whorl. Numerous fine secondarx spiral
cords cover the shell. S or more between each pair of
priinan cords with an occasional si'condarx spiral cord
intermediate in strength, not intersected by axial riblets.
Ribs on adult shells sometimes formed as thickened var-
ices with subse(juent growth originating from under the

Page 98

THE NAUTILUS. W

IH, No. 3

outer lip. Aperture elougate. Outer lip tliick, .siiiootli.
with a narrow, smooth shelf inside the lip. About 12
strong lirae extend from lip onlv a short way into the
aperture. A posterior canal is formed under the shoulder
and is further delineated In' a pustule on the parietal
Willi. Shield-like parietal callus well-developed. Colu-
mella with three strong folds, the anterior one weakest
and bordering the distinct siphonal fold before turning
to parallel the other folds. Folds do not e.xteud to the
end of the inductural Ciillus. Body whorl onlv slightK
constricted behind the siphonal fasciole. Umbilicus ab-
sent. Shell color off-white or beige: prinian' spiral cords
slightK' darker, light \ellow-bro\\Ti.

EtjTnolog>: Named for Dr. Patrice Ca\Te, head of
IRD's Department des Ressources \'i\ antes, in recog-
nition for his support to ta\onom\' and biodiversity' ex-
ploration.

Remarks: Minnulaphcra cai/rci differs from M. to-
saeiisis (Habe, 1961) and M. (mifiireims (\'erhecken.
1997) in having strong lirae within the aperture and a
more turreted, attenuate shell on which the spiral sculp-
ture is prominent in profile. Miranchiphera maestratii
has, among other differences, more numerous axial ribs,
stronger spiral sculpture, and a more rounded profile.

Miniiulaplwra iimc'-itratii new species
(Figiu'es -f-6)

T\pe Material: Holohpe (14.0 nun height X 8.4 mm
width! idd' in .\1NHN (Figure 4).

Type Locality: Olf northeast coast of New Caledonia,
Pas.se de Ilienghene, 20°33' S, 164°57' E, 533-610 m
[BATH US 4: sta. D\V948].

Other \taterial Examined: \anuatu: MUSORSTOM
S: sta. DWlOfil. Krl5' S. i67°20' E. 458-512 m, 1 dd;
Fiji: MUSORSTOM 10: sta. CP1341, 16°52.5' S,
177°43.7' E. 500-614 m, 1 dd; Wallis and Futuna: .MU-
SOR.STOM 7: sta. D\\Ti01, 13°19' S, 176°17' W, 350 m,
1 dd (Figures 5. 6),

Description: Profoeonch prominent, of 1.3 whorls, di-
ameter 850 [xm (Figure 5). Onset of teleoconch markeil
by a strong axial rib. Teleoconch of about 6 whorls or-
namented with high narrow axial ribs barcK wiiler than
interspaces. The axial ribs. 14-16 in number, extend over
the rounded slioukler and turn down to a narrow sutural
ramp, but do not extend all the wax to the moderat(>K
impres.sed suture. Spiral sculpture of narrow tleu.s(>l\
packed spiral coi'ds l^liat become denser and increase in
height at regular intenals, forming bead-like nodes on
the ribs. There are 8-10 nodes on the bod\ whorl ribs
with approximately 16 cfirds bcKveen centers of nodes.
Spiral cords intersected In finer incremental riblets that
form an intritacalx-like microsculpture (Figure 6). I'inal
rib is produced as a terminal \arix. Aperture elongate,
narrow. Outer lip thickened with 8 lirae within that ex-
tend to the outer edge btit do not tiescend dei^pK into

the aperture. Posterior can;il is formed under the sh(ml-
der and is further delineated b\- a pustule on the parietal
wall. Parietal callus well-developed but not extending
out as a shield. Columella with three strong folds, the
anterior one weakest and bordering the siphonal told
before turning to parallel the other folds. Folds do not
extend to the outer edge of the inductura. Bod\' whorl
slightlv constricted behind the small but w^ell-defined si-
phonal fasciole. Uml)ilicus absent. Shell color light
brow'nisli-\'ell(n\.

EtjTnoIogj': Named lor Philippe Maestrati, .MNHN
museum technician, in appreciation for his dedication to
processing and sorting much of the New Caledonia ma-
terial.

Remarks: Miraiulaplwra maestratii is separable from
M. caijivi h\ its closeK spaced axial ribs. The spiral cords
of M. maestratii are also unique, being so closelv spaced
tliat the interspaces appear as incised lines. The delicate
pattern the\' form is especialK' noticeable when there is
a \ailx on which the\' are \isible.

Genus Merica H. and .-K. Adanis, 1854

T\'pe species: CanccUaria mclanostoma Sowerby,
1849, b\' subsequent designation of Cossmann (1899).

Merica maiisca new species
(Figm-es 9-12)

T\pe Material: Holotvpe (25.7 mm height X 18.0
nnii width) iK ) (Figure 9) in MNHN.

T>'pe Locality: ( )tl northeast coast of New Caledonia,
Passe de Hienghene, 20°34' S, 164°58' E, 470-190 m
[BATHUS 4 sta. CP947].

Other Material Examined: New Caledonia: BA-
THUS 4: sta CP889, 2r01' S, 164°27' E, 416-433 m,
1 dd (Figure 12).— HALIPRO 1: sta CP851, 21°43' S,
166°37' E, 314-364 m, 1 Iv (Figure 11): Vanuatu: MU-
SORSTOM 8: sta. CP1136, 15°41' S, 167°02' E, 398-
400 m, 1 dd (Figure 10).

Description: Protocoucli smooth. shin\. with 1.0
wliorl iFiguri' 12). Transition to lcleoconi-li marked In'
a slight depression and onset of axial and spiral sculp-
tures. Teleoconch of up to 6 rovmded whorls. Shoulder
small, rounded, sloping back to an impressed suture
lorniing a narrow channel. Axial scul[)ture of about 1-1-
16 low axial ribs on b()d\ whorl of adult; slightlv more
on earlier whorls. Ribs rounded in profile and narnnver
than intencTiing spaces. Periodic inti-rnal \arices (see
Harasewxch and Petit, 1982: ill), noticeable on outer
surface of shell as thickened axial ribs, each followed by
a relati\('l\' flat area. Spiral sculpture oi strong, broad
cords ( 16-20 on bod\' whorl, 6-7 on pemiltimate whorl);
those on shoukk-r cnnvded. others usualK with a sec-
ondan cord in the interspaces and rareK a tertian cord.
.\pcrture lai'iif with ;i wideK elliptical, prosocline outtn"

p. Bouclu't

R. Petit. 2002

Page 99

Figures 9-17. Xe« deep-water Caneellariulue. 9-12. Mcrica mari.sca ne^^ spec.es. 9. 1 l..l..t>pe. l,ei>.|,l 2.yT mmu, oti ,K>rtheasten.
coast of New Caledonia. Passe de HienaliiMie. 2(f34' S. 164°5S' E. 470-490 >n. 10. Specn„e„ Iron. Xu.u.atn, he.gl.t - 1 , .9 .....K

MUSORSTO.M S: sta. CP11361. II. Specin.en troni New Cidctlon.a. I.eiglit IS., mm: SI 45 s.

12. Protoconcli of specimen fiom off the western coast o( New Caledonia.

CP.S891 13-14. Sieltiii ivrroii new species. 13. Ilolot\pe. I.eiglit 14.3 mm:

Onest. 23°20' S. 16S°01' E. .501-365 m [B.\TiIUS 3: sta. D\\S.30]. 14.

15°41' S. 167°02' E. 39S-400 m

166°3T' E. 314-.364 m [HALIPRO 1: sta. CPS.51]

21°01' S. 164°27' E. 416-433 m [B.-\THUS 4: sta

south of New C:aledonia. Xoifolk Riilge. Banc Jumean

Protoconc

15-1

167

23'

Sc;ile line

lull ui .\e\\ \..aie(.UMiia. -\vhiuii\ iiivii^^r. i.»txiis. uiiiv^n. ^..v.... — — . ri..i/-A/"«i i i^M'O"

otoconch of specimen fron. sonth of New Caledonia, S\\- of He des Pi.is. 23=0()' S. 16, 16 E. .5o( -n [BIOC.VL: sta. D\ 3,].
1-17. Svcltia splculklulu ,iew species. 13. Holot^pe. height 11.0 mm: so.,th ol New Cak^lonia. S\\ of He d.-s I ins. oO S.
i7^2P E 500-.504 m [SMIB 8: sta. D\\-2(l]]. 16. Parat\pe. height 10.1 nm.: south ol New (.aledcmi:.. S\\ ot He des I ms. - o9 -
;°00' S 167°21'-I67°23' E 491-558 m [SMIB 8: sta i:)\\lH.3-196l. 17. Protoco.ich: specime.i Worn s:mie local.t^ as l-ii;nre Hr

.500 M-m (12. 14. 17).

Page 100

THK XAITIIAS, \<)1. llfi. No

lip. Outer lip with a HiuK stTiati'd i'(l<ic; inner surface
witli 14 spiral lirae that (iiniinish aliout a (|uarter whorl
into the aperture. C'olnnulla with ■> folds, the posterior
one being slightK larger and the anterior one torniiug
the edge of the short, narrow canal. The anterior lold
extends to the edge of the inductural callus with smaller
fold-like features or pustules sometimes present on out-
er edge of inductnra. Color \ellow-hro\\ii with a dark
red-hrown hand at shoukler and smallei' lines or hands
on l)od\ whorl. \ white hand is sometimes present just
anterior to the shouldei' ami just anterior to the periph-
ery.

Etvinologv: Latin iiiaii^ca. f . a large t\pe of fig.

Reniark.s: Mcricd piniiii has many features in com-
mon with the Philippine Merica ekti/phos Petit and Har-
asew\ch, 2(100. which has stronger sculpture with lieavy
nodes formed at intersections of spiral cords and ;L\ial
rilis. In addition, M. ckfi/plnis has a roundetl and more
solid shell, and the protoconch is '4 whorl larger than in
M. piniin. Species of Merica li\e suhtidalK' and on the
continental shelf The batlnal M. piniiii is apparentK (he
deepest living species in the genus.

Cienns Svcltiii [oussi'anme. 1SS7

Type Species: "Svclliti i(nico\<i [sic| 15rocc." [= Vol-
uld rarricosa Broci-hi, fSf4l, h\ original di'signatiou.

Svcltid rocroii new species
(Figures 1:5-14)

T\pt' Material: Holot\pe (14.3 mm height X S.9 nun
width ! iK 1 (Figure 13) and one paratope (dd) from the
t\pc locality in MNHN.

Type Locality-: South of New Caledonia, Norfolk
Ridge, Banc jumean Ouest.. 23°20' S, lfiS°Of' F, 3fif-
365 m |B.\TIIUS 3: sta. DWS30|.

Other Material Examined: New C;aledonia: BiO-
CAL; sta. D\V.37, 2.3°()()' S. IfiT'lfi' E. 350 m. 1 ju\. K.
(Figure 14), 1 jnv. dd.— MUSOKSTOM 4; sta. DW'lSl.
18°57' S, 163°22' E, 350 m, 1 dd.— BATHUS 4: sta.
D\V931, 1S°55' S, 163°24' F, 360-377 m, 1 dd.— From
connnercial sources, boat Tui II, off Belep Islands, 3
spnis.

Description: Protoconch smooth, erect of I O whorl.
diameter ! ..30 mm. axis of protoconch tilted to teleo-
conch (Figure 14(. Transition to teleoconcli eleark
marked by a strong \arix-like line, angled at the shoulder
and followed b\- onset of axial ribs. Teleoconcli of about
five whorls, .\\ial sculpture of low, wideK spaced libs,
se\en on body whorl and ten on |iennltiiriate whoil. ex-
tending from the suture to the ba.se ol the shell Spiral
sculpture ol one cord situated niidwax on whorl and
forming an angled shoulder w ith sharp short sjiines |m'-
sent at intersections with iLxial ribs. A weaker spiral cord
is .sometimes present betvxeen the shoulder and the

slightK impressed suttu'e. About fi\c weak spiial colds
are located anterior to the shoulder, the first anterior one
widely spaced from shoukler and forming a slight angle
to the bod\ whorl Remaining spiral cords weaker and
more clost'K' spaceil. ,\pertnri' large, o\al. Outer lip pro-
socline. formed b\' a terminal \arix, smooth within. Pa-
rietal wall with weak callus, becoming stronger and
slightK' reflected at the inductnra. Columella with three
strong folds, the anterior one forming the edge of the
short siphonal canal. Shell thin, white, translucent.

Et\iiiolog\': Named for M. )ean-Paul Rocroi. .\INHN,
in appreciation ol liis contiibutiou to inollnscan iioinen-
clature.

Remarks: The new species ina\ be cmnparetl to oiiK
one Recent species other than Svcltia splendidula . de-
scribed herein, is S. 'gladiator (Petit, 1976), which occurs
off the Cialapagos Islanils at a depth of 200 m. Although
their protoconchs are entirelx different (the protoconch
ol S'. f^Iddiator is nmltispiral indicating planktotrophic
lar\al dexelopment), the two species share the "window-
like" sculpture below tlie shoulder formed h\ the wideK'
spaced cords and ribs. The spines on the shoulder of .S.
nx-niil are much shorter than in S. fjjadiator. The Elu-
ropean Tertian species Calcarata calcarata (Brocchi.
1S14) is startliugK- like S. rocroii in appearance but that
species differs, among other features, in being slightK'
umbilicate.

Sieltia splciididiild new species
(Figures 15-17)

Type Material: llolot\pt' ill.O mm height X 6.9 mm
widtli) (K) (Figure 15) and three paratxpes (1 l\. 2 dd(
in MNHN.

Type Locality: South of New (Caledonia. SW of Ik'
d(''s Pins, 22°5b' S, 167°21' E, 500-504 m [S.MIB S: sta.
l)\\'20fj.

Other Material Examined: South of New Caledonia.
SW of He des Pins; SMIB S: sta. D\\I93-196, 22°59'-
23°()0' S, 167°2r-167°23' E, 491-558 m, 1 Iv, 2 dd
iparatspes: Figures 16. 17).

Description: Protoconch smooth, erett. ol 0.9 whorls,
dianu'tei' 1.20 mm, axis of j^rotoconcli tilted to teleo-
conch (Figure 17'. Transition to teleoioiu-h inaikeil li\
the appearance ol an axial rib with angled shouldei'. Te-
leoconcli of .3+ lapidK expandinti whorls. Axial sciilp-
(iirc ol eveiiK spaced sharp ribs, about 12 on jioiK whoil
that ai'<' contiiinoiis Ircnii the suture to the base. I'he
libs are produced sli'.^lilK above and over siibseijneiit
grow til. .Spiral sciilplme ol about seven sliaq) sjiiral
cords. ( )ne cold, soiiiew ha( stronger. I onus the shoulder
over w I lid I the a\ial ribs cross ,md angle back to a sliglit-
K impressed suture. .\ second strong coril sometimes
|)reseiit at line of sntiiral attachment. On the final por-
tion ol the bodv whoil the ainj;led shoulder becomes ob-
solete and the ribs eiiivc siiiootlilv b.ick to an impressed

F. I^oiiclict and R. Petit. 2002

Page 101

Fissures 18-27. New ilccp-watiT ( laiici'lluriidac. 18-21. Mjtjxniiiiilirin pariliilis
nuitli ol New C:aleaiinia. CrMul l'assa-;r. 19 11(V S, 16.y29' K. 230 m [MrS()HST(

mm: north of New Caledonia. Craiul Passaiic 19 Di >. loo .>ii I'.. — ^i m | i.av..v_;.\. Ma, ii-^ji^- ' , .. i i

-.rid 111 ISMIB S; sla. 1)\\ ISl]. 21. .Mieroseiilptnrc ol teleoeoncli;

)rtli ol' New CaledonI;

/(A m-w >|H-eK'S. 18. l-araUpr. hciul.t 20.0 iiinu
lOM 4: sta. D\\1S5|. 19. liolotvpe. lieii;lil 2.5.5
ON sta 114S1. 20. rrotoeoiuii: spceiiiien IVoiii

South ol New

Scale lint

10(1 fjLiii !21. 24, 27' ami .500 ijliii

Pa2c> 102

THE NAUTILUS. Nol. 1 Ifi, NO. 3

suture. Thf .spiral cords do not cross over the a.\ial ribs.
Aperture large, ovate. Outer lip prosocliiie, tliin, without
internal lirations. Parietal area without callus. A small
callus is sometimes formed at \er\ base of intluctura.
Columella with three folds, the posterior one larger and
descending. The two anterior folds situated at ends of a
shelf-like structure bordering the short siphonal canal.
Bodv w'horl sliglitK- constricted behind the weak siphon-
al fasciole. .Shell color white, translucent.

Et\Tnolog\': Dimimitixc of the Latin adjectixc sjilcii-
(lidiis [-U. -HuiK meaning show^■ or magnificent.

Remarks: Sicltia s])hn(lid\d(i closeh' resembles S'. ro-
croii from which it differs in possessing more numerous
and stronger spiral cords and a\ial ribs, and in ha\ing a
much less angular shoulder. The unusual combination of
sculpture and translucence gi\es the shell a shoji-like
appearance. The bathvmetric distribution of the geuus
Sveltia is unclear due to the confused generic allocation
of included species, but Sveltia splcndkluhi is apparentK-
the deepest dwelling species in the genus.

Genus \ij)po)iiii)licra Ilabe, 1961

T^pe .Species: Xippiiii(ii)licni Itnhci Petit, 1972, In
iC.ZX Opinion 1052,

Nipponaphcrn fxinldlis new species
(Figures 1S-21>

Type Material: Ilolotvpe (25.5 m height X IS.O nun
width) (Iv) (Figure 19) and one parat\pe (Iv) (Figure 18)
in MNHN.

Type Locality: North of New Caledonia, Grand Pas-
sage, 19'()7' S, ](i:r:W K, 220 m [LAGON: sta. 114S].

Other Material E.xaininecl: New Caledonia: .\1U-
SORSTOM 4: sta. D\\T85, 19°0(i' S, lfi.3°29' E, 230 m,
I 1\- (parat\pe. Figure IS).— Sta. DW227. 22°46' S.
167°20' E, 300 m, 1 (kl— SMIB 5: ,sta. DWll. 23°42'
S, 168°0r E, 400 m. 1 dd [worn; allocation uncertain].—
SMIB 8: sta. nW'ISl, 23ns' S, lfiS''05' E. 311-330 m,
1 juv. Iv (Figures 20, 21).— Sta. 1)\\ 1S2-1S4, 23°18'-
23°19' S, 168°05' E, 305-367 m, 1 (kl.^BATHUS 2:
sta. D\V714, 22°38' S, I67°10' !■:. 121 m, 1 (kl,— BA-
THUS 4: sta. D\V942, 19°04' S, 163°27' E, 264-270 ni,
1 juv, 1\. — From commercial sources, boat Tui 11, off
Bele|) Islands, 3 spms.

Description: I'rotoconcii corrodid but appaicniK ccm-
.si,stiiig of about 1.5 whorls, teleoconch of 5.2 whorls.
Spire high. Sculpture of imbricated axial lam(>llae and
raised spiral cords that cross to form S(|narish intenals.
Aperiodical axial \arices, six on final whorl. fi\c on pen-
ultimate whorl, and two on antepenultimate whorl. Spi-
ral sculpture of about 10 spiral cords on peimltimale
whorl and about 20 on final whori Second oriler sculp-
ture of iTicriMuental lines crossing spiial threads and
forming fiucK reticulate sculptiuc in the s(|narish inter-

vals (Figure 21 ). Spire whorls regularly convex, gradualK'
l)ecoming angular at shoulder on final two whorls. Su-
ture deepK' impressed. Both' whorl onl\ slightK con-
stricted behind siphonal fasciole. Outer lip thin, smooth
innnediateh within but then with about 18 lirations that
extend deepK into the aperture. Inner lip reflected as a
parietal shield partly extending over the umbilical area
in the inductural region. Columella slightlv concave witli
three folds, the anterior two folds close together, unich
like one large bifiu'cate fold. Deep umbilicus well de-
veloped. Siphonal canal short but distinct. Shell liack-
ground color cream\-wliite with brownish blotches on
the periphen' of the whorls behind \arices; priman' spi-
ral cords brown on the varices.

EtymologA': The Latin panlalis. a female panther. To
be treated as a noun in apposition. Selected because of
tile brown blotches behind the varices.

Remarks: The protocouch of the holot\pe is corrod-
ed. The protoconcli of the paratxpe consists of 1.8
whorls, diameter 1.15 nun (Figure 20). Among the spe-
cies of Nipponapltcra known at present, this new species
most resembles .Y. tciriiiuicliii (Habe, 1961), new com-
bination, which is more distinctly angled at the shoul-
(lei; and has coarser sculpture, (The transfer of Tr/go/i-
iiplicra fcriiiiwchii to Xipjioiiaphcra will be discussed in
oui' later monograph. )

Nipponaplwni ci/phoniti new species
(Figures 22-24)'

Type Malei-ial: ilolotxpe (13.1 height X 9.2 nun
width) (ddl (Figures 22-24) in MNHN; 2 panit\pes.
Petit collectif)n.

Type Locality: North ul New C^aledonia. (irand Pas-
sage, 18°55' S' 163°24' E, 350-365 m [HALICAL 1; sta.
n\\04].

Other Material E.vaniinecl: From rommei'(i;il sourc-
es, off Hel(-p Island, dredged :it 400 ni In bo;it Tn II,
M;l\ 2001, 2 sjims (p;u";lt\]n's).

Description: Protocouch glassw white, with 0.9
whorls. dl;nneter 0.93 mm (Eigiu'e 23). Transition to te-
leoconch marked In ;i small \;nix-like axial rib followed
b\' spinil ;md ;ixi;il scnlptuic. Spiral sculpture of flat cords
ol xiUAiiig sizes, i'rimarv cords ni)ini;ilK separated b\'
slightlv smaller seconilan cortls with even smaller ter-
tian cords in iuters|i;ices between the jiriman- and see-
ondiUA ((lids li(i(l\ whorl willi 10-12 priuKUA spir;il
colds. ;ib()Mt 4 5 \isiblc on penultimate whorl. Spinil
(Olds sip;n'at('(l b\ ii;iirow sjiiices tluit iipjiear ;is iiKiscd
lines iiistc;i{l of s])aees i figure 24 >. .-\\i;il sculpture ol
loimded ribs, ;il)out 8 in number on the bo(l\ wlioii ;m(l
11 on the penultiiiKite wliori. SiikiII nodes ;ire present
where priin;u"S' s[)iral cords cross the axial ribs. On the
terminal \:uix there is a strong shoukk'r node. Shoulder
rounded b.ick to a b;ireK iiii|iresse(l sutur(>. Bodx whorl
loimded bill soiiiew luit coiistiicted behind the weiik si-

p. Boudiet ami H. Petit. 2002

Paw 10:3

phonal fasciolf. Onlci- lip thin, weakly serrati'. Iniici' pm-
ti(in of outer lip with a snmoth romuled shell' rnll(iw<'(l
In alniut 14 sharp short liiations. ( 'oliiiiiella Willi weak
callns on iiuliietural area. Ctiluniella with three lolds that
do not e.xtenil to the outer edge of the indnetiira. i'os-
terior told largest. Two anterior folds situated 011 ends
of a low shelf-like platlorin with the anterioiiiiost fold
becoming the edge ol the short well-toniied siphonal
canal. Umbilicus absent. Shell color mottled white and
light browii. ("olors arranged in wfak bands. Hmwii
bands are located at the shoulder, just antf^rior to the
periphery ami at the base.

EtyniologA': The (ireek ki/phoina, meaning hump,
with lelerenci' to the stiiixK axial ribs of tlie species. To
be treated as a noun in apposition.

Remarks: \ij)j>()iiili)lur(i ctipJutnui differs from \ip-
jxnifijiliciyi iiodosivtiricosfi (Petncli. 1979', new combi-
nation (originalK in .Agrz/ri.vi. in ha\ing iimeli finer spiiai
sculpture, fewer and more wideK spaceil axial ribs, and
a proportionalK- liroader shell. (The transfer of .Agrt?r/.v
uodosivaricosa to Sippoiuijiliird will be discussed in a
future monograph.) The batlnnietrie distribution ol the
genus Nipponapluni is imelear due to confused generic
allocation of di-scribed species, but Nippon(ij)lifni 11/-
plioma is apparenth' the species with the deepest occur-
rence in the genus.

Xippoiiaplicrii goiiiata new species
(Figiu-es 25-27)

Type Material: IIolot\pe (IfS.l iiini height
mm width) (dd) (Figure 25) in MNHN.

11,4

T^•pe Localit^•: .South of New Caledonia, Norfolk
lliilge. Banc A/ieijne jalso called Banc Antigonia on ma-
rine charts], 2:3°4r S, 16S°()1' E, 2S0 m [SMIB 5, sta.
DWTfi],

Other Material Examined: South of New Caledonia.
Norfolk Ridge. BIOCAL: sta. DW65. 24°48' S, 168°09'
E, 245-275^ll, 1 dd.— CALSUB: dive 21, 22°45' S,
lfi7°()9' E, 340 in, 1 K.— CHALCAL 2: sta. f^WTf,
24°42' S, ffiS°l()' E, 230 m, 3 dd.— SMIB S: sta.
D\VT54, 24°46' S, 1R8°()8' E. 235-252 m, 1 dd (Figure
27).— Sta. DW157, 24°4fi' S. 168°08' E, 251-255 m. 1
dd.— Sta. D\M58, 24°4fi' S, lfi8°0S' E, 262-290 m, 1
dd.— Sta. DW159, 24°4rV S, Ifi8°()8' E. 241-245 m. fi
dd (Figure 2fi),-^Sta. D\\T73, 23'41' S, 168°()0' E, 234-
242 m, 1 dd.— Sta. D\\T74, 23°40' S, lfi8°()f' E, 235-
240 m, 2 dd.— BERYX f f: sta. DWMS. 24M8' S, lfiS"09'
E, 250-270 in, f dd.

Description: Protoconch smooth, of sliglitK' more
than I whorl, tliameter 1.00 mm (F"igure26). Teleoconch
of 5 whorls, spire high, spire angle of 63-66°. Stnictnre
of strong lamellate axial ribs reflected adapertnralK' and
much weaker uneven spiral cords. Some axial ribs are
stronger than others ami form stronger \arices. Axial ribs
numlier about L3 on final whorl and 14 on penultimate

,uid .iiilepennltiinate whorls. I^ibs regnl.iiK cou\cx on
first two whorls hecoiniiig distiiictk angular at shoiikl(.M-.
Spiral sciil|)tme ol about S jirimaiA cords on spire whorl
ami about 15 on bod\ whorl, each with 5-8 secondaiy
cords in intenals. Spiral cords cross the man\- rib-like
growth lines loruiing finek reticulate reticulate micro-
sculpture (Figure 271. Suture well impressed. Bodv
whoH slightly constricted behind si])honal lasciole. \p-
ertnre with about 17 liiae within that do not extend to
the edge ol the outer lip hut extend deepK within. Inner
lip lorming parietal shield extending slightK' over the
narrow nmbilicns. Colnmella onl\- sliglitK' concave with
three folds, tlii' anterior two close together, much like
one large bibncate fold. Siphonal canal short but dis-
tinct. Shell white, mottled with lii^ht brown bK)tches
near the peiiphciA". Priniar\ cords brown, especiallv
wliere the\' cross the axial ribs on the last 1.5 whorls.
Some specimens are completeK white.

Etymology: The Latin adjective '^(iiiirilns i-a. -iiiii).
angnl.iled. in relereuce to the shell outline.

Remarks: Sippoitdphrni <^i)uiala diliers from .V. tcr-
(iiiuichii (Habe, 1961 1 In its proportionalK higher spire,
narrower nmbilicns, and its bnmn maculations. From .V.
parddlis it differs in having thicker ribs that are not im-
bricate. It also possesses a narrow umbilicus and has a
smaller adult si«'. The two species, .V. pardalis and .V.
goniata. coexist south of Ni'w Caledonia but have never
been taken together

LITER./VTURE CITED

.•\(laiiis. II. and .\. \(laiiis. I So 4 |in 1S53-5S]. The genera of
Recent .Molliisca; arraiii;e(l uccoiiling to their organiza-
tion. 2 vols. John \aii \boist. London. [1: f-256, pis. 1-
32, 1853; 1: 257-4S4, 2: 1-92, pis. 33-72, 1854; 2; 9:3-
284, pis, 73-96, 1855; 2; 285-112. pis, 97-112, 1856; 2;
41.3-540, pis. 113-128. 1857; 2: 541-660. pis. 129-138,
1858.]

15eii. .\. (;. and 1' \ Maxwell. 1987. .\ revision ol llie lossil
and living gastropods related to I'Icsiolrilon Fischer, 1884
(Familv (.'aiicellariidac, Snhlaniilv I'lesiotiitoninae n. sii-
bfani.) with an .Appendix: (ieneni ol Bnccinidae I'isaniinae
related to ('.nluhrnriri Schiiniacher, 1817. New Zealand
Geologiciil Siinev l';ileontoloijit;il Bnllctin 54: 1-140, pis.
1-30.

Bouchet. P. (ed . 1995. Hesultals des Campagnes .ML'SOR-
STOM. volume 14, Meiiioires dii Mns(''nni National
d'llistoire Naturelle 167: 1-654.

Bonchet. P and B. M;nshall ledsl 2001. Tropic;il Oeep-Sea
Benthos, volnnie 22, Menioires dii Mnseinii National
dllistoire N;itnrelle 185; 1-106.

Brocchi. (.'•. 13. 1814. Conchiologia lossile snhapennina con os-
senazioni scolopclie snt;li .Apennini e sni siiolo adiacenle.
Stamperia Heale. Milano. 2 vols. |l;l-240. 2:241-712. pis.
1-16.]

Crosuier A. and V. Boucliet (eds). 1991. Resnitats des Cam-
pagnes MIISORSTOM. volnmc 7, Meinoires du Mnseum
National d'llistoire Naturelle, ser A 150: 1-259.

Ginelin. |. F. 1791. Ciiroli a Linne Systenia Naturae per regiia
tria naturae. Kditio deciiiKi lertia. \'ol. 1, pt. 6 (\enncs):
3021-3910. Lipsiae.

Paiie 104

TIIK XAl'TILUS. \ol. IKS. No.

Hahe. T 1961. Coloured illii.stratiims of the sliell.s oCJapan (II).
lioiku.slui, Osaka. 1S2 pp., .Vppeiuli.x 42 pp., 66 pis. |.'\ii
luiglisli rdiliiin was issueil in 1964. with a lai<;cr pagi' si/.c;
.Shells ol the western I'aeifie in color, \'ol. II. Hoiknsha,
Osaka. 233 pp.. 66 pis. |

Haras(>wych, M. (;. and R. E. Petit, 1982. Notes on the nior-
pholoi^ ol Cduccllami reticulata (Gastropoda: C'aneellar-
iidaet". The Nantilus 96: 104-113.

Ha.segawa. K. 2()()(). Caneellaiioidea, pp. 5S1-5S,5. pis. 2SV)-
291. Ill: T. Okntani (ed.). Marine Mollusks in |apan. Tokai
Uni\ersit\ Press, Tok\Q, 1173 pp.

Houart, R. 2001. Ingensia gen. nov. and eleven new speeies ol
.Mnricidae (Gastropoda) from New Galedonia. X'annatn.
and Wallis and Fntnna Islands. In: P. Bonehet and B. Mar-
shall (edsl. Tropical Deep-Sea Benthos, \olnnie 22. .\le-
nioires dn Mnsenni National d'llistoire Natnrelle. Paris
1S5: 243-269.

Jousseannie, F. 1887, La hmnlle ili's Caiicellanulae i.Moll-
usques Gasteropodes). Le Natuniliste 9 [ = ser. 2, 1]:155-
157, 192-194, 213-214. 221-223.

.Marshall. B. 1991. Mollusca Gastropoda: Segnen/.iidae from
.New (Caledonia and the Lovalt\' Islands. In: A. Grosnier
and P. Bonehet (edsi. Resultats des Gampagnes MUSOH-
.STO.M, Nolnme 7. Memoires dn Musenm National
dTiistoire Natnrelle, Paris, ser. A 150: 41-109.

Petit. R. E. 1976. Notes on Gancellariidae (Mollusca: Gastro-
poda) - III. Tnlane Stndi(>s in Geolog\' and Paleontolog\'
12: 3.3-43. pis. 1-2.

Petit, R. E. and .M. G. llarasewvch. 2000. Three new species
ol the genus Mciica (Neogastropoda: Gancellariidae) troni
Soutli Alrica and llie Philippines. The Nautihis 1 14: 142-
148.

Petuch, E. 1979. TweKc new Indo-Pacific gastropods. Neni-
ouria23: 1-21.

Richer de Forges, B. 1990. Explorations loi liath\al lanna in
the New (Caledonian economic zone. In: .\. ( j'osnier led.).
Resultats des Gampagnes MUSORSTOM. \()lnnie 6. .Me-
moires iln Mnsenm National (.rilistoire Natnrelle. ser .\
145: 9-54.

Richer de Forges. B. 1991. Les tonds meuhles des lagons de
Nou\e!le-Galedonie: genenilites et eehantillonnages par
dragages. Pp. 7-148, in: B. Richer de Forges (ed.). Le
benthos des fonds meuliles des lagons de Non\elle-Gale-
donie. volume 1. Etudes et Theses [ORSTO.M. Paris].

Riclier de Forges, B. 1993. Champagnes d'exploration de la fau-
ne batlnale taites depuis mai 1989 dans la zone t^cono-
mique de la Nouvelle-Galedonie. Listes des stations. In:
A. Grosnier (ed.), Resultats des Gampagnes MUSOR-
STOM. \oKmie 10. Memoires dn .Museum National
<riIistoire Naturelle 156: 27-32.

Richer de Forges. B. and G. Ghe\illon. 1996. Les campagnes
d'echantillonnages dn benthos bath\al en Nomelle-Gale-
donie. en 1993 et 1994 (B.ATHUS la 4. SMIB 8 et HAL-
IPRO 1). In: A. Grosnier (ed.). Resultats des Gampagnes
MUSORSTOM, volume 15. Memoires dn Museum Na-
tional d'Histoire Naturelle 168: 33-53.

Scarabino. \'. 1995. Scaphopoda of the tropical Pacific and In-
dian Oceans, with description of 3 new genera and 42 new
species. In: P. Bouchet (ed.), Resultats des Gampagnes
MUSORSTOM, volume 14. Memoires dn Museum Na-
tional d'Histoire Naturelle 167: 189-379.

X'erhecken, .A. 1997. Mollusca Gastropoda: .\ralm'a Sea Gan-
cellariidae collected durint; the K.\Rl'B,\R cniise. Me-
moires du Museum National d'Histoire N;itnrclle 172:
295-323.

THE NAUTILUS 116(3):105-1()8. 200:2

Paijc 105

Spawn of the Patagonian gastropod Parcuthnii pluinhca
(Philippi, 1844) (Buccinidae)

Guiclo Pastorino

Miisro Ar<i;entino tie Ciencias

Xatiirales-CONICET
A\. Ancrel Ckillanlo 470
C1405 DJR Buenos Aires
ARGENTINA
r\pastor@cril)a.ed\i.ar

Pablo E. Penchaszadeh

Miiseo Anj;entiiu) de C^ieneias
Naturales-C'ONICET

Aw Angel CJallardo 470

CI405'dJR Buenos Aires

ARGENTINA

and

Faeulfad t\f Ciencias Exactas v
Naturales

Universidad de Buenos Aires

Buenos Aires

ARGENTINA

ABSTRACT

The spawn ot Piirctithrid plmnl)cii (Philippi. 1844) consists of
an egg mass of 4-19 egg capsules. The uncleaved egg, in num-
ber of 1-5 per capsule, measured about 300 |a.m. Pdniithrui
phimbca has direct development without niu'se eggs, hatclnng
at craw ling stage. Extra-vitelline substances in the intracapsular
lifjuid are likeK' to be the onK e.\tra\itelline tootl available for
the embrvos. Hatching occurs b\' nipture or dissolution ol the
capsule wall in a possibly random location (ju the (liin basal
membrane. The number of hatchlings varied troui 1-5 within
the same spawn. Hatchling muuber and shell m/.c are related.

INTHOnUCTION

The genus VdrriilJirid was proposcil hv Strcliel (1905)
in hi.s extensive accoinit ol tlie gastropod launa Ironi the
Strait of Magellan. He descrihed six new spe-cies under
I'lirciilliriii. Hovvevvr, the validitv ol these species re-
iriains uncertain.

Pairuthrid plnmheti (Philippi. IS44) is the oldest
name of a \en' poKnioiiihic species endemic to the .Ma-
gellanic province (Figures 1-3). It ranges from soiitlu'rn
C.'huhnt province {~45°S) in Argentina to 49°S in Chile
(Dell, 1971). It is usually found inti-rtidallv at lovy tide
inider rocks and in tide pools. Ciallardo and Penelias/.a-
deli (2001) mentioned Parciitliha jihniihiii as liaving di-
rect or non-pelagic development.

Bnceinids have a wide range of deyelopnicnlal iiiddes.
from tree-.svvimming planktotrophic veligt'rs (Bandel,
1975, 1976: Amio, 1963) to lack of a lanal stage { Fio-
roni, 1966: Mattel et al., 19Sfi: Pdrlniaun, 1925; Milos-
lavieh and Dnfresne, 1994; Miloskivicii aud Penchasza-
deh. 1994).

In this note, we describe the spawn ol Ptirculliria
pluuthi'd (Philippi, 1S44), including the ninnber and size

of the uncleaved egg. the development pattern cUid die
hatching stage, of a population from Puerto Deseado,
Argentina (—46° S) on the .\tlantic coiLst.

M.\TERIALS AND MKTll()i:)S

Adidts specimens and egg masses ol Parcutliria phuuhca
were collectetl at low tide in Dos Hermanas. Puerto De-
seado. Santa ilrnv. Province, .Xrgentina (47°45'S-
65°55'\^') during Fehrnarv 2000. The intertidal area i.s
occupied liv mvtilids toiuiing ininks on n)ck"v platforms.
Specimens and spawns ol /' phiDihca were freijuentlv
found in crevices and under rocks. Adults specimens and
egg capsules were fixed in o7( formalin on sea water.
Those with shelled embryos were presened in 70% etli-
anol. Most photographs of the eggs, capsules and em-
bryos were taken using a digital scanning camera at-
tached to a Zeiss Stemi 2000 stereoscopic microscope.
All iiuages were digitally processed.

RESL ITS

We examineil 15 egg masses, lor a tol.il number ol 106
egg capsules. Each egg mass is composed ol 4-19 egg
capsules (X = S.S3, SD = 5.07. n = 15) (Figure 6).
Pareiilhriti ))liiiiil>cri has transparent, yellowish, subcir-
cnlar, bnlliform capsules, which are often aggregated and
partialK ovcilapping t'ai'h other (I'ignres 7. 10), They are
attached to the substrate bv a circular ba,sal membrane
measuring 4.2-5.2 mm (X = 4.96, SD = 0.22, n = 10),
P'.ach capsule has its own basal membrane that is not
fused with adjacent ones. The ca|)snles nieasiu'e 3.2-3.S
nun in dianu-ter (X = 3.6, SD = O.IS, n = 10), There
is a suture line that splits the ca|)snle in two sides (Fig-
ures 9, II). Tliere is no pre-formed apical plate or es-
cape aperture as a plug, nor an escape slit. During iiatch-

Page 106

THK NAUTILUS. \n

Ifi. \o. 3

Figures 1-9. I'dirnlhria phinibi<i i rliilippi, lS44i. 1-3. MACN IO.'5i2. adult spceiiiicii IVoiii Caleta (.'oylc, S.iiita ( ju/, Ari^riiliiia.
Scale l)ar = I ( uj. 4. Detail of tlic protoconch. Siali- liar = 1 iniii. 5. Tiirce ciiibnos at tlio pre-liatciiliiin staijf. Scale liar = 1
mm. 6. Freslily Ian! spawn with \isilile cluster iif ei^i;s. Scale liar = .') iiiiii. 7. Several clusters of spawns, the tup ca]isnles with
three and two einhnns respcctixcK. 8. 'I'hree pws IroMi llie s.uiie capsule. Seale liar ^ oOH |Jini 9. Isolaled capsule with five earl\.
shelled enibnos. Scale liar = 1 nun.

G. Pastoriiio and P. P]. Pciifliaszadcli. 2002

Paw 107

""N^

^■^»!f^.■.i .' ..

.l^^-;"';.%. I

|7jfc-- ■ '-iM: .-^m %:■■ ;.•.#**?-;-.-. Vi;

■v--l.

^■.^■'7»»S^

10

..<*r^ " /■"

mt'-^.

.^'

Figures 10-12. Egg capsules of rtirciilliiin plmnhcd ■ I'liilippi. 1S441. 10. (.luster ( if i •<;<.; rapsulcs. II. Kulatcil capMilc, 12.
Middle section showing die capsule pnifilc. Scale Imis = 1 nini.

ing the rupture of the liasal iiu'iiiliianc tlial is imikIi
thinner than the upper wall (li the capsule was iccmded:
this could he an escape w"a\ for hatchlings.

The uncleaved eggs nieasiu'ed 280-340 |xni in diam-
eter (X = 300, SD = 15.6S, n = 14) (P^ignre S). They
are embedded in a \en dense intiacapsiilar licjnid that
becomes, when presened in ethxl alcoliol, opa(jue and
flocculates like denatured egg whites. This intra capsular
licjuid is conipleteK- translucent at the latest stages of
de\elopnient, suggesting that it contains nutritional sub-
stances that would be ingested In tlie cmbnos. \o
nurse egjis were recorded.

The number of embnos was 1-5 within an\ given
spawn. When fixe are present, one is generalK abortive
and is proliablv eaten iiv the remaining embnos. Hatch-
ling number and shell size are apparentlv iclatrd. When
a single embryo developed, the shell length at tin- [)ie-
hatching stage was: 2.14—2.19 mm (X = 2.1H nnn, n =
4, SD = ().()3), with t\vo embrvos the shell length was:
1.79-2.22 (X = 1.99 mm, n = 13, SD = 0.14), with
three; 1.60-1.97 (X = LSI mm. n = 21, SD = 0.09),
with four: 0.79-1.S4 mm (X = 1.62 nun, n = .33, SD =
0.17'. We did not find capsules at the late pre-hatching
stage with five embnos. The embnos hatch as crawling
juveniles, with a well-developed loot (Figure .5).

The protoconch has 'IVi-IVa convex, somewhat glo-
bose, wliorls (Figure 4). Whorls are completelv smooth
and browmish in color There is a gradual transition to
teleoconch. Therefore, changes in the axial firnamenta-
tion are nsualK' the onlv wav to recognize features of the
adult shell. The first teleoconch wiiorl has weak and ir-
rcg\ilar costae tliat disappear graduallv.

DISCUSSION

Development with planktonic lanae has been described
for several species within the familv Buccinidae. gener-

alK from ti'opical regions (e.g.. Visdnia ptisio (Linnaeus.
175S) and Pitsia fiiicfa (Onnad, 1S4B) (formerlv in the
genus Caiithani.s): Bandel, 1975, 1976). In temperate
and colder waters thi're are also examples of hatching of
veliger lanae (e.g., Bdhtjlonui japoiiicti (Reeve, 1S42);
Amio, 1963; Hivest, 19S3>. However the most common
mode of development within the Buccinidae involves the
lack of free-swimming lanal stag(>s, hatching of crawling
juveniles, and the presence nl muse eggs. The best-
kniown examples are Biicciiniin uiidfitniii Linnaeus. 175S
(Fioroni, 1966; Martcl ct al., i9S(-;: Portmann, 1925), B.
n/aucuui Bniguiere, 1792 ( Miloslavich and Dufresne,
1994), Scarlcsia (lira (Reeve. 1S46) (Rivest. 19S3i, and
Cuius stiiui)s()ui (M()rch. 1S67' iWcst. 1979i. anionic oth-
ers.

Other spei'ies of iinecinidac. such ;is F,li<J,iiiti()pliiis ilii-
iciitctiis (Sav, !S25i. Ii;ivc dirccl development vvitliout
nurse eggs, h;ililiin',; at tlie crawling pe(hveliger stage
(MiIosla\ich and Piiicliaszadcli. 1994). In this latter .spe-
cies, albumin in the intiacapsuhu' Ii(|uid is the most im-
portant extra food ;ivaihiblc foi' the embnos (Miloslav-
ich, 1999). (iiveii the hick of nurse eggs aTid tlu' large
difference between the nncleavcd egg diameter and the
shell length at hatching, I'firciilliria plumbed ])robablv
shows the same pattern of embnonic Icedinti on sub-
stances of the intracapsular llnid.

We found several capsules with onlv one cnibno in
an earlv stage of devi-lopincnt, and the rest ot them at
a later stage. The absence of empty shells inside the
capsule makes nnlikelv the possibilitv of cannibalism ;it
a late developmental stage. The same was ob.sencd in
Euiiouiophos uuiciurlus (Miloslavich and Penchaszadeh,
1994).

The lact tliat clusters loinied bv several c'gg masses
of /' pluinhcd are connnon could indicate a gregaiious

Page 108

THE NAUTllAiS. \ ol. lift, NO. 3

spawninif lH'lia\ior as demonstiatccl lor sexeial other
iieogastropotis (D'Asaro, 1970) and partieiilarlv for some
hiRX'inids siieli as E. iiiiiciiuiiis ( Milnslax icli and Pen-
ciuiszadeh, 1994).

The egg capsules ol some hnceiiiids present a pre-
formed exit liole or plug. This is the ease, among others,
u\ liucciniun tindattiDt. B. lu/dropliiiniiin Hancock, lS4fi
(Thorson, 1935) and all the species of the genus Ci>lii.s
(studied by Thorson, 1935 and 1946 as Sipho), including
C.jejfretjsiaruis (Fischer, 1868) ( = C. hoivsei in Lebour,
1937). In other buceinids the escape of hatchliugs takes
place through a widi' slit that is situated in difierent po-
sitions according to the species (e.g., Neptiinea antkjua
(Linnaeus, 1758), Beringiiis tiirtoni (Bean, 1834)), or by
an aperture tolkmang the suture line of the egg capsule,
as in E. uuiriiictiis (Milosla\'ich and Penchaszadeh,
1994). Despite the presence oi a sntiu'e line in the egg
capsules of F pl\nnhc(t. the opening iloes not occur along
tliis line, but b\' rupture or dissolution of the capsule
wall in an apparentK random location on the thin basal
membrane.

The large egg capsules oi \\>lutiij)sius muxciiicHs
(Gmelin, 1791) and Pijntlofusiis dcfoniiis (Ree\e, 1847)
do not liave anv kind of exit hole (Thorson. 1940). This
is also the ease in Parcuthiia pluinljca, although its egg
capsules are clearK smaller.

.ACKXOWLKDCMFA'TS

Special thanks are tine to .\. Pettoxclio and A. Dolcc-
maseolo who pro\ided hospitalitx at Puerto Deseado.
Veronica hanov kindly drew Figiu'es 10-12. Two anon-
ymous reviewers improved considerabK the original
manuscript. The fieldwork was supported in part b\' Pro-
ject PICT No. 01-04321 from the National Agency lor
Scientific and Technological Promotion, Argentina.

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